New Endemic Species of Capsicum (Solanaceae) from the Brazilian Caatinga: Comparison with the Re-Circumscribed C. parvifolium

Abstract

Two new species of Capsicum from the Caatinga Biome (Brazil) are described and illustrated. The two species are endemic to the north-eastern states of Brazil (C. caatingae: Bahia, Pernambuco, and north of Minas Gerais; C. longidentatum: center of Bahia and Pernambuco) and are morphologically similar to another caatinga species, C. parvifolium. Lectotypification and a complete description for C. parvifolium, which has been confused in the literature and herbaria, are also provided. The karyotype for the three species (2n = 2x = 24) is analyzed and discussed, and a key to differentiate the endemic Capsicum species from Brazil is included.

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Copyright (c) American Society for Plant Taxonomists. All rights reserved.
Systematic Botany (2011), 36(3): pp. 768–781
© Copyright 2011 by the American Society of Plant Taxonomists
DOI 10.1600/036364411X583718
768
Capsicum L. (Solanaceae) is an American genus of ca. 30 spe-
cies ( Moscone et al. 2007 ). It can be easily recognized by the
unlobed calyx that usually bears appendices or teeth, the con-
spicuous expansion of the filaments where they unite with the
corolla tube (stapet), which is functionally related to nectar
presentation, and berries that are usually pungent and with
giant cells in the innermost layer of the mesocarp ( Hunziker
2001 ). The genus is economically important for its five domes-
ticated species ( Bosland and Votava 2000 ; Reifschneider 2000 ;
Buckenhüskes 2003 ; Thampi 2003 ; Ravishankar et al. 2003 ;
Barceloux 2008 ; Yamamoto and Nawata 2009 ). According to
Pickersgill (1969) domestication probably began about 8,000
B. P. in Mexico for C. annuum , but starch fossils found in two
sites of southwestern Ecuador suggest that these plants must
have been domesticated elsewhere earlier than 7,000 B. P.
and brought into Ecuadorian sites from either the north or
the south ( Perry et al. 2007 ). After introduction of peppers to
Spain (mainly the members of the C. annuum complex) they
spread widely all over the world ( Basu and De 2003 ).
In the traditional classification of the family, Capsicum is
placed in tribe Solaneae, subtribe Capsicinae Yamazaki (sub-
fam. Solanoideae) with eight other genera ( Hunziker 2001 ),
but its phylogenetic placement based on different molecular
analyses, is in clade Capsiceae together with Lycianthes Hassl.,
its closest relative ( Olmstead et al. 2008 ). A classification of
the component species of Capsicum is still being studied. In
recent years, different classical and molecular cytogenetic
analyses ( Alcorcés de Guerra 2001 ; Moscone 1990 , 1993 , 1999 ;
Moscone et al. 1993 , 1995 , 1996 , 2003 , 2007 ; Park et al. 1999 ,
2000 ; Pozzobon et al. 2006 ; Scaldaferro et al. 2006 ), crossing
experiments ( Pickersgill 1991 ; Tong and Bosland 1999 ; Onus
and Pickersgill 2004 ), enzymatic studies ( McLeod et al. 1979a ,
b , 1982 , 1983 ; Loaiza-Figueroa et al. 1989 ; Quintero Barrera
et al. 2005 ), and plastid and nuclear DNA sequence studies
( Prince et al. 1995 ; Paran et al. 1998 ; Rodríguez et al. 1999 ;
Walsh and Hoot 2001 ; Buso et al. 2002 ; Kochieva and Ryzhova
2003 ; Votava et al. 2005 ; Guzmán et al. 2005 ; Cvikic et al. 2009 ;
Ince et al. 2009 ; Aguilar-Meléndez et al. 2009 ; Bahrami Rad
et al. 2009 ; Stágel et al. 2009 ) have facilitated progress in the
characterization and establishment of taxonomic groupings
in Capsicum . However, no formal infrageneric classification
has yet been proposed and the exact number of species is still
uncertain.
Species of Capsicum are native throughout Central and
South America but Brazil is the country with the greatest
concentration of species ( Barboza and Bianchetti 2005 ), espe-
cially in its southeastern coastal area (Espírito Santo to Rio
Grande do Sul). Only two taxa, C. annuum var. glabriuscu-
lum (Dunal) Heiser & Pickersgill and C. parvifolium Sendtn.
( Hunziker 2001 ; Barboza and Bianchetti 2005 ), have been
recorded in northern and north-eastern Brazil, an area with
a flora rich in endemism and diversity ( Cabrera and Willink
1980 ; Lleras 1997 ; Daly and Mitchell 2000 ; Giulietti et al. 2002 ,
2006 ; Queiroz 2006 ).
Northeastern Brazil is mostly occupied by the Caatinga
biogeographical Province, which constitutes one of three arid
nuclei in South America, together with the Guajira Peninsula
along the Caribbean coast of Colombia and Venezuela, and
the dry belt that extends from Argentina through Chile and
Peru into Ecuador ( Ab’Saber 1980 ; Andrade-Lima 1981 ,
1982 ). The Caatinga Biome covers an extensive area from the
north Atlantic coast of Ceará and Rio Grande do Norte states
to southern Bahia and part of Piauí and Mina Gerais states
( Ab’Saber 1977 ; Cabrera and Willink 1980 ). In this region,
the climate is semiarid, consistently hot, with summer and/
or autumn rains ( Daly and Mitchell 2000 ; Cavalcante and
Major 2006 ); it is covered by a xerophytic, spiny and decidu-
ous vegetation, unique in the world ( Cavalcante and Major
2006 ). General characteristics of plants of the Caatinga Biome
include small leaves with a waxy cuticle that are lost during
the dry season, dense branching of trees from the base (giving
them a shrubby appearance), and the presence of succulent
and crassulaceous species ( Lleras 1997 ).
To date, only C. parvifolium has been recorded from the
Caatinga ( Barboza and Bianchetti 2005 ). In the last 30 yr,
botanical exploration through this region has greatly inten-
sified by Brazilian researchers who have made many new
collections and reidentified existing collections of Capsicum .
New Endemic Species of Capsicum (Solanaceae) from the Brazilian Caatinga:
Comparison with the Re-circumscribed C. parvifolium
Gloria E. Barboza , 1
,
4 María F. Agra , 2 María V. Romero , 3 Marisel A. Scaldaferro , 3 and Eduardo A. Moscone 3
1 Instituto Multidisciplinario de Biología Vegetal (IMBIV-CONICET) and Facultad de Ciencias Químicas,
Universidad Nacional de Córdoba. CC 495, CP 5000. Córdoba, Argentina.
2 Laboratório de Tecnologia Farmacêutica Prof. Delby Fernandes de Medeiros, Universidade Federal da Paraíba,
Caixa Postal 5009, 58015-970, João Pessoa, PB, Brasil.
3 Instituto Multidisciplinario de Biología Vegetal (IMBIV-CONICET) and Facultad de Ciencias Exactas, Físicas y Naturales,
Universidad Nacional de Córdoba. CC 495, CP 5000. Córdoba, Argentina.
4 Author for correspondence: ( gbarboza@imbiv.unc.edu.ar )
Communicating Editor: Anne Bruneau
Abstract— Two new species of Capsicum from the Caatinga Biome (Brazil) are described and illustrated. The two species are endemic to the
north-eastern states of Brazil ( C . caatingae : Bahia, Pernambuco, and north of Minas Gerais; C . longidentatum : center of Bahia and Pernambuco)
and are morphologically similar to another caatinga species, C . parvifolium . Lectotypification and a complete description for C . parvifolium ,
which has been confused in the literature and herbaria, are also provided. The karyotype for the three species (2 n = 2 x = 24) is analyzed and
discussed, and a key to differentiate the endemic Capsicum species from Brazil is included.
Keywords— Caatinga Biome , endemism , karyotypes , lectotypification , morphology , peppers.

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The analysis of these herbarium collections and observations
in the field revealed the need for the description of new taxa
and the recircumscription of C. parvifolium . Here we present
a detailed description of C. parvifolium , which has been con-
fused in the literature, and karyotype diagrams for C. longi-
dentatum and C. parvifolium .
Materials and Methods
Taxonomy— Descriptions were based on measurements of living
plants taken during field work in the Caatinga of Bahia and Paraíba and
examination of herbarium specimens loaned from or inspected at the 28
herbaria listed in the Acknowledgments.
Field observations and morphological examinations of material pre-
served in FAA solution and dried material using a stereomicroscope were
conducted. Measurements on dried material were made from dissections
of flowers rehydrated in hot water. Information about flower, fruit, and
seed color was taken mainly from our own observations in the field; we
also tested pungency of the fruits by tasting them in the field. Vernacular
names were taken from herbarium specimen labels.
The map with gray shading indicating the Caatinga region was
adapted from Lleras (1997) . The geographic distribution for each species
is based on all herbarium collections analyzed. The conservation status
proposed here follows the IUCN ( IUCN 2010 ).
For C. parvifolium and its synonyms, all the type specimens found were
studied in the herbaria where they are housed with the exception of one
Fregirardia leptoclada fragment, which is held at F and was analyzed from
the digital image. The specimens selected as lectotypes correspond to the
best preserved and most diagnostic material.
Karyology— One population each of C. longidentatum ( Agra and Barboza
7086 ) and C. parvifolium ( Agra and Barboza 7075 ) were studied. Somatic
chromosomes were observed in squashed root meristems obtained from
germinated seeds. The root apices were fixed in 1:3 acetic acid:ethanol
mixture for 12 hr after a pretreatment in p-dichlorobenzene-saturated
solution for two hr at room temperature.
The material was kept at -20°C until examination. In all cases root tips
were then stained according to Feulgen’s technique using Schiff’s reagent
for one hr in darkness, after hydrolysis in 5 N hydrochloric acid for
50 min at room temperature ( Jong 1997 ). Meristem cells were isolated,
macerated, and squashed in a drop of 2% aceto-carmine after the staining.
Slides were made permanent by freezing with liquid CO
2 ( Bowen 1956 ),
removal of the coverslip, and mounting in Euparal. Chromosomes were
observed and photographed with a Leica DMLB microscope equipped
with a Leica DC 250 digital camera and the Leica IM 1000 image manage-
ment system. Somatic chromosomes measurements (l = long arm length,
s = short arm length and c = chromosome length) were taken from six meta-
phase plates. The arm ratio (r = l/s) was used to classify the chromosomes
as recognized by Levan et al. (1964) as metacentric or “m” (r = 1.00–1.69),
and submetacentric or “sm” (r = 1.70–2.99). Battaglia’s (1955) terminology
for satellites was used as follows: microsatellite, having a shorter diameter
than the chromosome diameter and being small in size; macrosatellite,
having a diameter equal to the chromosome diameter and being large in
size. The satellite lengths were added to the lengths of the corresponding
arms. Idiograms were prepared by organizing the chromosomes into two
groups according to their increasing arm ratio (metacentric to submeta-
centric), and then according to decreasing length within each group.
Taxonomic Treatment
Capsicum caatingae Barboza & Agra, sp. nov. — TYPE:
BRAZIL. Bahia. Munic. Cachoeira: Estação da Embasa,
Cachoeira/Bahia, Vale dos Rios Paraguaçu e Jacuipe,
39°05’W, 12°32’S, 40–120 m, Jun 1980, P. do Cavalo et al.
162 (holotype: HUEFS 00920!; isotype: ALCB 07938!).
A Capsicum parvifolium Sendtn. inflorescentiis multifloribus
(floribus 5–18), calyce edentato, seminibus flavidis differt.
Shrub or small trees 2–3.5 m tall; stems few branched, grey,
slender, scandent, striate; old stems glabrous; young stems gla-
brescent with antrorse simple hairs, and sometimes with tiny
glandular hairs. Leaves mostly solitary, rarely geminate, 2–2.5
times longer than broad, 3–4.9(–6) cm long, 1.3–2(–2.4) cm
wide, ovate to elliptic, membranous or papery, the adaxial
surface dark green, the abaxial surface light green, glabres-
cent to pubescent on both sides, with simple hairs and small
glandular hairs especially on the lower surface veins, the
margin entire, the apex scarcely acuminate, the base shortly
attenuate and unequal; petioles (0.5–)0.7–2 cm long, pubes-
cent. Flowers in fascicles of 5–13(–18); flowering pedicels
pendant, not geniculate (straight) at anthesis, 0.7–2.1(–2.8) cm
long, glabrescent or pubescent, the hairs simple and antrorse.
Flower buds ellipsoid, white or greenish white. Calyx green,
cup-shaped, toothless, with short uniseriate verrucose nong-
landular trichomes, 1.2–1.7(–2) mm long. Corolla stellate,
4.5–6(–8) mm long, lobed nearly halfway to the base, with
bright purple lobes within except for narrow white marginal
band, and greenish white center, pubescent adaxially with
small glandular hairs with a unicellular head and a bicellular
stalk at the base of the lobes and in the throat; lobes longer
than the tube, ca. 2.9–3.5 mm long, 1.7–2.4 mm wide, broadly
triangular, the tip cucullate, pubescent with bicellular simple
hairs on the tips and involute margins. Filaments greenish
white, (0.8–)1.1–1.75 mm long; anthers yellowish cream, ca.
1.4–2.1 mm long; stapet ca. 2 mm long. Ovary pale green, sub-
globose, ca. 1.1–1.4 diam; style pale green, (4.3–)4.6–4.8 mm,
widening distally; stigma greenish yellow, somewhat bilobed.
Berry globose, slightly flattened at the apex, ca. 7–11 mm diam,
green when immature, yellow at maturity; fruiting pedicels
pendant, the fruiting calyx persistent, discoid with the margin
entire or sometimes easily torn, the pericarp hot to the taste
and lacking stone cells. Seeds (9–)11–17 per fruit, pale yellow,
3.2–3.7 mm long, 2.2–2.8 mm wide; testa thick, foveolate, with
spine-like projections, the fovea deep at the seed margin and
shallow in the central zone. Figures 1 , 2 , 3G .
Vernacular Names— “Pimenta brava” (Paraíba), “pimenta”
(Paraíba).
Etymology— The new species is named in allusion to its
habitat since it is distributed through the Caatinga Biome in
Brazil.
Phenology— Flowering from December to June; fruiting
from February to June. Some specimens have been collected
in flower in September.
Distribution and Habitat— Endemic to the northeastern
states of Brazil (Bahia, Pernambuco and the northern part of
Minas Gerais, Fig. 2 ) between 100–775 m in the arborescent-
shrubby open Caatinga with shrubs ( Jatropha mollissima (Pohl)
Baill., Varronia leucocephala (Moric.) J. S. Mill. and many spe-
cies of Croton L.), cacti, bromeliads, and some trees ( Spondias
tuberosa Arruda, Ziziphus joazeiro Mart., and species of Mimosa
L., among others). It is usually found in the margins of
Caatinga forest and amongst granitic and gneissic outcrops
(‘inselbergs’).
Conservation Status— Widespread disturbance (agricul-
tural expansion, grazing, hunting, and burning) is observed
in the caatinga, and therefore the geographical and ecologi-
cal area of distribution of this species is relatively affected.
Following the IUCN ( IUCN 2010 ), we suggest that this spe-
cies be assigned a conservation status of vulnerable (VU).
Observations— The pubescence and leaf shape of C. caatin-
gae are variable. In general, plants are glabrescent with simple,
multicellular (3–5-celled), and verrucose nonglandular hairs
on young stems, leaves, pedicels, and calyx ( Fig. 1 H ), and
tiny glandular hairs (multicellular or unicellular head, short
2–3-celled stalk) on leaf veins and calyx ( Fig. 1E, J, N ). Some
specimens ( Guedes et al. 13245, 13297 ) are pubescent with

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770 SYSTEMATIC BOTANY [Volume 36
Fig . 1. Capsicum caatingae ( Hunziker 25233 ). A. Branched trichome from leaf. B. Open corolla sector, internal view. C. Flower. D. Gynoecium. E, J, N.
Glandular trichomes from calyx and leaf venation. F. Apex of a flowering branch. G. Node of a fruiting branch. H. Verrucose nonglandular trichomes from
calyx. I. Embryo. K. Simple nonglandular trichome from the corolla lobes. L. Fruit in cross section. M. Calyx. N. Ovary in cross section. O. Seed in cross
section. P. Seed in lateral view. Adapted from Hunziker (2001) .

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2011] BARBOZA ET AL.: THE CAPSICUM SPECIES FROM THE BRAZILIAN CAATINGA 771
longer trichomes (up to seven cells) and occasionally also
have branched trichomes ( Fig. 1A ). Leaves are small to
medium-sized (2–6 cm long, 1.3–3 cm wide) but they can mea-
sure up to 9 cm long and 4 cm wide in some specimens ( Melo
4336; Bautista and Pinto 1023; Silva 18; Miranda 1642, Sobral
7646 ).
Capsicum caatingae is distinguished from other species of
Capsicum by the combination of number of flowers, non-
geniculate (straight) pedicel, nontoothed calyx, and flower
color. The presence of fascicles up to 18-flowered is an exclu-
sive trait of this species. In fact, in the majority of Capsicum
species, the inflorescences consist of (1–)2–4 flowers with the
exception of nine species ( C. ceratocalyx M. Nee, C. coccineum
(Rusby) Hunz., C. flexuosum Sendtn., C. hookerianum (Miers)
Kuntze, C. lanceolatum (Greenm.) Morton & Standley, C. parvi-
folium Sendtn., C. rhomboideum (Dunal) Kuntze, C. schottianum
Sendtn., and C. tovarii Eshbaugh, P. G. Sm. & Nickrent) where
inflorescences are 6–9-flowered. Among the four Brazilian
wild species with a toothless and cup-shaped calyx ( Barboza
and Bianchetti 2005 ), C. caatingae differs in corolla color from
C. flexuosum and C. campylopodium Sendtn., which have a
corolla that is white with yellowish-green or golden spots on
the internal surface at the base of the lobes and limb, respec-
tively, while in C. caatingae the corolla has bright purple lobes
within except for a narrow white marginal band, and green-
ish-white center. In the other two species, C. pereirae Barboza &
Bianchetti and C. schottianum , the corolla has purple spots
inside that do not extend through all lobe surfaces as seen in
C. caatingae . The nongeniculate pedicel at anthesis ( Fig. 1F )
is a character shared with other species from Brazil ( Barboza
and Bianchetti 2005 ; see key), and with C. dimorphum (Miers)
Kuntze, C. rhomboideum, and C. hookerianum from the Andean
region (Barboza unpubl. data).
Capsicum caatingae most closely resembles C. parvifolium
in its habit, indumentum, straight pedicels, size, shape, and
color of the corolla, and in its fruit. However, C. caatingae has
a toothless calyx ( Fig. 1M ), a many-flowered inflorescence
( Fig. 1F ), and pale yellow seeds ( Fig. 3G ).
Karyology— This species has 2 n = 2 x = 24. Two differ-
ent cytotypes have been described for C. caatingae (both as
C. parvifolium , Moscone 1993 ; Moscone et al. 2007 ). The first
consists of a karyotype composed entirely of m chromosomes
( Moscone 1993 ) while the second cytotype exhibits 11 m + 1
subtelocentric (st) chromosomes ( Moscone et al. 2007 ). In both
cytotypes, pair no. 12 possesses nucleolar organizing regions
(NORs) and satellites in the short arms.
Additional Material Examined— BRAZIL. Bahia: Cachoeira, Mata
a NE. da B. Bananeiras, Cachoeira/Bahia-Vale dos Rios Paraguaçu e
Jacuipe, 12°32’S, 39°05’W, 40/120 m, Jul 1980, P. do Cavalo 438 & 453 (ALCB,
CEPEC, HUEFS, NY). At the same place, P. do Cavalo 331 & 771, Jul-Sep
1980 (ALCB; HUEFS). Caraíba, Apr 1994, Sobral & Ganev 7646 (HUEFS).
Feira de Santana, parque da cidade, 12°15’S, 39°4’W, 15 May 2007, Moraes
790 (HUEFS). Feira de Santana, Distrito de São José da Itapororoca,
Serra de São José, Faz. Boa Vista, 12°15’S, 38°58’W, 11 May1984, Noblick
3207 (CEPEC, CTES, HUEFS, MBM, NY). Distrito Ipuaçu, face oeste,
12°13′55″S, 39°04′39″W, 240 m, 30 Jun 2005, Couto et al.106 (HUEFS). Monte
Alto, Caatinga de Grotão, 14°13′55″S, 39°4′34″W, 8 Jul 2006, Lima et al. 38
(HUEFS). Ibiquera, Gruta da Lapinha, 12°27′51″S, 40°58′47″W, 610–710 m,
1 May 2004, Franca et al. 4956 (HUEFS). Ipirá, Faz. Nova Fanela, ca. 2.5 km
S de Ipirá, 12°10′45″S, 39°46′12″, 278 m, s. f., Queiroz et al. 10598 (HUEFS,
JPB). Itatim, Morro do Quixaba, 12°44′21″,3″S, 39°47′33″,9″W, 273 m, 3
Apr 2009, Agra et al. 7085 (CORD, JPB). Inselbergue Morro da Queixaba,
12°44′26″S, 39°47′37″W, 290 m, 12 Feb 2006, Moraes et al. 159 & 160 (HUEFS).
Itatim, Pedra Grande, 12°42′59″S, 39°45′41″W, 320 m, 9 Apr 2005, Silva et
al. 18 (HUEFS). Itatim, interior da mata da base do Inselberg, 12°45′12″S,
39°46’W, 300 m, 26 Jan 1997, Melo et al. 1969 (HUEFS). Itiúba, Fazenda
Experim. de Epaba, 10°43’S, 39°50’W, 27 Jun 1983, Pinto & Bautista 104/83
(HRB). Jacobina-Barracão de Cima, 11°01′38″S, 40°33′38″W, 662 m, 6 Jul
1996, Harley et al. 3445 (CEPEC). Santo Estevão, BR 116, 53 km SW de Feira
de Santana, 200 m, 12°30′40″S, 39°22′27″W, 27 May 1987, Queiroz et al. 1520
(CEPEC, HUEFS, NY). Serra Preta, Faz. Santa Clara Nova, 12°09′24″S,
39°22′46″ W, 262 m, 2 Apr 2009, Agra et al. 7081 (JPB). 6 km W do Ponto
de Serra Preta, Faz. Santa Clara, 12°10’S, 39°20″W, 262 m, 17 Jul 1985,
Noblick & Lemos 4227 (HUEFS). Jaguarari, Caminho do Engenho, estr. para
Grotas, 10°08’S, 40°13’W, 24 Jun 2005, Souza-Silva & Rapini 21 (HUEFS,
JPB). Miguel Calmon, Piemonte da Diamantina, entorno do Parque Sete
Passagens, Ponto 230, 11°19′56″S, 40°35′53″W, 525 m, 23 Dec 2006, Guedes
et al. 13245 & 13297 (ALCB). Euclides da Cunha, Sítio do Jaime, 10°30’S,
39°00’W, 20 Mar 2004, Guedes et al. 10847 (ALCB). Reserva de Cotigipe,
22 Sep 1994, Guedes et al. s. n. (ALCB). Caem, Estrada BR 324 Caem, Km
4, 11°09′15″S, 40°24′19″W, 450 m, 18 Feb 2004, Pereira-Silva et al. 8481
(HUEFS). Campo Formoso, 10°30′32″S, 40°25′59″W, 739 m, 14 Apr 2006,
Barreto et al. 218 (HUEFS); Senhor do Bonfim e Mun. Itiuba, plantas de
semillas cultivadas en Viçosa, 13 Apr 1988, Hunziker 25333 (CORD). Minas
Gerais: Monte Azul, Sentido Jaíba, estrada de chão, 15°12′36″S, 42°53′49″W,
775 m, 22 Apr 2006, Carneiro-Torres et al. 717 (HUEFS, KPB). Permambuco:
Buique, Parque Nacional Serra do Catibau, trilha dos Torres, 8°34’33″S,
37°14′45″W, 740 m, 27 Jun 2007, Santos et al. 1574 (HUEFS). Garanhuns,
ca. 58 km Arcoverde, 08°44′02″S, 38°44′05″W, 955 m, 19 Apr 2005, Melo et
al. 3862 (HUEFS, JPB). Inajá, Res. Ecol. Serra Negra, 23 Mar1994, Miranda
et al. s. n. (ALCB). Lagoa dos Gatos, Pau Ferrado, 21 Apr 1994, Miranda &
Félix 1642 (ALCB). Estrada Ibimirim-Petrolândia, 3 Jun 1995, Sales et al.
604 (JPB).
Capsicum longidentatum Agra & Barboza, sp. nov. — TYPE:
BRAZIL. Bahia: Itatim, Morro da Pedra Grande, base do
Fig . 2. Distribution of Capsicum species in the Brazilian Caatinga
(gray area). C. caatingae (circle), C. longidentatum (square), C. parvifolium
(asterisk). Abbreviations: PI, Piauí; CE, Ceará; RGN, Rio Grande do Norte;
PA, Paraíba; PE, Pernambuco; AL, Alagoas; SE, Sergipe; BA, Bahia; MG,
Minas Gerais.

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772 SYSTEMATIC BOTANY [Volume 36
Fig . 3. Capsicum parvifolium (A-E), C. longidentatum (F, H, I), C. caatingae (G). A. Fruiting branch. B-D. Details of the flower (in D, a nongeniculate pen-
dant flower is observed). E-G. Seeds. H. Typical aspect of the plant in the arid caatinga. I. Detail of the pendant fruit.
Morro, 12°42′57″S, 39°45′46″W, 285 m, 8 Apr 2006 (fl), E.
Melo, F. França, C. T. Lima and C. Cunha 4344 (holotype:
HUEFS 109249!).
A Capsicum parvifolium Sendtn. denso indumento ramosis
trichomatibus, calyce longis dentibus conspicuis, corolla alba
viridulis vel flavidis maculis differt.

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Slender or semiscandent shrubs 1.5–4 m tall; stems few
branched, grayish, fragile, striate; old stems glabrous;
young stems densely pubescent with branched hairs. Leaves
geminate, or solitary, 2–2.5 times longer than broad, (3–)3.5–
5.4 cm long, 1.5–2.5 cm wide, ovate, membranous, slightly
discolored, pubescent, with abundant branched hairs on both
surfaces, especially abaxially, the margin entire, the apex
acuminate, the base short-attenuate or truncate and unequal;
petioles (0.3–)0.5–1.5 cm. Flowers in fascicles of (1–)2–4; the
flowering pedicels pendant, not geniculate (straight) at anthe-
sis, (0.65–)0.8–2.1 cm, with dendritic hairs. Flower buds ovoid,
whitish green. Calyx green, densely pubescent with branched
hairs, the tube 2–5 mm, with 5 conspicuous linear and sub-
equal teeth (4.5–)5–8.5 mm long. Corolla stellate, 6–7.2 mm
long, lobed nearly half way to the base, pure white abaxi-
ally, white with two pale yellow or greenish yellow spots and
small glandular hairs at the base of each lobe and in the throat
adaxially; lobes equal to the tube, ca. 3.3–3.6(–4) mm long,
2–2.5 mm wide, broadly triangular, the tip cucullate, densely
pubescent on the tips and involute margins. Filaments
greenish white, ca. 1.25(–2) mm; anthers yellowish cream, ca.
2–2.5 mm long; stapet 1.5 mm long. Ovary greenish white,
ovoid, ca. 2 mm long, 1.3 mm wide; style greenish white, 3.5–
3.75 mm, widening distally; stigma green, somewhat discoid.
Berry globose, slightly flattened at the apex, ca. 7.5–9.5 mm
long, 7–8.5 mm wide, green when immature, yellowish green
at maturity; fruiting pedicels pendant, the fruiting calyx per-
sistent, the pericarp sweet to the taste and lacking stone cells.
Seeds (3–)5–15(–17) per fruit, brownish tan, 3–3.7 mm long,
2.5–2.8 mm wide; testa thick, foveolate, with spine-like pro-
jections, the fovea deep at the seed margin and shallow on
the central zone, with sinuate and thick cell walls. Figures 2 ,
3F, H, I , 4 .
Etymology— The species epithet refers to the conspicuous
calyx teeth which are the longest observed in any Capsicum
species.
Phenology— Flowering from December to April; fruiting
from January to April.
Distribution and Habitat— This species is endemic to the
center of Bahia and Pernambuco states ( Fig. 2 ), growing on
granitic hillsides with shrubby open dry Caatinga vegetation
and in gallery forests along small rivers.
Conservation Status— Capsicum longidentatum is a narrowly
distributed species restricted to areas that are experiencing
significant human disturbance. According to the IUCN cat-
egories, it should be considered a threatened species and we
tentatively assign it a status of critically endangered (CR).
Observations— Capsicum longidentatum differs from other
members of the genus by its relatively long calyx teeth and
its peculiar pubescence. This species is the only one in which
the calyx teeth reach 8.5 mm in length. Two other Capsicum
species from Brazil (i.e. C. cornutum (Hiern) Hunz., C. hun-
zikerianum Barboza & Bianchetti), and C. hookerianum (Miers)
Kuntze) from Ecuador and Perú, have calyces with relatively
long teeth but in these taxa the teeth are not more than 6 mm
in length ( Barboza and Bianchetti 2005 ). Capsicum longiden-
tatum is the only species of the genus where the indumen-
tum consists of a variety of types of branched hairs (bifurcate,
trifurcate, and dendritic hairs; Fig. 5 ) on the young stems,
leaves, pedicels, and calyces; these give the species a distinc-
tive shaggy appearance. Branched hairs are uncommon in
Capsicum and in general if they are present they are mixed
with simple nonglandular trichomes covering stems and
leaves (e.g. C. rhomboideum ) or only present on abaxial leaf
veins (e.g. C. scolnikianum ); branched hairs occasionally occur
on some pubescent specimens of C. parvifolium and C. caat-
ingae . Finally, C. longidentatum fruits are completely free of
pungency, as in some other Capsicum : C. lanceolatum, C. dimor-
phum (Barboza, pers. obs.), and C. rhomboideum ( Bosland and
Zwedie 2001 ).
Karyology— A somatic chromosome number of 2 n = 2 x = 24
was found ( Table 1 ). The karyotype of the species comprises
12 m chromosome pairs, where pair no. 12 has NORs on the
short arms ( Fig. 6A ). A heteromorphic condition is found with
one homologous chromosome bearing a microsatellite and
the other a macrosatellite. The total length of the basic chro-
mosome complement is 76.11 μm.
Additional Material Examined— BRAZIL. Bahia: Brumado, Fazenda
Lagoa Redonda, 14°17’S, 41°33’W, 500 m, 26 Mar1984, de Oliveira Filho &
Lima 131 (CEPEC, MO). Itatim, Cerca de Pedra Resoles, Morro detrás del
Restaurant Resoles, 12°44′21.8″S, 39°47′01.8″W, 295 m, 3 Apr 2009, Agra
et al. 7083 (CORD, JPB). Morro do Quixaba, 12°44′21.3″S, 39°47′33.9″W,
273 m, 3 Apr 2009, Agra et Barboza 7086 (CORD, JPB). Inselbergue Morro
da Quixaba, 12°44′26″S, 39°47′37″W, 290 m, 12 Feb 2006, Moraes et al.
161 (HUEFS). Morro do Quixaba, 12°44’S, 39°47’W, 276 m, 14 Dec 1996,
Melo et al. 1859 (HUEFS). Morro do Bastião, 12°45′12″S, 39°46’W, 282 m,
25 Jan 1997, Melo et al. 1920 (HUEFS, JPB). Serra de Itiúba, about 6 km
E of Itiúba, 10°41’S, 39°48’W, 500 m, 19 Feb 1974, Harley 16184 (CEPEC,
RB). Pernambuco : Mirandiba, Serra do Tigre, 08°03′35″S, 38°43′07″W,
495 m, 18 Apr 2007, Pinheiro 253 (CORD, JPB, UFP).
Capsicum parvifolium Sendtn., Fl. Bras. 10: 145. 1846.—
TYPE: [BRAZIL]. “In prov. Bahiensi: Blanchet (n. 2823) ”
(holotype: B†,[photo F neg. #2872]; lectotype here
designed: Blanchet n. 2823, W-118245!; isolectotypes: BM!
[BM001016675, BM000617948], F! (fragment), G! [F neg. #
6864], K! [sub nom. Fregirardia leptoclada ], P-124/76!).
Fregirardia leptoclada Dunal, Prodr. 13 (1): 505. 1852.—TYPE:
BRAZIL. “In Brasiliae provinciâ Bahiensi ad pedem
montis Acurua in Sertao do rio de S. Francisco ( Blanchet,
n. 2823 , in h. DC. H. Moric. et h. Boiss.)” (holotype: G!;
isotypes: BM! [BM001016675, BM000617948], F! (frag-
ment), G! [F neg. # 6864], K !, P!).
Bassovia ciliata J. R. Johnston, Proc. Amer. Acad. Arts 40: 694.
1905.—TYPE: VENEZUELA. [St. Nueva Esparta: Is.
Margarita] El Valle, 30 Aug 1903, J. R. Johnston 75 (lecto-
type designated here: GH!; isolectotype: US-531916!).
Shrub 1–4(–5) m tall, with an almost fastigiate habit; stems
much branched, slender, fragile, grayish, striate; old stems gla-
brous; young stems pubescent with multicellular nonglandu-
lar hairs. Leaves solitary or geminate, 1.5–3 times longer than
broad, (1–)2.3–6.5(–9) cm long, 0.7–4.2 cm wide, ovate, mem-
branous, slightly discolor, upper surface light green, sparsely
pubescent, underside paler and sometimes pubescent, the
hairs long and simple on both surfaces, rarely with a few den-
dritic hairs on the veins abaxially, the margin entire, the apex
acuminate, the base shortly attenuate and unequal; petioles
canaliculate adaxially, (0.3–)0.5–2.4 cm, pubescent. Flowers in
fascicles of 3–6; pedicels pendant, not geniculate (straight) at
anthesis, 0.9–1.8(–2) cm, pubescent, with short simple nong-
landular and antrorse hairs. Flower buds ovoid, violet at the
apex and cream near the base. Calyx green, pubescent like
the pedicels, the tube (1.2–)1.5–2.4 mm long, 5-toothed or
exceptionally up to 7-toothed, the teeth conspicuous, patent
or oblique, subequal, 0.7–2(–2.2) mm long. Corolla stellate,
4.5–7.3 mm long, 1–2 cm diam, whitish green abaxially, the
lobes and part of the limb purple adaxially, the tube cream or
yellowish white, the margin white with small glandular hairs

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774 SYSTEMATIC BOTANY [Volume 36
Fig . 4. Capsicum longidentatum (A, B, D, G, H, L: Melo et al. 4344 ; C, E, F, I, J, K: Agra and Barboza 7086 ). A, D. Anthers, ventral and dorsal view, respec-
tively. B. Flowering branch. C. Flower bud. E. Detail of a flowering node. F. Embryo. G. Open corolla (one stamen has been removed). H. Gynoecium.
I. Seed. J. Fruit. K. Fruiting calyx. L. Flower.

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Fig . 5. Types of trichomes in C. longidentatum . A-C, F. Leaf trichomes. D, E, G, J. Calyx trichomes. H, I, K-N. Corolla trichomes. A, N. Bifurcate. B, D,
L, M. Trifurcate. C, E, G. Dendritic. F, J. Simple, glandular hairs. H, I, K. Simple, non glandular hairs. Scale bar = 50 μm.

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776 SYSTEMATIC BOTANY [Volume 36
(unicellular head, bicellular stalk) at the base of the lobes
and in the throat adaxially; lobes ca. (2.1–)2.9–3.5 mm long,
2–3.1 mm wide, broadly triangular, densely papillate on the
tips and the involute margins, or with long simple hairs, the
tip cucullate. Filaments greenish white, 1.5–2.5 mm; anthers
yellow, 1.7–2.3 mm long; stapet 1.3–1.5 mm long. Ovary
greenish white, ovoid, ca. 1–1.4 mm long, 1 mm wide; style
whitish green, 3.5–5.1 mm long, widening distally; stigma
green, somewhat discoid. Berry globose, slightly flattened,
ca. 8.5–9.5 mm diam, dark green when immature, yellowish
green at maturity; fruiting pedicels pendant and curved, not
geniculate, the fruiting calyx persistent, the teeth reflexed, the
pericarp hot to the taste and lacking stone cells. Seeds (2–)4–
13 per fruit, blackish brown, 3.8 mm long, 2.7–3 mm wide;
testa foveolate with spine-like projections, the fovea deep
at the margin and central area, with thick sinuate cell walls.
Figures 2 , 3A-E , 7 .
Typification— Capsicum parvifolium was described based on
a specimen of Blanchet 2823 from Bahia; it is often assumed
that Sendtner’s types were at B. The sheet of Blanchet 2823 at
B was destroyed during the Second World War ( Hiepko 1987 ).
A photograph of this sheet (F neg. # 2872) is held in the Type
Photograph Collection at F. Dunal (1852) described Fregirardia
leptoclada from a duplicate of the same collection housed at G
(F neg. # 6864); this name is a nomenclatural synonym (ICBN
art. 14.4, McNeill et al. 2006 ). Duplicates of Blanchet’s collec-
tions were distributed in many herbaria ( Stafleu and Cowan
1976 ), and the localization of the collections from the state
of Bahia remains difficult ( Hind 1999 ). Among the syntypes
found, we select the collection at W as lectotype of Capsicum
parvifolium since it is the best preserved sheet consisting of a
complete branch, with flowers and fruits.
Johnston (1905) described Bassovia ciliata from two speci-
mens collected by him on the Venezuelan Isla Margarita (“El
Valle, river trail, Miller & Johnston, n° 255 , July 15, 1901, and
Johnston, n° 75 , Aug. 30, 1903”). Both specimens agree with the
diagnosis; the first syntype ( Miller and Johnston 255 ) is depos-
ited at GH, NY, and US, but the material from the three sheets
is poor; in contrast, Johnston’s collection designed here as lec-
totype is of better quality with many buds and flowers.
Vernacular Names— “Pimentinha” (Pernambuco), “Piment-
eira” (Paraíba) “Alecrim-quebrado (Piauí)”, “Jiriquiti” (Piauí).
Phenology— Flowering from November to April and
fruiting from February to June in the caatinga; flowering
from April to October and fruiting from May to October in
Venezuela and Colombia.
Distribution and Habitat— This species has a discontin-
uous distribution in South America (northeastern Brazil,
Venezuela, and Colombia). It is a common element of the
arborescent-shrubby closed or open Brazilian Caatinga (Bahia,
Minas Gerais, Paraíba, Ceará, Pernambuco, Piauí, Río Grande
do Norte), in areas with sandy and stony soils (‘inselbergs’),
at 600–1,200 m. In Venezuela and Colombia, C. parvifolium
also inhabits dry forests, or occurs on steep rocky slopes at
lower altitudes (0–350 m).
Conservation Status— Capsicum parvifolium is not consid-
ered to be under threat at this time; it is widely distributed
and is common where it occurs.
Notes— Capsicum parvifolium is an enigmatic species that
has been confused in the literature. Morphologically, the fruits
have been described as red or orange-colored ( Pozzobon et
al. 2006 ; Moscone et al. 2007 ), with pale yellow or brownish
yellow seeds ( Barboza and Bianchetti 2005 ; Moscone et al.
2007 ). According to our observations in the field, the fruits at
maturity are clearly yellowish green ( Fig. 3A ) as occurs in the
remaining species from Brazil, with the exception of C. ann-
uum var. glabriusculum , C. baccatum , C. praetermissum Heiser
& Smith, C. coccineum , and C. flexuosum where fruits are red
( Barboza and Bianchetti 2005 ). The seeds of C. parvifolium are
blackish brown, not pale ( Fig. 3E ).
Hunziker (2001 : 235, Fig. 80) published a detailed illustra-
tion under the name Capsicum parvifolium based on different
specimens: Pickersgill 72–366 and Hunziker 25233 . Hunziker
had a broad and inclusive species concept for C. parvifolium
(Hunziker in mss.), especially with respect to the number
of flowers per inflorescence and the presence or absence of
calyx teeth. The two specimens used by Hunziker (2001) were
analyzed here and we consider Pickersgill 72–366 to belong to
C. parvifolium and Hunziker 25233 to C. caatingae .
Capsicum parvifolium resembles C. caatingae (see above);
nevertheless, it is clearly distinguishable by the 5-toothed
calyx ( Fig. 7E ), the fascicle with fewer than nine flowers
( Fig. 7C ), and the blackish brown seeds ( Fig. 3E ). Specimens
of C. parvifolium from Venezuela have been confused in the
Table 1. Chromosome measurements in Capsicum longidentatum ( Agra & Barboza 7086 ) and Capsicum parvifolium ( Agra & Barboza 7075 ). Abbreviations:
r, arm ratio; l, long arm length; s, short arm length; HKL, haploid karyotype length; x, mean value; sd, standard deviation; m, metacentric chromosome;
sm, submetacentric chromosome; m-sat, metacentric chromosome with secondary constriction and satellite.
Pair
C. longidentatum C. parvifolium
Length r
(l/s)
Type
Length r
(l/s)
Type
Absolute (μm)
X (sd) Relative (% of HKL) X (sd) Absolute (μm)
X (sd) Relative (% of HKL) X (sd)
1 7.61 (0.72) 10.00 1.11 (0.06) m 8.38 (1.05) 10.44 1.06 (0.03) m
2 7.26 (0.59) 9.54 1.17 (0.08) m 7.61 (0.94) 9.48 1.21 (0.14) m
3 7.02 (0.55) 9.22 1.31 (0.18) m 7.29 (1.08) 9.09 1.13 (0.12) m
4 6.89 (0.47) 9.05 1.17 (0.16) m 7.15 (0.93) 8.91 1.26 (0.07) m
5 6.73 (0.46) 8.84 1.19 (0.12) m 7.02 (0.82) 8.74 1.29 (0.14) m
6 6.47 (0.35) 8.50 1.20 (0.19) m 6.86 (1.06) 8.56 1.17 (0.04) m
7 6.29 (0.52) 8.26 1.21 (0.14) m 6.65 (0.76) 8.29 1.20 (0.18) m
8 6.26 (0.48) 8.23 1.20 (0.15) m 6.42 (0.59) 7.99 1.24 (0.11) m
9 6.07 (0.42) 7.97 1.19 (0.13) m 6.27 (0.96) 7.81 1.21 (0.13) m
10 5.87 (0.30) 7.71 1.15 (0.07) m 6.07 (1.97) 7.56 1.28 (0.09) m
11 5.15 (1.14) 6.77 1.41 (0.07) m 5.55 (1.02) 6.92 1.50 (0.09) m
12 4.49 (0.42) 5.89 1.65 (0.21) m-sat 4.99 (1.55) 6.22 1.84 (0.12) sm-sat

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2011] BARBOZA ET AL.: THE CAPSICUM SPECIES FROM THE BRAZILIAN CAATINGA 777
herbaria with C. rhomboideum due to their similarities in the
calyx; however they can easily be differentiated by the corolla
shape and color (campanulate and yellow in C. rhomboideum ),
and fruit color at maturity (red in C. rhomboideum ).
Karyology— A somatic chromosome number of 2 n = 2 x = 24
was found ( Table 1 ). The karyotype of the species comprises
11 m chromosome pairs and 1 sm pair, where pair no. 12 pos-
sesses NORs on the short arms ( Fig. 6B ). The heteromorphic
condition found is with one homologous chromosome bearing
a microsatellite and the other one carrying a macrosatellite. The
total length of the basic chromosome complement is 80.26 μm.
Additional Material Examined— BRAZIL. Bahía: Maracas, km 6–7 de
la route Maracas-Contendas da Sincorá, 25 Nov 1986, Chautems et al. 165
(CEPEC). Chapada Diamantina, 29 Apr 2006, Guedes et al. 12268 (ALCB).
Jequié, Morro da Torre, 13°53′27″S, 40°07′20″W, 671 m, 13 Apr 2007, Queiroz
et al. 12964 (HUEFS). Camino para Serra Preta, 12°31′55″S, 41°35′14″W, 800
m, 28 Mar 2003, Melo et al. 3646 (HUEFS). Minas Gerais: Manga, Gleba B,
2 Dec 1989, Horta et al. 126 (BHCB). Paraíba: Maturéia, Pico do Jabre, 30
Mar 2009, Agra et Barboza 7075 (JPB). Campina Grande, 24 Jun 1935, Pickel
3892 (US). Distr. São José da Mata, Fazenda Pedro da Costa Agra, estrada
para Soledade, 16 km O do centro de Campina Grande, 7°46’S, 35°52’W,
500–510 m, 2 Apr 1988, Agra 658 (JPB). Pico do Jabre, ao Norte da sede da
Telpa, em direção Leste, 07°11′10″N 037°25′53″W, 800–1010 m, 22/23 Apr
1998, Agra 5249 (JPB, MO). São José dos Cordeiros, RPPN-Fazenda Almas,
Area I, trilha para casa de Comadre Florzinha, 07°28′37″S, 36°53′16″W,
15 Feb 2003, Barbosa et al. 2674 (JPB); RPPN-Fazenda Almas, camino para
Pedra da Bola, 3 Mar 2004, Barbosa et al. 2895 (JPB). Ceará: Ceará, s. f., F.
Allemão 1229 (US); Fazenda Itatiaia, Santa Quitéria, 26 Apr 1984, Fernández
et al. s. n. (JPB). Pernambuco: Alagoinha, no Km 8 da BR 232, faz. Cajueiro
Seco, 14 Jan 1998, Melo et al. 221 (SP). Tapéra, Gravatá, Mount “Cruzeiro,”
Jul 1926, Pickel 2492 (US). Sertânia, Centro da Caprino-Ovino-Cultura do
I. P. A., 8°4’25″S, 37°12′24″W, 610 m, 4 Apr 2001, Harley & Giulietti 54171
(HUEFS). Buique, 27 Feb 2004, Lucena 91 (JPB). Piauí: Serra da Lagoa, Jan
1907, Ule 7475 (L). São José do Piauí, Morro do Baixio, 21 Feb 2000, Mendez
et al. 292 (JPB). Río Grande do Norte: South of Cerro Cora, 700 m, Mar1972,
Pickersgill RU72–366 (CORD).
COLOMBIA. Atlántico: Savanilla, 31 May 1874, Kuntze 1811 (NY).
Barranquilla and vicinity, Oct 1928, Elías 576 (US). Magdalena: Santa
Marta, near Mamatoca, 100 ft, 23 Aug 1898/1899, Smith 1176 (BM, CORD,
E, GH, K, L, MO, NY, P, US). Santa Marta, hill 3 miles inland from Playa
Brava, 500 ft, 6 Jun 1898/1899, Smith 1183 (F, K, NY, US).
VENEZUELA. Andes, Jul 1930, Rutten-Peckel Harig 4 (U). Distrito
Federal: Vargas, Parroquia Catia La Mar, Esc. Naval, ca. 10°36’N, 67°02’W,
5 m, 10 Oct 1988, Ramírez 2548 (VEN). Aragua: Ocumare, s. f., Pittier 14032
(US); M. Briceño Iragorry, 4–5 km al NE de Bahía de Cata, 150 m, 13
Apr 1981, Carnevali et al. 531 (VEN). Sendero Cata-Catica, PN H. Pittier,
10 May 1992, Castro & Magallanes 39 (MY, VEN). Carabobo: Distrito
Valencia, margen derecha del Río Las Marías, Carretera La Belén-Las
Marías desde Trincheras, 350 m, 25 Aug 1991, Benítez de Rojas & Rojas 4202
Fig . 6. Somatic metaphase chromosomes and idiograms of Capsicum longidentatum (A) and C. parvifolium (B), both with 2 n = 24.

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778 SYSTEMATIC BOTANY [Volume 36
Fig . 7. Capsicum parvifolium (A-D, F, H, I: Agra and Barboza 7075 ; E, G: Pickersgill 72–366 ). A. Flower. B, D. Anthers, dorsal and ventral view, respec-
tively. C. Fruiting branch. E. Calyx. F. Gynoecium. G. Dendritic trichome. H. Open corolla. I. Fruits.

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Key to Differentiate the Endemic Species from Brazil
The 11 endemic species of Capsicum from Brazil and C. parvifolium are characterized by having yellowish green fruits at matu-
rity. The three species from the Caatinga share the following suite of characters: habit (relatively low slender shrubs), stems
(fragile, striate, grayish, and much-branched), leaf size (in general, small and deciduous), and flower size (small, not more than
8 mm long). The following key to the Brazilian species is adapted from Barboza and Bianchetti (2005) and modified to include
C. caatingae and C. longidentatum .
1. Corolla entirely pink or lilac, clearly campanulate to urceolate, tube (5.5–)7–9(–11) mm;
leaves generally ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1. C. friburgense Bianchetti & Barboza
1. Corolla white with yellowish green and sometimes also with purple spots inside, or purple with a yellowish green center,
stellate, never campanulate-urceolate, the tube (2–)2.6–6(–8) mm; leaves ovate, elliptic or narrowly elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
2. Corolla white with two yellowish green or golden spots at the base of each lobe and on the internal surface of the throat . . . . . . . . . . . . . . . . . . 3
3. Calyx toothless; corolla with golden spots; ovules 2 per locule; androecium heterodynamous
with 3 short stamens and 2 long stamens; fruits globose-compressed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2. C. campylopodium Sendtn.
3. Calyx with 5, or 6–9 teeth; corolla with greenish yellow spots; ovules 5–10 or more per locule;
androecium homodynamous with all stamens equal in length; fruits globose or globose-flattened . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4
4. Plants entirely covered by branched hairs; pedicels pendant, nongeniculate (straight at anthesis);
calyx teeth (4.5) 5–8.5 mm long, erect or oblique. Plants from the Caatinga . . . . . . . . . . . . . . . . . . . . 3. C. longidentatum Agra & Barboza
4. Plants entirely covered by simple hairs, branched hairs absent; pedicels erect, geniculate at anthesis
(flowers longitudinal axis at 90° to pedicel); calyx teeth 1–2 mm long, recurved.
Plants from the Atlantic Forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4. C. recurvatum Witas.
2. Corolla white with purple spots near the base of each lobe with an interrupted yellowish-green zone in the lobes and
limb or lobes more distally, almost completely purple with yellowish-green throat inside . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
5. Pedicels pendant and nongeniculate (straight) at anthesis . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
6. Flowers solitary or the fascicles 2–3-flowered; corolla 9–10 mm long, with small purple spots at the base of the lobes;
leaves coriaceous, elliptic to narrowly elliptic, 3–5.5(–10) times longer than broad, glabrate.
Plants from the Atlantic Forest . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5. C. pereirae Barboza & Bianchetti
6. Fascicles (3–)6–18-flowered; corolla 4–8 mm long, almost entirely purple, except the white margin;
leaves membranous, ovate or elliptic, 2–2.5 times longer than broad, glabrescent to pubescent. Plants from the Caatinga . . . . . . . . . . 7
7. Calyx 5-toothed; inflorescence 3–6-flowered; seeds blackish brown . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6. C. parvifolium Sendtn.
7. Calyx toothless; inflorescence 5–13(-18)-flowered; seeds yellowish tan . . . . . . . . . . . . . . . . . . . . . . . . . . 7. C. caatingae Barboza & Agra
5. Pedicels erect and geniculate at anthesis (flowers longitudinal axis at 90° to pedicel) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8
8. Calyx toothless or sometimes with 5 tiny teeth . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 8. C. schottianum Sendtn.
8. Calyx 5–10-toothed . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
9. Calyx with only 5 short teeth (0.5–3 mm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10
10. Plants glabrescent, the hairs antrorse; leaves elliptic to narrowly elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9. C. mirabile Mart.
10. Plants densely pubescent, the hairs flexuous and spreading on stems, petioles, pedicels,
and sometimes also on the leaf veins beneath; leaves ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 10. C. villosum Sendtn.
9. Calyx up to 6–10 long teeth (3.2–6 mm long) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
11. Plants densely pubescent; corolla (8–)9–12(–14) mm long, the lobes mostly white;
leaves membranous, ovate to broadly ovate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11. C. cornutum (Hiern) Hunz.
11. Plants glabrate; corolla 10–14(–16) mm long, the lobes mostly purple; leaves coriaceous,
slightly ovate to elliptic . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12. C. hunzikerianum Barboza & Bianchetti
(VEN, MY). Porto Cabello, [year] 1917, Curran & Haman 1181 (GH, US).
Lara: Iribarren, arriba del caserío Simara, ca. 8 km de Bobare, 20 May 1978,
Burandt et al. 0193 (VEN); Urdaneta, 8 km vía desde Aguada Grande vía
Siquisique, 15 Jul 1980, Smith 9349 (VEN). Sucre: Isla Los Venados, south
of Playa Las Barracas Azules, 0–50 m, 10 Sep 1973, Steyermark et al. 108086
(US). Playa Manzanillo, west of Hotel Cumanagoto and Cumaná, 0–2 m,
8 Sep 1973, Steyermark et al. 108194 (MO, US).
Discussion
The two species examined are diploid with x = 12 as are all
other Capsicum species ( Hunziker 1979 ). While the presence
of satellites is not a constant feature in Solanaceae ( Wijsman
et al. 1983 ; Moscone 1989 ; Chiarini and Barboza 2008 ), it is
common in the species studied here, as is the rule in Capsicum
in general ( Pickersgill 1971 ; Moscone 1990 ). The polymor-
phism found in C. longidentatum and C. parvifolium is related
only to the sizes of satellites. The total karyotype length of
both species is among the largest of the species studied in
Capsicum ( Moscone et al. 2007 ).
The results for the sample of C. parvifolium studied here
are in disagreement with cytotype 1 and 2 described under
the same name (recognized here as C. caatingae ) by Moscone
et al. (2007) . Capsicum caatingae was until now the only species
in the genus with a karyotype composed entirely of m chro-
mosomes ( Moscone 1993 , as C. parvifolium ), but our results
show this also occurs in C. longidentatum . Considering karyo-
type formula, haploid karyotype length (HKL), number and
position of satellites, as well as morphological features and
geographical distribution, C. longidentatum could be closely
related to C. caatingae (as C. parvifolium cytotype 2), but addi-
tional data from chromosome banding may be needed to
show these relationships ( Moscone et al. 2007 ).
Acknowledgments. We are indebted to the curators and assistants
of the herbaria at ALCB, BHCB, BM, CEPEC, COL, CORD, CTES, E, F,
G, GH, HRB, HUEFS, JPB, K, KPB, L, MBM, MO, MY, NY, P, RB, SP, U,
US, VEN, and W who either loaned material for study or allowed us to
examine material under their care, to C. Niezgoda (F) for providing digi-
tal images of type specimens, S. Knapp and the two anonymous review-
ers for helpful comments on the manuscript; thanks are also tendered to
L. Ribulgo and P. Peralta for making illustrations and maps. Financial assis-
tance was provided by Consejo Nacional de Investigaciones Científicas
y Técnicas (CONICET, PID 6188; PIP 01686), Agencia Nacional de
Promoción Científica y Tecnológica (PICT 20196), Ministerio de Ciencia y
Tecnología de la Provincia de Córdoba, Secretaría de Ciencia y Tecnología
(SECYT-UNC), Myndel Botanica Foundation, and Conselho Nacional de
Desenvolvimento Científico e Tecnológico (CNPq), Ministério da Ciência
e Tecnologia do Brasil.
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