Taxonomic Revision of Aloysia (Verbenaceae, Lantaneae) In
South America
Author(s): Nataly O'Leary , Patricia Lu-Irving , Pablo Moroni Stephen Siedo
Source: Annals of the Missouri Botanical Garden, 101(3):568-609.
Published By: Missouri Botanical Garden
DOI: http://dx.doi.org/10.3417/2013015
URL: http://www.bioone.org/doi/full/10.3417/2013015
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TAXONOMIC REVISION OF
ALOYSIA (VERBENACEAE,
LANTANEAE) IN SOUTH
AMERICA1
Nataly O’Leary,2 Patricia Lu-Irving,3 Pablo
Moroni,2 and Stephen Siedo4
ABSTRACT
Aloysia Paláu is the third largest genus of tribe Lantaneae, after Lippia L. and Lantana L., in the Verbenaceae. Recent
molecular phylogenetic studies have circumscribed genus Aloysia as 31 species, with the transfer of most species of
Acantholippia Griseb. and the inclusion of the monotypic Xeroaloysia Tronc., as well as the exclusion of several North American
Aloysia species that nest within a Lippia–Lantana clade. Newly circumscribed Aloysia are found mostly in South America, where
the genus is represented by 28 species and six varieties. Only four Aloysia species are found in North America, A. coalcomana
Siedo, A. macrostachya (Torr.) Moldenke, A. wrightii A. Heller, and A. gratissima (Gillies & Hook.) Tronc. var. gratissima, this
last being the only taxon found in both North and South America. A taxonomic revision of the genus Aloysia for South America is
provided with detailed morphological descriptions, as well as keys for taxonomic identification, illustrations or indication of
iconography, and distribution and herbarium specimen lists. The genus Xeroaloysia Tronc. is here considered a synonym of
Aloysia, and nine new taxonomic synonyms are here established. Lectotypification is designated for Verbena L. sect. Aloysioides
Walp., and neotypification is designated for V. salviifolia Hook. & Arn.
Key words: Acantholippia, Aloysia, South America, Verbenaceae.
Recent molecular phylogenetic studies dealing
with generic limits in the Lantaneae (Verbenaceae)
have suggested that Aloysia Paláu, as traditionally
circumscribed, may be a polyphyletic group (Marx et
al., 2010; Lu-Irving & Olmstead, 2013). A subsequent study that focused on resolving the position of
Aloysia and related genera (Lu-Irving et al., 2014),
using both chloroplast and nuclear gene sequences,
confirmed the non-monophyly of traditional Aloysia.
These authors proposed the inclusion within Aloysia
of most taxa in Acantholippia Griseb. and the
monotypic genus Xeroaloysia Tronc., and the exclusion of several North American taxa nested in the
Lantana L.–Lippia L. clade (Lu-Irving et al., 2014:
649, fig. 4), to be able to recover a well-supported
monophyletic Aloysia. Consequently, this new circumscription of Aloysia comprises 31 species, which
occur in four major and consistently inferred clades.
One clade groups the majority of Aloysia taxa; a
second clade groups the type species of Aloysia, A.
citrodora Paláu, plus another species that shares its
inflorescence morphology; a third clade reunites two
South American Aloysia taxa; and a fourth clade
groups the type species of Acantholippia together with
the rest of the taxa formerly in Acantholippia. The
Lantana–Lippia clade is a strongly supported group
that includes Phyla Lour. and Nashia Millsp., as well
as three Mexican species of Aloysia (A. barbata
(Brandegee) Moldenke, A. chiapensis Moldenke, and
A. sonorensis Moldenke), and Acantholippia seriphioides (A. Gray) Moldenke.
Aloysia is the third largest genus of tribe
Lantaneae, after Lippia and Lantana. Aloysia is
distinguished from Lantana by its dry fruit, being
fleshy in the last, and from Lippia by its long
racemose florescences with the rachis longer than the
peduncle, and alternate or opposite flowers; in Lippia,
florescences are capituliform with a brief rachis and
longer peduncle, and flowers are spiraled. Sanders
(2001: 310) states Aloysia is based on two synapo-
1 We thank the curators of the herbaria cited in the text for the loan of specimens and assistance in the search of type
material and bibliography, especially S. Beck from LPS; A. Smith from E; G. Pieszko from CTES; M. Garcı́a from MVM; R.
Russell from US; T. Wendt from TEX; and T. Zanoni from NY. This study was possible thanks to a grant
(PIP112200801000177/09, Consejo Nacional de Investigaciones Cientı́ficas y Técnicas [CONICET]) given to the authors. A
special thanks to Marı́a E. Múlgura who preferred not to be an author despite her important contribution. An earlier version
of the manuscript benefited greatly thanks to suggestions given by M. Belgrano. Special thanks also to Federico Luebert,
from Bonn, Germany, for just-in-time comments, and to Victoria Hollowell for her significant editorial work. Illustrations
were done by Jennifer Castelo, Vladimiro Dudas, and Elio Vega, to whom the authors are very grateful.
2 Instituto de Botánica Darwinion, Labardén 200, CC 22 (B1642HYD), San Isidro, Buenos Aires, Argentina. Author for
correspondence: noleary@darwin.edu.ar.
3 Department of Ecology and Evolutionary Biology, 1401 E Lowell Street, University of Arizona, Tucson, Arizona 85721,
U.S.A.
4 Plant Resources Center, University of Texas, Austin, Texas 78712, U.S.A.
doi: 10.3417/2013015
ANN. MISSOURI BOT. GARD. 101: 568–609. PUBLISHED
ON
27 APRIL 2016.
Volume 101, Number 3
2016
O’Leary et al.
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Revision of Aloysia (Verbenaceae) in South
America
morphies, ‘‘inconspicuous, ephemeral floral bracts
and involute calyx lobes.’’ Lu-Irving et al. (2014)
noted the 4-lobed calyx as the defining feature for
newly circumscribed Aloysia, as had been suggested
by previous studies (O’Leary et al., 2012).
Aloysia as newly circumscribed is almost exclusively found in South America, with only three
species restricted to North America. Aloysia wrightii
(A. Gray) A. Heller and A. macrostachya (Torr.)
Moldenke occur in the southwestern United States,
extending into Baja California and across northern
Mexico; A. coalcomana is only known from one
locality in Michocán, Mexico. These three species are
not further considered here. Lu-Irving et al. (2014)
stated that shifts in geographic range from subtropical
South America to North America must have occurred
at least twice in Aloysia, based on the reconstruction
of the geographical distribution of species in the
phylogenetic trees (2014: 651, fig. 5). However,
based on these results, Lu-Irving et al. (2014) could
not ascertain if the three North American Aloysia taxa
were the result of long-distance dispersal or northward migration via the Andes.
Lu-Irving and Olmstead (2011; 2013) and LuIrving et al. (2014) found South American Aloysia
comprised of several small lineages and one larger
lineage containing the majority of Aloysia species.
This large clade includes the formerly monotypic
genus Xeroaloysia as well as the North American
Aloysia taxa; all exemplars have homothetic pleiobotrya (only axillary florescences). This species assemblage seems to represent a recent radiation, because
branch lengths were short throughout the clade (LuIrving et al., 2014). Aloysia species with heterothetic
pleiobotrya (both axillary and terminal florescences)
grouped together in another clade, in basal position
relative to the larger Aloysia clade. Taxa formerly in
Acantholippia with monobotrya (only terminal florescences) also formed a clade, basal to the rest of
Aloysia. Consequently, inflorescence morphology
seems to have phylogenetic significance in Aloysia.
There has been no comprehensive taxonomic
revision of Aloysia. A Ph.D. dissertation, which
unfortunately lacks illustrations (Siedo, 2006), considered 30 species and 14 varieties, seven species
from North America and the remaining 23 species
from South America. This work included a phylogenetic analysis based on 50 morphological characters,
and the results supported the monophyly of Aloysia.
However, the author mentioned that preliminary
molecular phylogentic analysis conducted by the
Olmstead et al. laboratory suggested the contrary,
resulting in a polyphyletic Aloysia, and that additional studies were needed. This further investigation
was performed by Lu-Irving et al. (2014), with the
taxonomic implications being followed in this present
work.
Partial treatments have been done for Aloysia in
Argentina (Troncoso, 1962, 1974; Botta, 1979, 1993;
Múlgura et al., 2012), recognizing ca. 10 species. In
Bolivia, Wood (2009) recognized seven species, and
in Venezuela, López Palacios (1977) mentioned one
species. However, no overall revision of the entire
genus has ever been published. Four new species of
Aloysia were recently described by Siedo (2012) for
Brazil, Peru, and Mexico.
Revision of Aloysia for South America is presented
here, following new taxonomic circumscription as
suggested by the phylogenetic evidence of Lu-Irving
et al. (2014). Illustration or reference to iconography,
citation of representative material, synonymy, and
any pertinent typification are included for the 28
species and six varieties in South America as
considered herein.
MATERIALS
AND
METHODS
This taxonomic revision is based on collections
from the following herbaria: BAF, CONC, F, K, LIL,
MO, NY, P, SGO, SI, ULS, US, and WTU. Flower
measurements were taken from material rehydrated
by boiling. Fruit measurements were taken from dried
specimens. The descriptive terminology of the
inflorescences used here is in accordance with
Múlgura et al. (2002), the morphological terms follow
Hickey (1974), and the description of pubescence
corresponds to that of Lawrence (1951). Inflorescence
morphology follows terminology used by Múlgura et
al. (2002), which follows Troll (1964–1969) and Sell
(1976, 1980). Distribution and habitat data for taxa
were taken from the herbarium specimen labels. A
list of accepted species and varities of Aloysia
(Appendix 1), an index to collectors (Appendix 2),
and a list of taxa newly synonymized here with their
accepted range (Appendix 3) are provided.
TAXONOMIC TREATMENT
Aloysia Paláu, Parte Práct. Bot. 1: 767. 1784. TYPE:
Aloysia citrodora Paláu.
Aloysia Ort. ex Juss., Ann. Mus. Hist. Nat. 7: 73. 1806,
nom. illeg. Lippia L. sect. Aloysia (Ort.) Schauer in
DC., Prod. [de Candolle] 11: 572. 1847, nom. illeg.
Lippia subg. Aloysia (Ort.) Schauer in Engl. & Prantl,
Nat. Pflanzenfam. Teil 4(Abt. 3a): 151. 1897, nom.
illeg. TYPE: Verbena triphylla L’Hér. [[ A. triphylla
(L’Hér.) Britton; ¼ A. citrodora Paláu].
Verbena L. sect. Aloysioides Walp., Repert. Bot. Syst. 4: 13.
1845. TYPE: Verbena gratissima Gillies & Hook. [[
Aloysia gratissima (Gillies & Hook.) Tronc.], lectotype, designated here.
570
Annals of the
Missouri Botanical Garden
Zapania Lam., Tabl. Encycl. 1: 59. 1791, nom. illeg. orth.,
Zappania Scop., 1786.
Xeroaloysia Tronc., Darwiniana 12: 50. 1960. syn. nov.
TYPE: Xeroaloysia ovatifolia (Moldenke) Troncoso [¼
Aloysia ovatifolia Moldenke, Lilloa 5: 379. 1940].
faces not connate, exceptionally orbicular in cross
section, commissural faces connate.
Plants suffruticose or shrubby, mostly aromatic;
stems 4-angled when young, rounded with age,
branches sometimes spiny, pubescent or glabrate.
Leaves simple, mostly opposite or ternate, 3(4)whorled, occasionally alternate or clustered into
fascicles of 4 to 8 leaves; blades entire or 3- to 5lobed, exceptionally 3-parted, linear, elliptic, oblong,
obovate, ovate, orbicular, rhomboidal, or cordate,
sessile, subsessile, or briefly petiolate, basally
attenuate, acute, round, truncate, or cordate, apically
obtuse, subobtuse, acute, acuminate, or round; blade
margins entire, crenate, serrate, dentate, or lobed,
sometimes revolute or subrevolute; adaxially glabrous
or scabrous, hispidulous, or strigose; abaxially
strigose, incanous, hirsute or hispid, often with
glandular trichomes, membranaceous, coriaceous, or
somewhat thickened texture. Inflorescences in spicate racemes, occasionally paniculate; florescences
globose, or filiform in anthesis, elongated or not in
fructification, sessile or subsessile, in monobotrya or
grouped as pleiobotrya in terminal and axillary
(heterothetic) or only axillary (homothetic) positions;
flowers often in clusters of 3 to 6; sessile to briefly
pedicellate, subtended by floral bracts; bracts inferior
to flowers, linear, elliptic, obovate, or ovate; abaxially
strigose to setose; apically acute to acuminate. Flower
with the calyx 4-toothed, exceptionally bilobed,
subactinomorphic to zygomorphic, externally strigose,
setose, hirsute, or velutinous, often subsessile
glandular, internally glabrous; fully acrescent in fruit
and persistently enclosing mature schizocarp; corollas subactinomorphic to zygomorphic, infundibuliform, white, lavender, purple, pink, or blue; tube
cylindrical to gibbous, glabrous to variously pubescent externally, internally villous along distal half;
limb 4-lobed, superior lobe often cleft; styles filiform
and usually glabrous, occasionally villous along base;
stigmas capitate to subcapitate, bilobed, stigmatic
lobes approximately equal or oblique, apically or
laterally disposed; stamens 4, epipetalous, subequal
to didynamous, the superior pair sometimes weakly
exerted, exceptionally with glandular anther connective appendices; thecae longitudinally dehiscent.
Fruit a dry schizocarp, exceptionally drupaceous;
the dry schizocarp composed of 2 cluses (each cluse
is a unit representing half a carpel, derived from a
unicarpellate ovary splitting into 2), ellipsoid to
obovoid, often 6 cordate, basally truncate, apically
rounded to bilobed, glabrous or setose pubescence,
typically elliptic in cross section and commissural
Distribution. Aloysia is an American genus,
occurring from southwestern United States and
Mexico to Chile and central Argentina. It is most
diverse in South America, with 28 species and six
varieties occurring there (Appendix 1). One taxon, A.
gratissima var. gratissima (Gillies & Hook.) Tronc., is
found in both North America (Texas, Arizona) and
South America, with a disjunction in distribution
across the tropics. Only three Aloysia species are
found exclusively in North America, A. coalcomana
Siedo, A. macrostachya (Torr.) Moldenke, and A.
wrightii A. Heller, and are not further considered
here.
Notes. Walpers (1845) established Verbena sect.
Aloysioides, mentioning the two species V. salviifolia
Hook. & Arn. and V. gratissima Gillies & Hook. The
latter name is selected as lectotype (McNeill et al., 2012,
Art. 10.2) because it is a species that certainly belongs
within Aloysia and is representative of the genus.
The fruit in Aloysia is a schizocarp that derives
from a unicarpellate ovary that normally (except in A.
ovatifolia Moldenke, which has an undivided drupaceous fruit) separates into two units along the medial
plane of the ovary; each unit represents half a carpel
and is called a cluse (O’Leary et al., 2012).
The term florescence is used to define a minimum
expression of flower arrangement. In the Verbenaceae, florescences are either racemes or spikes that
vary in the arrangement, spacing, number of flowers,
and development of its rachis. Florescences are
organized into simple inflorescences, as monobotrya,
or compound inflorescences, as pleiobotrya. In
pleiobotrya, florescences may be either terminal or
axillary, as heterothetic pleiobotrya, or grouped only
as axillary florescences, as homothetic pleiobotrya
(Múlgura et al., 2002; O’Leary et al., 2012). Classical
views of inflorescence evolution (Troll, 1964–1969;
Sell, 1976; 1980) suggest that heterothetic pleiobotrya gave rise, on the one hand, to homothetic
pleiobotrya by loss of the terminal florescence, and,
on the other hand, to monobotrya, by loss of axillary
florescences (O’Leary et al., 2012). Homothetic
pleiobotrya and monobotrya can therefore be considered derived conditions. This analysis also confirms
the derivation of monobotrya from heterothetic
pleiobotrya by loss of axillary florescences. However,
unexpectedly, according to classical assumptions
about the evolution of inflorescences, reversions from
monobotrya to heterothetic pleiobotrya also occur.
Within Aloysia, the most frequent inflorescence
morphology in South America is a homothetic
Volume 101, Number 3
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O’Leary et al.
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Revision of Aloysia (Verbenaceae) in South
America
pleiobotrya, found in 19 out of the 28 taxa. There are
only five species with heterothetic pleiobotrya: A.
arequipensis Siedo (cf. Fig. 1), A. citrodora, A. fiebrigii
(Hayek) Moldenke, A. herrerae Moldenke (Fig. 7),
and A. velutina Siedo (Fig. 13). Monobotrya are found
in four out of the five Acantholippia species recently
transferred to Aloysia (Lu-Irving et al., 2014): A.
deserticola (Phil.) Lu-Irving & N. O’Leary, A. riojana
(Hieron. ex Moldenke) Lu-Irving & N. O’Leary, A.
salsoloides (Griseb.) Lu-Irving & N. O’Leary, and A.
tarapacana (Botta) Lu-Irving & N. O’Leary.
The base chromosome number in Aloysia seems to
be x ¼ 9, based on a specimen of A. gratissima,
although this is the lowest observed count in the
genus (Powell et al., 2010). This is in agreement with
other Lantaneae, and both Lippia and Phyla would
also appear to have x ¼ 9 (Sanders, 1987; Munir,
1993). In contrast, Lantana is reported to have a base
chromosome number of x ¼ 11 or 12 (Sanders, 1987).
TAXONOMIC KEY
AMERICA
1.
10.
2(1).
20.
3(2 0 ).
30.
4(3 0 ).
40.
5(1 0 ).
50.
6(5 0 ).
60.
7(6 0 ).
70.
8(7).
80.
TO
SPECIES
OF
ALOYSIA
FROM
SOUTH
Plants with spiny branches and reduced leaf
blades, always shorter than 0.5 cm long . . . . . . 2
Plants with no spiny branches and developed leaf blades, always longer than 0.5 cm
long . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5
Leaves alternate, not squamiform, nor imbricate, blades 5-lobed . . . . . . . . . . . . . . . . . . .
. . . . . . . . . 22. A. salsoloides (Griseb.) Lu-Irving &
N. O’Leary
Leaves opposite, squamiform, densely imbricate, blades entire or 3-lobed . . . . . . . . . . . . . 3
Leaf blades entire and no evident furrows on
abaxial surface, dark green colored, endemic
to Chile . . . . . . . . . . . . . . 25. A. tarapacana (Botta)
Lu-Irving & N. O’Leary
Leaf blades 3-lobed, with a conspicuous
furrow on each blade lobe, light green or
yellow-green colored, Argentina and Chile . . . . 4
Fruits typical of the genus with cluses (each
cluse is a unit representing half a carpel,
derived from a unicarpellate ovary splitting
into two) of elliptic cross section and commissural faces not connate . . . . . . . . . . 9. A. deserticola
(Phil.) Lu-Irving & N. O’Leary
Fruits with cluses of orbicular cross section
and connate commissural faces . . . 21. A. riojana
(Hieron. ex Moldenke) Lu-Irving & N. O’Leary
Fruits undivided and drupaceous . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . 16. A. ovatifolia Moldenke
Fruits schizocarpic and divided into two dry
cluses . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6
Leaves alternate . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . 19. A. polystachya (Griseb.) Moldenke
Leaves opposite or verticillate (ternate) . . . . . . . 7
Leaves mostly verticillate (ternate), sometimes opposite at some nodes . . . . . . . . . . . . . . . . 8
Leaves mostly opposite, sometimes verticillate at some nodes . . . . . . . . . . . . . . . . . . . . . . . . . 13
Florescences only axillary . . . . . . . . . . . . . . . . . . . . 9
Florescences axillary and terminal . . . . . . . . . . 12
9(8).
Leaves evenly crenate toward apex . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . 8. A. crenata Moldenke
90.
Leaves with entire margins . . . . . . . . . . . . . . . . . . 10
10(9 0 ). Leaves adpressed to the stem, sessile, with
ovate, elliptic, or cordate blades . . . . . . . . . . . . . 11
10 0 .
Leaves not adpressed to the stem, with
sessile to subpetiolate, elliptic blades . . . . .
. . . . . . . . . . . . . . . . . . . . 2. A. brasiliensis Moldenke
11(10). Leaf blades ovate to elliptic . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . 18. A. polygalifolia Cham.
11 0 .
Leaf blades cordate . . . . . . . . . 7. A. cordata Siedo
12(8 0 ). Leaf blades smaller, 1.8–3 3 0.2–0.5 cm,
with entire margins . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . 11. A. fiebrigii (Hayek) Moldenke
12 0 .
Leaf blades larger, 2–8 3 1–2.5 cm, with
entire or slightly serrate margins . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . 6. A. citrodora Paláu
13(7 0 ). Florescences terminal and axillary; plants
endemic to Peru and Bolivia . . . . . . . . . . . . . . . . 14
13 0 .
Florescences only axillary . . . . . . . . . . . . . . . . . . . 16
14(13). Leaf margins entire, with minute scabrous
pubescence on abaxial blade surface, glabrous adaxially . . . . . . . 14. A. herrerae Moldenke
Leaf margins not entire, with velutinous,
14 0 .
incanous, strigose, or tomentose pubescence,
never glabrous . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
15(14 0 ). Margins slightly crenate along entire blade,
adaxially velutinous, abaxially incanous . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . 27. A. velutina Siedo
15 0 .
Margins finely serrate along apical 2/3 to 1/2
of blade length, adaxially strigose, abaxially
tomentose . . . . . . . . . . . . . 1. A. arequipensis Siedo
16(13 0 ). Flowers with a bilobulated calyx . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . 10. A. dusenii Moldenke
16 0 .
Flowers with a 4-toothed calyx . . . . . . . . . . . . . . 17
0
17(16 ). Superior pair of stamens with anther connective appendices glandular . . . . . . . . . . . . .
. . . . . 26. A. trifida (Gay) Lu-Irving & N. O’Leary
17 0 .
Superior pair of stamens with anther connective appendices not glandular . . . . . . . . . . . . 18
18(17 0 ). Leaf margins completely entire, sometimes
slightly serrate toward apex . . . . . . . . . . . . . . . . . 19
18 0 .
Leaf margins not entire, ranging from serrate
or crenate to dentate, or basally entire with
some serration, crenation, or dentition in
some portion of the blade; if margins entire,
then only on leaves on some part of the
plant, the remaining leaves not entire . . . . . . . 22
19(18). Leaves smaller, narrowly elliptic, less than 2
3 0.3 cm . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . 12. A. gratissima (Gillies & Hook.) Tronc.
19 0 .
Leaves larger, elliptic, ovate, cordate, or
obovate, more than 2 3 0.3 cm . . . . . . . . . . . . . 20
20(19 0 ). Leaf adaxial surfaces without a conspicuous
midvein, leaves briefly petiolate, with blades
elliptic, obovate, or ovate . . . . . . . . . . . . . . . . . . . 21
20 0 .
Leaf adaxial surfaces with conspicuous midvein, leaves sessile, with blades obovate to
oblanceolate . . . . . . 15. A. oblanceolata Moldenke
21(20). Leaf blades elliptic to obovate, with obtuse
apex . . . . . . . . . . . 20. A. pulchra (Briq.) Moldenke
Leaf blades elliptic to ovate, with acute to
21 0 .
subobtuse apex . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . 12. A. gratissima (Gillies & Hook. ex
Hook.) Tronc.
22(18 0 ). Leaf margins almost entirely dentate, crenate, or serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 23
572
Annals of the
Missouri Botanical Garden
Figure 1. Aloysia arequipensis Siedo. —A. Floriferous branch with florescences in both axillary and terminal positions
(heterothetic pleiobotrya). —B. Lateral view of intact flower with calyx and floral bract. —C. Intact leaf, adaxial surface. —D.
Detail of tomentose pubescence on abaxial blade surface. —E. Detail of strigose pubescence on adaxial blade surface. A–E,
illustrated from Pennell 13079 (type, US).
22 0 .
Leaf margins with some serration or dentition in some portion of the blade, but always
entire basally . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 31
23(22). Leaf margins with 3 to 5 deep teeth coarsely
dentate or serrate on each side . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . 5. A. chamaedryfolia Cham.
Leaf margins evenly dentate, crenate, or
23 0 .
serrate . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
24(23 0 ). Leaf margins minutely serrate or crenate;
corolla less than 4 mm long; calyx with long
teeth, equaling or exceeding calyx tube in
length . . . . . . . 28. A. virgata (Ruiz & Pav.) Pers.
0
24 .
Leaf margins notoriously crenate or dentate;
corollas more than 4 mm long; calyx teeth
never exceeding calyx tube in length . . . . . . . . 25
25(24 0 ). Florescences dense, subrhomboidal in anthesis, cylindrical in fructification, 2–4 cm long,
1 or 2(3) per leaf axil or grouped toward plant
apex . . . . . . . . . . . . . . 4. A. catamarcensis Moldenke
Florescences dense or lax, cylindrical, 1–12
25 0 .
cm long, 1 per leaf axil . . . . . . . . . . . . . . . . . . . . . 26
26(25 0 ). Leaf blades oblong, 1–4.5 3 0.15–1 cm . .
. . . . . . . . . . . . . . . . . . . 3. A. castellanosii Moldenke
26 0 .
Leaf blades elliptic, ovate, or orbicular, 1–
5( 7) 3 1–3( 4) cm . . . . . . . . . . . . . . . . . . . . . . . . 27
27(25 0 ). Leaves membranaceous and plane; corolla
tube more than 6 mm long . . . . . . . . . . . . . . .
. . . . . . . . . . . . 17. A. peruviana (Turcz.) Moldenke
Leaves rugose and creased; corolla tube less
27 0 .
than 6 mm long; if leaves membranaceous
then corolla tube less than 4.5 mm long . .
. . . . . . . . . . . 24. A. scorodonioides (Kunth) Cham.
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2016
28(22). Floral bracts 4–4.5 mm long, surpassing
calyx, endemic to Chile . . . . . . . . . . . . . . . . . .
. . . . . 23. A. salviifolia (Hook. & Arn.) Moldenke
28 0 .
Floral bracts 1–2 mm long, not surpassing
calyx . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
29(28 0 ). Leaf blades elliptic, with slightly serrate
margins in apical half, venation conspicuous, reddish brown, pinnate, impressed on
abaxial surface, endemic to Brazil (Paraná)
. . . . . . . . . . . . . . . . . . 13. A. hatschbachii Moldenke
29 0 .
Leaf blades elliptic or ovate, with entire or
serrate margins, midvein impressed only on
abaxial surface, or if venation pinnate and
conspicuous then not reddish brown . . . . . .
. . . . . . 12. A. gratissima (Gillies & Hook.) Tronc.
1. Aloysia arequipensis Siedo, Lundellia 15: 38, fig.
1. 2012. TYPE: Peru. Arequipa: Mpio. Arequipa, Tiabaya, open, rocky slope, 8 Apr. 1925, F.
W. Pennell 13079 (holotype, NY [barcode]
NY01911741 not seen, NY image!; isotypes, F
[bc] F0093716F not seen, F image!, GH [bc]
GH00359317 not seen, GH image!, US!). Figure
1.
Shrubs 0.5–1.5 m tall; stems glabrous. Leaves
opposite, rarely ternate; petioles brief, 1–2 mm;
blades elliptic, 1–2 3 0.5–1.2 cm, apex acute to
obtuse, base acute, margins finely serrate along
apical 2/3 to 1/2 of blade length, adaxially strigose,
abaxially tomentose, with an understory of subsessile,
glandular trichomes. Florescences terminal and
axillary (heterothetic pleiobotrya), cylindrical, loosely
spicate, 1–5 cm; peduncles 0.5–2.8 cm; flowers
lavender to pink, often with a whitish limb, floral
bracts elliptic, 1–3 3 0.5–1 mm, apex acuminate,
with strigose pubescence and an understory of
subsessile, glandular trichomes. Flower with the
calyx 2–4 mm, densely hirsute, with an understory
of subsessile, glandular trichomes with 4 brief teeth,
unequal, triangular; corolla tube 4–7 mm, externally
finely pulverulent, with villous fauce. Cluse 1–1.5 3
1 mm, glabrous.
Distribution and habitat. Aloysia arequipensis is
endemic to Peru, in arid sites from Arequipa and
southern Lima Provinces, at elevations of 2000–3000
m.
Discussion. Aloysia arequipensis is one of the five
South American taxa distinguished by its heterothetic
pleiobotrya, also present in A. citrodora, A. fiebrigii,
A. herrerae, and A. velutina. However, A. arequipensis
may be differentiated from A. citrodora and A.
fiebrigii by its opposite leaves, being ternate in these
last two species. It is distinguished from A. herrerae
by its finely serrate leaf blade margins, with blade
margins entire in this last taxon. Aloysia arequipensis
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Revision of Aloysia (Verbenaceae) in South
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may be differentiated from A. velutina because in this
last species leaf blade margins are crenate. Aloysia
arequipensis is also morphologically similar to A.
scorodonioides (Kunth) Moldenke var. hypoleuca
(Briq.) Moldenke from which it is readily distinguished by inflorescence morphology, having only
axillary florescences present (homothetic pleiobotrya)
in this last taxon.
Selected specimens examined. PERU. Arequipa: Arequipa, cerros de Jesús, Vargas 12671 (US). Lima: Mpio.
Yauyos, Aiza, entre Catahuas y Tupe, Pradera, E. Cerrate
1282 (MO).
2. Aloysia brasiliensis Moldenke, Phytologia 3: 162.
1949. TYPE: Brazil. Paraná, 4 Jan. 1904, P. K.
Dusén s.n. (holotype, R [barcode] R000046798
not seen, R image!; isotypes, NY [bc]
NY00103867 not seen, NY image!, SI [bc]
SI003386!). Figure 2.
Shrubs 1–3 m tall, stems glabrous. Leaves ternate,
not adpressed to the stem, sclerophyllous, sessile to
subpetiolate, petiole 0.2–1.5 mm, blades narrowly
elliptic to elliptic, 1–4( 5) 3 0.5–1.5 cm, apex acute
or sometimes subobtuse, base acute, margins entire,
sometimes subrevolute, scabrous to strigose pubescence on both surfaces. Florescences axillary,
solitary, lax, 4–10( 16) cm; peduncles 1–5 cm;
flowers lilac; pedicels 0.5–1 mm; floral bracts linear,
2–3.5 mm, apex acuminate, with strigose pubescence. Flower with the calyx 2–4 mm, hispid, 4toothed, the teeth unequal, triangular; corolla tube 4–
5.5 mm, externally finely pulverulent, internally, with
villous fauce. Cluse 1.5 3 1 mm, glabrous.
Distribution and habitat. Aloysia brasiliensis is
endemic to southern Brazil, from the states of Rio
Grande do Sul, Paraná, and Santa Catarina. The
species has been collected from disturbed habitats, at
elevations from 700 to 900 m.
Discussion. Aloysia brasiliensis, A. cordata, and
A. polygalifolia all have glabrous stems but puberulent rachises, and sclerophyllous leaves with entire
margins, and all three are endemic to Brazil. Aloysia
brasiliensis differs because leaves are not adpressed
to the stem, with elliptic blades, versus leaves
adpressed to the stem, with ovate, elliptic, or cordate
blades in the other two species. In addition, A.
brasiliensis has larger leaves (1–4[ 5] 3 0.5–1.5 cm)
with an acute base, versus smaller and a different
basal shape in the other two species. Leaf blades in
A. polygalifolia (0.5–2 3 0.3–1 cm) have a truncate
base; blades in A. cordata (0.3–1 3 0.3–0.9 cm) have
a cordate base.
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Figure 2. Aloysia brasiliensis Moldenke. —A. Floriferous branch, showing the axillary florescences (homothetic pleiobotrya).
—B. Intact flower in lateral view, with floral bract subtending the hispid calyx. —C. Corolla dissected open to reveal the
androecium. —D. Intact leaf, adaxial surface. —E. Detail of strigose pubescence on blade adaxial surface. —F. Detail of
strigose pubescence on abaxial blade surface. A, taken from Klein 3489 (SI); B–F, from Hatschbach 14905 (SI).
Selected specimens examined. BRAZIL. Paraná: Uniao
da Vitoria, Sao Domingos, Silva 7037 (SI); Uniao da Vitoria,
Hatschbach 14905 (SI). Rio Grande do Sul: Nonoai, ad fl.
Uruguay, Rambo 28141 (NY).
¸ Santa Catarina: Irani, Smith
& Klein 13029 (NY); CaCador, faz. dos Carneiros, Klein
3489 (SI), Smith & Reitz 9012 (SI).
3. Aloysia castellanosii Moldenke, Lilloa 5: 372.
1940. TYPE: Argentina. San Juan: Quebrada
del Zonda, 28 Feb. 1926, A. Castellanos s.n.
(holotype, BA-26/602!; isotypes, CORD [barcode] CORD00003815 not seen, CORD image!,
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NY [bc] NY00103870 not seen, NY image!, SI
[bc] SI003376!).
Aloysia castellanosii var. magna Moldenke, Known Geogr.
Distrib. Verb. Avicenn., 76. 1942. TYPE: Argentina.
Tucumán: Valle de Amaicha, Zurita, 4 Feb. 1917, L.
Castillón 85 (holotype, NY [barcode] NY00103871
not seen, NY image!; isotypes, LIL [bc] LIL001434
not seen, LIL image!, LP [bc] LP006684 not seen, LP
image!, SI [bc] SI003387!).
Aloysia decorticans Ravenna, Onira 11(2): 3( 4). 2007.
TYPE: Argentina. Salta: Quebrada de Guachipas, 23
Jan. 1943, Castellanos s.n. (holotype, BA-46961!).
Plants suffruticose, 1 m tall; stems cylindrical,
pilose to subglabrous, internodes 1–4.5 cm, smooth,
red, with fallen bark. Leaves opposite; petioles brief,
0.8–4 mm; blades oblong, 1–4.5 3 0.15–1 cm, apex
obtuse, base truncate, margins prominently and
evenly crenate, blade margins subrevolute, adaxially
scabrous, creased, abaxially densely hispid. Florescences axillary, solitary, dense, cylindrical, 1–6 cm
in fructification; peduncles 0.5–2.8 cm; flowers
white, pink, or violet; floral bracts ovate, obovate, or
elliptic, 2.5–10.5 3 1.5–6 mm, sticky glandular on
both bract surfaces. Flower with the calyx 2.5–5 mm,
densely hirsute, with 4 brief teeth, unequal, triangular; corolla tube 4–7 mm, externally glabrous, with
villous fauce. Cluse 2 3 0.5 mm, glabrous.
Iconography. Botta (1979: 73, fig. 1).
Distribution and habitat. Aloysia castellanosii is
endemic to Argentina, known only from the provinces
of Catamarca, La Rioja, Salta, San Juan, and
Tucumán. The species is found in the phytogeo˜ (Cabrera &
graphic provinces of monte and chaquena
Willink, 1973), growing at elevations of 2500–2800
m.
Discussion. Aloysia castellanosii is distinguished
by its leaf morphology, with oblong blades and
margins prominently and evenly crenate, and creased
texture. This feature makes it hard to confuse with
any other species of the genus.
Selected specimens examined. ARGENTINA. Catamarca: Andalgalá: 16 km SE from Andalgalá, Cantino 693 (SI).
La Rioja: General Lavalle, Parque Nac. Talampaya, Botta
695 (SI). Salta: San Carlos, ruta 40 de Cafayate a Cachi,
sierra de Quilmes, Cialdella 218 (SI). San Juan: Caucete,
Marayes, Haene 93 (SI). Tucumán: Tafı́, Amaicha del
Valle, Burkart 22066 (SI).
4. Aloysia catamarcensis Moldenke, Known Geogr.
Distrib. Verb. Avicenn. 76. 1942. TYPE:
Argentina. Catamarca: Quebrada del Tala, 15
Mar. 1909, L. Castillón 956 (holotype, NY
[barcode] NY00103872 not seen, NY image!;
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isotypes, CORD [bc] CORD00003817 not seen,
CORD image!, GH [bc] GH00299000 not seen,
GH image!, LIL [bc] LIL001435 not seen, LIL
image!, LIL [bc] LIL001436 not seen, LIL
image!, SI [bc] SI003377!).
Plants suffruticose, 1–1.5 m tall, stems cylindrical,
pilose to glabrous, with fallen bark, internodes 3–7 cm.
Leaves opposite, sometimes ternate; petioles 0.3–1.5
mm; blades elliptic, 1.5–5 3 1–3 cm, apex acute to
subobtuse, base acute, margins prominently and
evenly crenate to dentate, scabrous, adaxially rugose,
abaxially hispid. Florescences axillary, 1 or 2(3) per
leaf axil, grouped toward stem apex, dense, subrhomboidal at anthesis, cylindrical in fructification, 2–4 cm,
pedunculate; flowers white, pink, or violet; floral bracts
elliptic to obovate, apex attenuate, acuminate, 2.5–3.5
3 1–1.5 mm, surpassing calyx, puberulous. Flower
with the calyx short, 1–2 mm, hispid, with 4 brief
teeth, unequal, triangular; corolla tube 4–7.5 mm, with
glabrous fauce. Cluse 1 3 0.5 mm, glabrous.
Iconography. Botta (1979: 79, fig. 3).
Distribution and habitat. Aloysia catamarcensis
is endemic to northwestern Argentina, being found in
the provinces of Santiago del Estero, La Rioja,
Catamarca, and Salta. The species has been observed
to grow on slopes of small hills or rises.
Discussion. Aloysia catamarcensis is distinguished by its elliptic leaf blades with margins
prominently and evenly crenate to dentate. It can be
confused with A. scorodonioides var. scorodonioides or
A. peruviana. However, these two last taxa have
florescences cylindrical in anthesis, 4.5–10 cm long,
one per leaf axil, in contrast to the shorter
florescences of A. catamarcensis, subrhomboidal in
anthesis, cylindrical in fructification, 2–4 cm long,
one or two (three) per leaf axil or grouped toward the
apex of the plant in A. catamarcensis.
Selected specimens examined. ARGENTINA. Catamarca: Belén, cerrito de la Cruz, Ulibarri 332 (SI); Capital,
Quebrada de los Nacimientos, Ulibarri 950 (SI). La Rioja:
Chilecito, Miranda, rt. 40 a 16 km de Chilecito, Olmstead
˜ Km. 77, camino de Cafayate a
2007-82 (SI). Salta: La Vina,
Salta, Correa 4318 (SI). Santiago del Estero: Alberdi, entre
Donaden y Campo Gallo, s. coll. 43 (SI).
5. Aloysia chamaedryfolia Cham., Linnaea 7: 234.
1832. Lippia chamaedryfolia (Cham.) Steud.,
Nomencl. Bot. [Steudel], ed. 2 1: 62. 1841.
TYPE: ‘‘Brasilia,’’ s.d., F. Sellow s.n. (lectotype,
designated by Siedo [2010: 200], W-0032437!;
isolectotypes, BM-643743 not seen, BM image!,
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G [barcode] G00208776 not seen, G image!, WRchb. 1889-0290180 not seen, W image!).
Shrubs erect, 0.5–2 m tall; stems tetragonous,
densely pubescent with retrorse hairs. Leaves
opposite, sometimes ternate, subsessile; blades ovate
to suborbicular 0.3–3 3 0.7–2 cm, apex subobtuse,
base subtruncate, margin slightly revolute, deeply
toothed, with 3 to 5 deep teeth coarsely dentate or
serrate on each side, adaxially scabrous, abaxially
densely strigose, venation conspicuous. Florescences
axillary, solitary, lax, 5–15 cm; peduncles 2–4 cm;
flowers lilac or purple; floral bracts ovate, apex
subulate, 1.5 mm, hispid. Flower with the calyx 2.5
mm, hispid in inferior half, slightly hispid toward
apex, calyx with 4 brief teeth, unequal, triangular;
corolla tube 3–4 mm, with villous fauce. Cluse 1 3
0.5 mm, pubescent at apex.
Iconography. Botta (1979: 91, fig. 7).
Distribution and habitat. Aloysia chamaedryfolia
grows in southern Brazil, Uruguay, and northeastern
Argentina where it is found in rocky soils.
Discussion. Aloysia chamaedryfolia is distinguished by its leaf blades ovate to suborbicular with
margins with three to five deep teeth on each side,
which makes this species not easily mistaken for any
other member of the genus.
Selected specimens examined. ARGENTINA. Misiones:
San Javier, Balneario 4 Bocas, 11 km SE de San Javier,
Krapovickas 28868 (CTES, SI). BRAZIL. Rio Grande do
Sul: Arroio dos Ratos, Hagelund 10590 (SI). URUGUAY.
Artigas: Arroyo Sepulturas, Bonifacino et al. 1953 (SI).
Rivera: Herter 158 (NY). Tacuarembó: Valle Edén,
Rosengurtt 4967 (SI).
6. Aloysia citrodora Paláu, Parte Prácte Bot. 1:
768. 1784, as ‘‘citriodora.’’ Verbena citriodora
(Paláu) Cav., Descr. Pl. (Cavanilles) 68. 1802.
Aloysia citriodora Ort. ex Pers., Syn. Pl. 2(1):
139. 1806, nom. superfl. illeg. [cf. Art. 52.1;
cites Verbena triphylla L’Hér.]. TYPE: unnumbered illustration in Paláu, Part. Prácte. Bot. 1:
768. 1784 (lectotype, designated by Armada &
Barra [1992: 89], icon s.n. in Paláu [1784:
768]).
Verbena triphylla L’Hér., Stirp. Nov. 1: 21. 1785. Zapania
citrodora Lam., Tab. Encycl. 1: 59. 1791, nom. illeg.
[Art. 52.1, cites Verbena triphylla L’Hér., the epithet
that ought to have been used]. Lippia citrodora Kunth,
Nov. Gen. Sp. 2: 269. 1818, as ‘‘citriodora,’’ nom.
illeg. [cf. Art. 52.1, comb. based on ‘‘Zapania
citrodora’’ Lam.]. Lippia triphylla (L’Hér.) Kuntze,
Revis. Gen. Pl. 3(3): 253. 1898. Aloysia triphylla
(L’Hér.) Britton, Sci. Surv. Porto Rico & Virgin
Islands 6: 140. 1925, nom. superfl. illeg., non Aloysia
triphylla Royle, Ill. Bot. Himal. Mts., 299. 1833.
TYPE: France. Ile de France, cultivated plant in
Jardin de Plantes, Paris, s.d., C. L. L’Héritier s.n.
(lectotype, designated by Moldenke & Moldenke
[1983: 232], P not seen, P image!).
Aloysia sleumeri Moldenke, Phytologia 10: 170. 1964.
TYPE: Argentina. Catamarca: Belén, Pozo de Piedra,
25–31 ene. 1952, H. Sleumer & F. Vervoost 2370
(holotype, US [barcode] US00118880 not seen, US
image!; isotypes, TEX-LL [barcode] LL00374941 not
seen, TEX-LL image!, SI [bc] SI003401!).
Aloysia triphylla (L’Hér.) Britton f. serrulata Moldenke,
Phytologia 50: 308–309. 1982. TYPE: [cultivated]
U.S.A. Indiana: Floyd Co., New Albany, 3 Oct. 1893,
L. H. Bailey 160 (holotype, BH not seen).
Shrubs 1–3 m tall, aromatic, stems glabrous at
maturity, subpendulous. Leaves ternate, briefly
petiolate, petioles 1–5 mm; blades elliptic, 2–8 3
1–2.5 cm, apex acute, base acute, margins entire or
slightly serrate, blade adaxially scabrous, abaxially
glabrate with subsessile glandular trichomes, midvein
and pinnate venation conspicuous. Florescences
terminal and axillary (heterothetic pleiobotrya), lax,
1–5 cm, the terminal ones grouped as paniculiform
inflorescences; flowers white, small; floral bracts
reduced, ovate, 1–1.5 mm, scabrous. Flower with the
calyx 2.5–3 mm, puberulous, with 4 brief teeth,
unequal, triangular; corolla tube 5–6 mm, externally
puberulous, with villous fauce. Cluse 2 3 1 mm,
glabrous or pubescent at apex.
Iconography. Botta (1979: 104, fig. 12).
Distribution and habitat. Aloysia citrodora is
native to the dry areas of northwestern Argentina,
including the provinces of Catamarca, Jujuy, La
Rioja, Salta, San Juan, and Tucumán, to southern
Bolivia and Paraguay. The species is cultivated all
around the world.
Discussion. Aloysia citrodora has strongly lemonscented leaves, which easily distinguishes it from
other species, and is often cultivated for the
preparation of aromatic teas with medicinal properties. For this reason the taxon is widely cultivated in
Chile, Uruguay, southern Brazil, and Paraguay
(Arambarri et al., 2009: 18). It can be confused with
A. friebrigii, which also has ternate leaves. However,
in this last species, blades are smaller, 1.8–3 3 0.2–
0.5 cm, and the blade margin is always entire, in
contrast to the blades 2–8 3 1–2.5 cm in A. citrodora,
which are also entire but sometimes with a slightly
serrate margin.
Moldenke (1982: 309) noted that the type for
Aloysia triphylla f. serrulata was collected ‘‘from the
garden of Cornell Experiment Station. . .from material
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Figure 3. Aloysia cordata Siedo. —A. Floriferous branches, showing the axillary florescences (homothetic pleiobotrya). —B.
Detail of stem with leaf scars and antrorse pubescence. —C. Lateral view of intact flower, with hispid floral bract. —D. Cordate
shape of leaf adaxial surface. —E. Detail of leaf adaxial surface, glabrous with minutely scabrous margins. —F. Detail of leaf
abaxial surface, glabrous with minutely scabrous margins. A–F, taken from Hatschbach 20792 (isotype, SI).
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secured from Ernest Walker of New Albany,
Indiana.’’ Even though described from North America, this was a cultivated form of A. citrodora native to
Argentina.
Cavanilles spelled the epithet citriodora with a
second ‘‘i.’’ This spelling is correctable (McNeill et
al., 2012, Art. 60.8) by omitting the second ‘‘i’’ for
citrodora.
species occur in Brazil, while A. cordata is restricted
to eastern Paraná. Aloysia polygalifolia is more
broadly distributed, from the states of Paraná, Rio
Grande do Sul, and Santa Catarina. Aloysia brasiliensis also occurs in Paraná, in sympatry with these
two species; however, it is easily distinguished by its
leaves not adpressed to the stem, with elliptic blades
(see discussion under A. brasiliensis).
Selected specimens examined. ARGENTINA. Catamarca: Belén, Pozo de Piedra, Troncoso 1897 (NY, SI). Jujuy:
Tumbaya, Chicayo, finca del Sr. Gronda subiendo cerro,
Zuloaga 10188 (SI); Volcán Chicayo, Cabrera 16877 (SI).
La Rioja: Famatina, La Aguadita, Cabrera 24637 (SI).
Salta: Rosario de Lerma, rt. 51 de Salta a San Antonio de
los Cobres, Olmstead 2007-13 (SI). San Juan: Valle Fértil,
de Sa. De Elizondo a Sa. De Chavez, Kiesling 6620 (SI).
Tucumán: Juan Bautista Alberdi, El Chorro, cumbres
Calchaquies, D. Rodrı́guez 1214 (NY, SI). BOLIVIA.
Cochabamba: Carrasco, E. Martinez 479 (NY). La Paz:
Murillo, Calacoto, Solomon 15755 (SI). Tarija: Mendez, rio
Pilaya, Camaron, Gerold 161 (SI). PARAGUAY. Cordillera:
Schinini 6767 (NY). Dpto. Central: Itá, Arenas 1918
(BACP).
Selected specimens examined. BRAZIL. Paraná: San
˜ Hatschbach 20792 (SI);
José dos Pinhais, rio Pequeno,
Piraquara, Medianeira, Costa & Barbosa 6 (SI).
7. Aloysia cordata Siedo, Lundellia 15: 42, fig. 3.
˜ José dos
2012. TYPE: Brazil. Paraná: Mpio. Sao
´ Pequeno, do brejo, 17 Jan. 1969,
Pinhaes; Rıo
G. Hatschbach 20792 (holotype, NY [barcode]
NY01911743 not seen, NY image!; isotypes, K!,
MICH!, MO!, SI [bc] SI041007!, SI [bc]
SI041008!, UC!). Figure 3.
Shrubs 1–2 m tall, slender, few-branched. Leaves
ternate, antrorsely adpressed to the stem, internodes
highly regular in length, sessile; blades cordate,
sclerophyllous, 0.3–1 3 0.3–0.9 cm, apex mucronulate, base cordate, margins entire, minutely scaberulous, adaxially glabrous, smooth, lustrous, abaxially
glabrous, smooth, satin-lustrous. Florescences axillary, solitary, lax, 4–12 cm; peduncles 1–3 cm,
strigulose; flowers lilac; pedicels 0.5–1 mm; floral
bracts linear, 1–1.5 mm, apex acuminate, with
strigose pubescence. Flower with the calyx 1.5–2
mm, setose, glandular, 4-toothed, the teeth unequal,
triangular; corolla tube 2.5–3.5 mm, externally finely
pulverulent, internally, with villous fauce. Cluse 1.5
3 1 mm, glabrous.
8. Aloysia crenata Moldenke, Phytologia 9: 182.
1963. TYPE: Paraguay. Alto Paraná: in regione
fluminis, 1909–1910, K. Fiebrig 6137 (holotype,
US [barcode] US00118876 not seen, US image!;
isotypes, BM [bc] BM000098774 not seen, BM
image!, GH [bc] GH00282999 not seen, GH
image!, LIL [bc] LIL001364 not seen, LIL
image!, TEX-LL [bc] LL00374936 not seen,
TEX-LL image!, SI [bc] SI003382!, SI [bc]
SI003381!, W not seen).
Aloysia krapovickasii Moldenke, Phytologia 47: 330. 1981.
TYPE: Argentina. Corrientes: Ituzaingó, 24 Sep.
1974, C. L. Cristobal, J. M. Gonzalez, A. Schinini, C.
Quarin, M. M. Arbo & A. Krapovickas 26439
(holotype, TEX-LL [barcode] LL00374493 not seen,
TEX-LL image!; isotypes, CTES [bc] CTES0013814
not seen, CTES image!, SI [bc] SI003402!).
Shrubs 1–2 m tall; stems cylindrical, densely
strigose. Leaves ternate, rarely 2 or 4 per node,
subsessile, blades elliptic to obovate, 3.5–7 3 1–2.5
cm, apex acute to subobtuse, base acute, margin
entire toward base, evenly crenate toward apex,
conspicuously reticulate venation, adaxially scabrous, abaxially dense strigose with subsessile
glandular trichomes underneath. Florescences axillary, solitary, lax, 10–15 cm; peduncles 2–4 cm;
flowers white, floral bracts linear, with subulate apex,
2.5–4 mm, hispid. Flower with the calyx 4 mm,
hispid in the inferior half, slightly hispid toward apex,
with 4 brief teeth, unequal, triangular; corolla tube 7
mm, with villous fauce. Cluse 2 3 0.5 mm, pubescent
in apex.
Iconography. Botta (1979: 100, fig. 10).
Distribution and habitat. Aloysia cordata is
endemic to Brazil, from the state of Paraná, where
it is found on wet soils.
Discussion. Aloysia cordata is similar to A.
polygalifolia in having leaves adpressed to the stem
and sessile, but it is distinguished by its cordate leaf
blades (vs. ovate to elliptic in A. polygalifolia) and its
glabrate leaf surfaces (vs. scabrous to strigose). Both
Distribution and habitat. Aloysia crenata is found
from eastern Paraguay to northeastern Argentina
(Corrientes) and in Brazil (Paraná).
Discussion. Aloysia crenata is distinguished by
its ternate leaves with evenly crenate blade margins
toward the apex and the presence of only axillary
florescences. Other species that have ternate leaves
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and only axillary florescences (A. brasiliensis, A.
cordata, and A. polygalifolia) can be differentiated by
their entire leaf blade margins.
The type specimens of both species Aloysia
crenata and A. krapovickasii are from areas in close
proximity to one another, and both specimens are
nearly identical morphologically. Thus, A. krapovickasii is conspecific and considered a synonym of A.
crenata, as previously established in Múlgura et al.
(2012).
Discussion. The four species Aloysia deserticola,
A. tarapacana (Botta) Lu-Irving & N. O’Leary, A.
riojana (Hieron. ex Moldenke) Lu-Irving & N.
O’Leary, and A. salsoloides (Griseb.) Lu-Irving & N.
O’Leary were formerly recognized in Acantholippia
and have short floriferous branches that develop over
the previous year’s stems, as stated by Múlgura et al.
(2002). In addition, each of these species has erect,
imbricate leaves with inverse dorsiventral mesophyll
(Carmona & Ancibor, 1995).
Moldenke (1961) mentioned Verbena deserticola,
described by Philippi (1860), as a synonym of
Acantholippia deserticola. However, V. deserticola is
actually a synonym referred to Junellia origenes
(Phil.) N. O’Leary & P. Peralta (O’Leary et al., 2011).
Selected specimens examined. ARGENTINA. Corrientes: Ituzaingó, rte. 39 a 10 km de rte. 14, Cabrera
29106 (SI). BRAZIL. Paraná: Hatschbach 26325 (SI).
PARAGUAY. Caazapá: San Juan Neponuseno, Rojas 5903
(SI).
9. Aloysia deserticola (Phil.) Lu-Irving & N.
O’Leary, Syst. Bot. 39(2): 653. 2014. Basionym:
Lippia deserticola Phil., Anales Univ. Chile 59:
262. 1881, replacement name. Replaced synonym: Lippia microphylla Phil., Anal. Univ. Chile
27: 350. 1865, non Lippia microphylla Cham.,
Linnaea 7 : 226. 1832. Acantholippia deserticola
(Phil.) Moldenke, Lilloa 5: 370. 1940. Acantholippia punensis (Phil.) Botta, Hickenia 1(35):
195. 1979, nom. illeg. superfl. TYPE: Chile.
‘‘Frequens in parte boreali deserti Atacama,’’
s.d., R. A. Philippi s.n. (lectotype, designated by
Múlgura et al. [2012: 5] SGO-4230 not seen,
SGO-4230 image!).
Shrubs 0.4–1.5 m tall, with spiny branches; stems
hispid, but glabrous at maturity. Leaves small,
opposite, adpressed to the stem, sessile, squamiform,
imbricate; blades rhomboidal, 1.5 3 1.5–2 mm, 3lobed with a large apical and 1 lateral small lobe on
each side, somewhat thicker texture, apex subobtuse,
base rounded, margin entire, revolute, adaxially
scabrous, abaxially with a conspicuous hirsute furrow
on each blade lobe, light green or yellow-green in
color. Florescences terminal, solitary, dense, 12–15
mm; flowers lilac; floral bracts ovate or obovate, 3–
3.5 mm long, apex acute or obtuse, slightly strigose.
Flower with the calyx 3.5–4.5 mm long, hispid, with 4
brief teeth, unequal, triangular; corolla tube 4–6 mm,
with villous fauce. Cluse 2–3 3 0.5 mm, elliptical in
cross section, commissural faces not connate.
Iconography. Botta (1980: 518, fig. 1); Caro
(1982: 17, fig. 3).
Distribution and habitat. Aloysia deserticola
grows in northwestern Argentina, southern Bolivia,
and Chile, Region II. This species is found at
elevations of 2300–3500 m and is noted to occur
frequently on rocky to sandy soils.
Selected specimens examined. ARGENTINA. Catamarca: Antofagasta de la Sierra, Ulibarri et al. 679 (SI). Jujuy:
Susques, ca. de Olacapato, esquina Azul, Cabrera et al.
31792 (SI). Salta: Los Andes, camino de Pocitos a
Canchari, Ruthsatz 213 (SI). San Juan: Dpto. Iglesia,
Parque Nac. San Guillermo, E. Haene et al. 2121 (SI).
BOLIVIA. Oruro: Atahualpa, al S de Sabaya, salar de
´
Coipasa, Beck 21555 (SI). Potosı́: Nor Lıpez,
E. Garcı́a
1209 (SI). CHILE. Region II: Antofagasta, El Loa, 60 km E
ruta 5, on rte. to Solar de Aguas Calientes, Dillon et al.
6018 (SI).
10. Aloysia dusenii Moldenke, Phytologia 1: 440.
1940. TYPE: Brazil. Paraná: shrubby campo at
Tamandré, 4 Oct. 1914, P. Dusén 1050a
(lectotype, designated by Siedo [2010: 200],
S11-10477 not seen, S image!; isolectotype, NY
[barcode] NY00103874 not seen, NY image!).
Figure 4.
Aloysia ternifolia Moldenke, Phytologia 2: 309. 1947.
TYPE: Brazil. Paraná: Itaiacoca, Ponta Grossa, 17
Mar. 1904, P. K. Dusén 4228 (holotype, S11-10485
not seen, S image!; isotypes, NY [barcode]
NY00103891 not seen, NY image!, NY [bc]
NY00103892 not seen, NY image!, US [bc]
US00118881 not seen, US image!).
Aloysia ternifolia Moldenke f. oppositifolia Moldenke,
Phytologia 28: 192. 1974. TYPE: Brazil. Paraná:
Pitanga, rio Bonito, 25 Feb. 1971, G. Hatschbach
26516 (holotype, TEX-LL [barcode] LL00374942 not
seen, TEX-LL image!; isotypes, SI [bc] SI041146!, UC
[bc] UC1426950 not seen, UC image!, US [bc]
US00118882 not seen, US image!).
Shrubs 1–2 m tall; stems glabrous. Leaves mostly
opposite, sometimes 3-whorled, sessile to subpetiolate, petiole 2–4 mm; blades narrowly elliptic to
elliptic, 3–4( 5) 3 1–1.5 cm, apex acute to
subobtuse, base acute, margins entire toward base,
minutely serrate toward apex, abaxially strigose on
midvein, adaxially with adpressed strigose pubescence over entire surface. Florescences axillary,
solitary or sometimes 2 per leaf axil, lax, 3–8 cm;
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Figure 4. Aloysia dusenii Moldenke. —A. Floriferous branch, showing the axillary florescences (homothetic pleiobotrya).
—B. Intact flower in lateral view, with floral bract subtending the lightly strigose calyx. —C. Detail of bilobulated calyx. —D.
Detail of dissected bilobulated calyx, internal view. —E. Gynoecium. —F. Detail of leaf abaxial surface, strigose on midvein.
—G. Detail of leaf abaxial surface, with adpressed strigose pubescence. A–G, taken from Hatschbach 22546 (SI).
peduncles 1–3.5 cm; flowers white; pedicels 0.2–0.5
mm; floral bracts narrowly elliptic, 2–2.5 mm, apex
acute, strigose. Flower with the calyx bilobulated, 2–
2.5 mm, lightly strigose; corolla tube 3–4 mm,
externally glabrous, with villous fauce. Cluse 1.5 3 1
mm, glabrous.
Distribution and habitat. Aloysii dusenii is endemic to Brazil (Paraná, Rio Grande do Sul, Santa
Catarina) and has been observed to grow on
riverbanks or on marginal moist forests.
Discussion. Aloysia dusenii is remarkable in
relation to its bilobulated calyx, which is 4-toothed
in the rest of the South American Aloysia taxa.
Bilobulated calyces are seen among North American
Aloysia such as A. coalcomana Siedo and are frequent
within Lippia.
Selected specimens examined. BRAZIL. Paraná: Guarapuava, Rio Jordao, Hatschbach 22546 (SI); General
Carneiro, faz. dos Souzas, Hatschbach 28366 (SI). Rio
Grande do Sul: Bom Jesus, Pelotas, de S. Joaquim a
Ronchinha, Krapovickas 38344 (SI). Santa Catarina: Rio
Pelotas, crossing rd. from Silveira to Sao Joaquim, Olmstead
2010-217 (SI); Mun. Sao Joaquim, Lajes, Lourteig 2189
(SI).
11. Aloysia fiebrigii (Hayek) Moldenke, Revista
Sudamer. Bot. 4: 15. 1937. Basionym: Lippia
fiebrigii Hayek, Bot. Jahrb. Syst. 42: 165. 1909.
TYPE: Bolivia. Tarija, 4 Feb. 1904, K. Fiebrig
3036 (lectotype, designated by Wood [2009: 515],
K [barcode] K000470999 not seen, K image!;
isolectotypes, A [bc] A00069310 not seen, A
image!, BM [bc] BM000643747 not seen, BM
image!, E [bc] E00373278 not seen, E image!,
F [bc] F0092410F not seen, F image!, F [bc]
F0093491F not seen, F image!, G [bc] G00386442
not seen, G image!, GH [bc] GH00069311 not
seen, GH image!, GOET [bc] GOET011516 not
seen, GOET image!, M [bc] M0111834 not seen,
M image!, NY [bc] NY00137758 not seen, NY
image!, P [bc] P00713756 not seen, P image!, P
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2016
[bc] P00713757 not seen, P image!, S11-10531
not seen, S image!, S11-10526 not seen, S image!,
SI [bc] SI003507!, SI [bc] SI003383!, SI [bc]
SI003508!, US [bc] US00118823 not seen, US
image!, W 1911-0001742 not seen, W image!).
Aloysia arcuifolia G. L. Nesom, Phytologia 70: 145. 1991.
TYPE: Bolivia. Potosi: Valle de Palqui, 7 Feb. 1987,
R. Ehrich 339 (holotype, TEX [barcode]
TEX00374932 not seen, TEX image!; isotype, LPB
[bc] LPB0000837 not seen, LPB image!).
Shrubs 1–2 m tall. Leaves verticillate, in whorls of
3, sometimes 4, rarely opposite, subsessile; blades
linear to elliptic, 1.8–3 3 0.2–0.5 cm, apex acute,
base acute, attenuate, margin entire, adaxially
scabrous, abaxially glabrate, with underlayer of
glandular trichomes, midvein abaxially conspicuous.
Florescences terminal and axillary (heterothetic
pleiobotrya), dense, brief, 1–2 cm; peduncles 0.5–1
cm; flowers white; floral bracts ovate, apex acuminate, 2–2.5 mm, lightly strigose, margin ciliate.
Flower with the calyx 1.5 mm, strigose, with 4 brief
teeth, unequal, triangular; corolla tube 5.5 mm, with
glabrous fauce. Cluse 1 3 0.5 mm, glabrous.
Iconography. Botta (1979: 102, fig. 11).
Distribution and habitat. Aloysia fiebrigii has
been observed to grow in southern Bolivia and
northwestern Argentina (Jujuy, Salta).
Discussion. Siedo (2010) lectotypified this taxon
superfluously, because Wood had previously designated the lectotype for Aloysia fiebrigii (Wood,
2009).
Aloysia fiebrigii shares with A. arequipensis, A.
citrodora, A. herrerae, and A. velutina the presence of
axillary and terminal florescences, a group of five
species in Aloysia with heterothetic pleiobotrya. It is
distinguished from these species, except from A.
citrodora, by its ternate leaves, with the leaves being
opposite in the rest. Aloysia citrodora and A. fiebrigii
may be differentiated by their leaves, being smaller
(blades 1.8–3 3 0.2–0.5 cm) and with entire margins
in the latter, and larger (blades 2–8 3 1–2.5 cm) and
with entire or slightly serrate margins in the former.
Selected specimens examined. ARGENTINA. Jujuy:
Santa Catalina, Cieneguillas, Arenas 927 (SI). Salta:
Hunziker 8055 (CORD). BOLIVIA. Chuquisaca: entre
Muyuquiri y Camargo, Cocucci 3366 (SI). Tarija: Yunchará, Rupaska, Beck 26988 (SI); s. loc., s.d., Fiebrig
3040 (SI).
12. Aloysia gratissima (Gillies & Hook. ex Hook.)
Tronc., Darwiniana 12: 527. 1962. Basionym:
Verbena gratissima Gillies & Hook. ex Hook.,
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Bot. Misc. 1: 160. 1829. TYPE: Argentina.
Mendoza, 1829, G. Gillies s.n. (holotype, K
[barcode] K000470994 not seen, K image!;
isotypes, BM [bc] BM000643772 not seen, BM
image!, BM [bc] BM000643773 not seen, BM
image!, E [bc] E00320620 not seen, E image!, E
[bc] E00320623 not seen, E image!, E [bc]
E00320622 not seen, E image!, GH [bc]
GH00069316 not seen, GH image!, GH [bc]
GH00299005 not seen, GH image!, OXF not
seen, SI [bc] SI003701!). Figure 5.
Shrubs 1–3 m tall; stems glabrous, sometimes
puberulous when young. Leaves opposite, subsessile,
or briefly petiolate to 3 mm, sometimes fasciculate,
membranaceous or coriaceous; blades elliptic to
ovate, 0.15–7 3 0.05–3 cm, apex acute or sometimes
subobtuse, base acute, margins entire, dentate, or
serrate, or basally entire and serrate toward apex,
adaxially scabrous, abaxially glabrate to lightly
strigose, midvein only impressed on abaxial surface
or the venation pinnate, conspicuous. Florescences
axillary, solitary, lax, or dense, 3–13 cm; peduncles
0.5–3 cm; flowers white; floral bracts ovate, apex
acute, 1–2 mm, lightly strigose, margin ciliate.
Flower with the calyx 2–4 mm, hispid in inferior
half, lightly hispid toward apex, calyx with 4 teeth,
unequal, triangular; corolla tube 3–5 mm, externally
puberulous toward apex, with villous fauce. Cluse 1–
2 3 0.5 mm, glabrous.
Distribution and habitat. Aloysia gratissima is an
amply distributed species, growing throughout the
Americas. Aloysia gratissima var. gratissima has the
most far-reaching distribution, being the only taxon in
the genus that occurs in both South and North
America. Its geographic distribution extends from the
southwestern United States and northern Mexico, into
southern South America. Three other varieties of A.
gratissima are here recognized (A. gratissima var.
schulziana (Moldenke) Botta, A. gratissima var.
angustifolia (Tronc.) Botta, and A. gratissima var.
chacoensis (Moldenke) Botta) and have distributions
restricted to southern South America. Consequently,
in South America four varieties of A. gratissima are
recognized.
Discussion. Ragonese (1955) stated that Aloysia
gratissima is an aromatic shrub known to be toxic for
cattle.
Aloysia gratissima is a problematic taxon, and
throughout the years botanists (Troncoso, 1964; Botta,
1979; Siedo, 2006) have differed in the significance of
singular characters to delimit this complex, which has
been reflected in the variable number of species and
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Figure 5. Aloysia gratissima (Gillies & Hook.) Tronc. A–F. Aloysia gratissima var. gratissima. —A. Basal branch. —B. Apical
branch. —C. Fructiferous branch. —D. Leaf. —E. Intact flower with calyx hispid in lower half, with three teeth (of four) visible.
—F. Dissected calyx, internal view. G–J. Aloysia gratissima var. chacoensis (Moldenke) Botta. —G. Floriferous branch with dentate
leaves. —H. Apical branch with entire leaves. —I. Basal leaf with dentate margin in upper half. —J. Apical leaf with entire
margins. K–M. Aloysia gratissima var. angustifolia (Tronc.) Botta. —K. Fructiferous branch with apical leaves. —L. Floriferous
branch with basal leaves. —M. Mature leaf. N–Q. Aloysia gratissima var. schulziana (Moldenke) Botta. —N. Floriferous branch.
—O. Fructiferous branch. —P. Leaf, abaxial surface with conspicuous pinnate venation. —Q. Leaf, adaxial surface. A–F from
Schinini 10366 (SI); G–J from Schulz 1494 (holotype, A. gratissima var. chocoensis, CTES); K from Burkart 23805 (SI); L–M from
Burkart 6695 (holotype, A. gratissima var. angustifolia, SI); N from Krapovickas 30899 (SI); O–Q from Cabrera 14633 (SI).
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varieties recognized over time. Lu-Irving et al. (2014)
hypothesized that this group may represent a recent
radiation, given that branch lengths are short
throughout the A. gratissima clade. Therefore, a
population-level approach to sampling may be required to elucidate the identities and evolutionary
histories of taxa belonging to this clade. Recently,
Moroni et al. (2016) applied a statistical approach to
try to resolve relationships within A. gratissima.
margins, and a conspicuous midvein abaxially.
Florescences dense, 3–7 cm long.
Cytology. A mitotic chromosome count of 2n ¼
54 has been reported for the species Aloysia
gratissima (Powell et al., 2010).
TAXONOMIC KEY
TO THE
VARIETIES
OF
ALOYSIA
GRATISSIMA
1.
Apical leaf blades small, less than 2 cm long, or
small throughout the entire stem axis; florescences dense, 3–7 cm long . . . . . . . . . . . . . . . . . . . . . . . . . . . 2
1 0 . Apical leaf blades larger than lower ones, more
than 2 cm long; florescences lax, 5–13 cm long . . . 3
2. Leaf blades with entire margins, less than 2
cm long throughout the entire plant . . . . . .
. . . . . . . . . . . 12a. A. gratissima (Gillies & Hook.
ex Hook.) Tronc. var. angustifolia (Tronc.) Botta
2 0 . Leaf blades on upper plant stems with entire
margins or sometimes 1- to 3-dentate,
blades less than 1 cm long, leaf blades on
lower plant stems with 1 to 4 teeth, blades
more than 2.5 cm long . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . 12b. A. gratissima (Gillies & Hook.
ex Hook.) Tronc. var. chacoensis Moldenke
3. Leaves with an acute apex and serrate
margins; abaxial blades with conspicuous pinnate venation . . . . . . . . . . . . . .
. . . 12d. A. gratissima (Gillies & Hook. ex
Hook.) var. schulziana (Moldenke) Botta
3 0 . Leaves with an acute or subobtuse
apex and entire margins, sometimes
serrate; abaxial blades with only the
midvein impressed . . . . . . . . . . . . . . . . .
. . . 12c. A. gratissima (Gillies & Hook. ex
Hook.) Tronc. var. gratissima
12a. Aloysia gratissima (Gillies & Hook. ex Hook.)
Tronc. var. angustifolia (Tronc.) Botta, Darwiniana 22: 89. 1979. Basionym: Aloysia chacoensis
var. angustifolia Tronc., Darwiniana 13: 630.
1964. TYPE: Argentina. Buenos Aires: Campana, 28 Oct. 1934, A. Burkart 6695 (holotype, SI
[barcode] SI003379!; isotype, SI [bc]
SI003378!). Figure 5K–M.
Aloysia decipiens Ravenna, Onira 11(4): 14–15. 2007.
TYPE: Argentina. Santiago del Estero: Dpto. Robles,
colonia Jaime, 19 Nov. 1948, B. Luna Ruiz s.n.
(holotype, BA [barcode] BA53388!).
Xerophytic shrubs; leaves small, narrow elliptic,
up to 2 3 0.3 cm, with an acute apex and base, entire
Distribution and habitat. Aloysia gratissima var.
angustifolia grows in central Argentina (Buenos
Aires, Chaco, Córdoba, Entre Rios, La Pampa, La
Rioja, Santa Fé, and Santiago del Estero) and also in
western Paraguay.
Discussion. Ravenna (2007) stated that Aloysia
decipiens was closely related to A. beckii Moldenke;
however, this name is treated as a synonym of A.
gratissima var. gratissima herein. Study of the type
material of A. decipiens evinces that it shares a similar
leaf morphology with A. gratissima var. angustifolia,
and therefore Múlgura et al. (2012) considered the
species name a synonym of this variety.
Selected specimens examined. ARGENTINA. Buenos
Aires: Campana, barranca Paraná, Krapovickas 2597 (NY).
Chaco: Resistencia, Malvarez 1367 (NY). Córdoba: Punilla,
Villa del Lago, Cabrera et al. 29664 (SI). Diamante:
Barranca, Burkart 28067 (SI). Entre Rı́os: Paraná, Paracao,
Burkart et al. 23805 (SI). La Pampa: Guatraché, Estancia
Remecó, Rúgolo 1275 (SI). La Rioja: Chilecito, Meyer
4087 (NY). Santa Fé: General Obligado, Villa Ana, Villa
Ocampo, Ragonese 3120 (SI). Santiago del Estero: San
Martı́n, ruta 34, Km. 571, Aliscioni et al. 692 (SI).
Tucumán: s.d., D. Cozzo s.n. (NY). PARAGUAY. Boquerón: Colonia Menno, Arenas 1085 (SI). Chaco: Loma Porá,
Rojas 2542 (SI).
12b. Aloysia gratissima (Gillies & Hook. ex Hook.)
Tronc. var. chacoensis (Moldenke) Botta, Darwiniana 22: 89. 1979. Basionym: Aloysia
chacoensis Moldenke, Lilloa 5: 373. 1940.
TYPE: Argentina. Chaco: Tirol, on side of
mtn., Feb. 1934, A. G. Schulz 1494 (holotype,
NY [barcode] NY00103873 not seen, NY
image!; isotype, CTES [bc] CTES0013810 not
seen, CTES image!). Figure 5G–J.
Aloysia casadensis Hassl. ex Moldenke, Phytologia 3: 106–
107. 1949. TYPE: Paraguay. Puerto Casado, Feb.
1917, T. Rojas 2529 (holotype, MVM not seen;
isotypes, NY [barcode] NY00103869 not seen, NY
image!, NY [bc] NY00103868 not seen, NY image!, SI
[bc] SI003388!).
Leaves small, elliptic, with lengths of less than 1
cm toward apical portion of plant stems, but lengths
to 4 cm with 1 to 4 teeth, toward basal portions.
Florescences dense, 3–7 cm long.
Distribution and habitat. Most specimens of
Aloysia gratissima var. chacoensis grow in the
Argentinean provinces of Chaco and Formosa. This
variety is less commonly found in Santa Fé, Santiago
del Estero, and Córdoba, and also occurs in western
Paraguay.
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Selected specimens examined. ARGENTINA. Chaco:
Enrique Urién, Schulz 772 (NY); Pampa del Infierno, Meyer
5066 (NY). Córdoba: Ischilı́n, Dean Funes a San Vicente,
de la Sota 198 (NY). Formosa: Camino de Fortı́n Nuevo,
Pilcomayo a La Esmeralda, Cordini 60 (SI). Santa Fé: San
Bernardo, Alonso 548 (SI). Santiago del Estero: Espada,
˜ ´ s 61 (SI). PARAGUAY. Alto Paraguay: Puerto
Arganara
Casado, Rojas 7701 (SI). Boquerón: Colonia Fernheim,
Filadelfia, Arenas 1879 (SI). Presidente Hayes: Ea. Loma
Pyta, Arenas 664 (SI).
Leaves elliptic to ovate, generally glabrate, with an
acute to subobtuse apex and blade margins entire,
sometimes slightly serrate; abaxial blade surface with
only the midvein impressed. Florescences lax, 5–13
cm long.
12c. Aloysia gratissima (Gillies & Hook. ex Hook.)
Tronc. var. gratissima. Figure 5A–F.
Aloysia lycioides Cham., Linnaea 7: 237. 1832. Lippia
lycioides (Cham.) Steud., Nomencl. Bot. [Ed. 2] 2: 62.
1841. TYPE: Brazil meridian, s.d., F. Sellow s.n.
(lectotype, designated by Múlgura et al. [2012: 17], K
[barcode] K000487005 not seen, K image!, K;
isolectotypes, BM [bc] BM001118329 not seen, BM
image!, BR [bc] BR0000005720156 not seen, BR
image!, HAL [bc] HAL0107063 not seen, HAL
image!, HAL [bc] HAL0107062 not seen, HAL
image!, K [bc] K000487006 not seen, K image!, G
[bc] G00386444 not seen, G image!, M [bc]
M0111831 not seen, M image!, NY not seen, NY
photo at SI!, US [bc] US01049793 not seen, US
image!, SI [bc] SI003390!, SI [bc] SI003391!).
Aloysia floribunda M. Martens & Galeotti, Bull. Acad. Roy.
Sci. Bruxelles, 11(2): 320. 1844, syn. nov. TYPE:
Mexico. Veracruz: June–Oct. 1840, H. Galeotti 774
(holotype, BR [barcode] BR005187300 not seen, BR
image!).
Lippia ligustrina (Lag.) Britton var. lasiodonta Briq.,
Annuaire Conserv. Jard. Bot. Genève 7–8: 305.
´ 15 Mar. 1881,
1904. TYPE: Paraguay. Paraguariı,
B. Balansa 3117 (holotype, G [barcode] G00166265
not seen, G image!; isotypes, BM [bc] BM000643660
not seen, BM image!, GH [bc] GH00299001 not seen,
GH image!, F [bc] F0092411F not seen, F image!, K
[bc] K000487007 not seen, K image!, SI!).
Lippia ligustrina (Lag.) Britton var. paraguariensis Briq.,
` 7–8: 305. 1904.
Annuaire Conserv. Jard. Bot. Geneve
Aloysia ligustrina var. paraguariensis (Briq.) Moldenke, Phytologia 1: 167. 1935. Aloysia lycioides var.
paraguariensis (Briq.) Moldenke, Phytologia 2: 464.
1948. Aloysia gratissima (Gillies & Hook.) Tronc. var.
paraguariensis (Briq.) Moldenke, Phytologia 9: 500.
1964. TYPE: Paraguay. Paraguarı́, Jan. 1875, B.
Balansa 1015 (holotype, G [barcode] G00166413 not
seen, G image; isotypes, BR not seen, BR photo at SI!,
G [bc] G00166414 not seen, G photo at SI!, G image!,
SI-67609!).
Aloysia beckii Moldenke, Phytologia 52(1): 18. 1982. TYPE:
Bolivia. Cochabamba: Carrasco, Cochabamba hacia
Santa Cruz, 27 Sep. 1981, S. Beck 7036 (holotype,
TEX-LL [barcode] LL00374934 not seen, TEX-LL
image!; isotypes, LPB [bc] LPB0000839 not seen,
LPB image!, M [bc] M0112713 not seen, M image!, SI
[bc] SI003374!).
Aloysia famatinensis Ravenna, Onira 11(4): 15–16. 2007.
TYPE: Argentina. La Rioja: Sierra de Famatina,
Guanchı́n Viejo, 25 Jan. 1928, A. Castellanos s.n.
(holotype, BA-28328!).
Distribution and habitat. Aloysia gratissima var.
gratissima is amply distributed across northern,
northeastern, and northwestern Argentina, extending
south to La Pampa. The autonymic variety also occurs
in Chile, Bolivia, Paraguay, Uruguay, and southern
Brazil. It is found across northern Mexico and in the
southwestern United States. This is the only taxon of
Aloysia known to occur in both South and North
America, with an interesting distributional disjunction between northern populations (southwestern
United States and adjacent Mexico) and southern
ones (Brazil and south). Aloysia gratissima var.
gratissima has been collected and noted as frequent
along riverbanks or in xeromorphic scrublands.
Discussion. In northeastern Corrientes Province,
Argentina, as well as Paraguay, a phenotype exists
(Morel 1257 [NY]; Jorgensen 2473 [NY]) that has
slightly obovate or rounded leaves, with an entire
margin and obtuse apex, with a prominent midvein
abaxially and multiple fasciculate leaves at stem
nodes. This resembles what Moldenke (1935)
referred to as Aloysia ligustrina var. paraguariensis,
which is herein considered a synonym of A.
gratissima var. gratissima.
Study of the type material of Aloysia floribunda
indicates that this species is the same taxon as A.
gratissima var. gratissima. This was first proposed by
Múlgura et al. (2012), given that the type specimen
has lax florescences longer than 5 cm, with leaves
with a subobtuse apex and entire margin. Wood
(2009) mentioned that the holotype of A. lycioides was
housed at LE, but none of the authors have seen it. A
lectotype was selected by Múlgura et al. (2012) from
a specimen housed at K.
Selected specimens examined. ARGENTINA. Buenos
Aires: Campana, Burkart 3083 (SI). Catamarca: Tinogasta,
cuesta de Zapata, Cabrera 24648 (SI). Chaco: Colonia
Benı́tez, Schulz 8317 (SI). Córdoba: Punilla, Los Gigantes,
Sayago 940 (SI). Corrientes: Santo Tomé, Arroyo Chimiray,
´ Uruguay, Schinini 10366 (CTES; SI). Curuzú Cuatiá:
Rıo
Est. Marı́a Azucena, Schinini 13902 (SI). Entre Rios:
Concordia, Salto Grande, Cabrera 19245 (SI). Formosa: s.
loc., Jörgensen 2473 (NY). Jujuy: Tumbaya, Volcán,
Laguna, Burkart 30608 (SI). La Pampa: Sierra de Lihuel
Calel, 30 Nov. 1959, Troncoso s.n. (SI 20589). La Rioja:
´ Okada 2747 (SI). Mendoza: Tunuyán,
Chilecito, Quanchın,
Ruiz Leal 1102 (SI). Misiones: Cainguás, rte. 7, camino de
Aristóbulo del Valle a Jardin América, Morrone 635 (SI).
Patino:
˜ Las Lomitas, 3–5 km SO por rte. 81, Schinini
24130 (SI). Pilcomayo: Clorinda, Morel 1257 (NY). Salta:
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Anta, finca Pozo Largo, Saravia Toledo 716 (SI). San Juan:
Valle Fértil, Sa. De Valle Fértil, Kiesling 4954 (SI). San
Luis: La Capital, San Martin del Alto Negro, L. Anderson
1521 (SI). Santa Fe: Garay, entre Santa Rosa y Cayastá,
Ragonese 2829 (SI). Santiago del Estero: Ojo de Agua,
Fabris 2729 (SI). Tucumán: Tafı́ del Valle, Schulz 11465
(SI). BOLIVIA. Chuquisaca: Luis Calvo, clausura El Huare,
Saravia Toledo 11830 (SI). Cochabamba: Mizque, canton
Molinero, Sigle 137 (SI). Santa Cruz: Samaipata, Steinbach
8248 (NY). Tarija: de Tarija a Narvaez, Kiesling 3708 (SI).
BRAZIL. Rio Grande do Sul: Alegrete a Uruguayana,
Nicora 4726 (SI). CHILE. Region V: Limache, Garaventa
7092 (SI). ECUADOR. Pichincha: Quito, Parque Italia,
Barrio Las Casas, Cerón 12387 (SI). MEXICO. Chihuahua:
Bachimba canyon, 23 Mar. 1885, Pringle s.n. (SI).
Guanajuato: 60 mi. S San Luis Potosi, Dunn 20552
(MO). Nuevo Leon: valley near Monterey, 31 Aug. 1903,
Pringle s.n. (SI). PARAGUAY. Boquerón: Colonia Menno,
Lolita, Vanni 1830 (SI). Chaco: Santa Elisa, Hassler 2635
(SI). Cordillera: San Bernardino, Hassler 53 (SI). Guaira:
Cordillera de Ybytyruzú, Cerro. Polilla, Zardini 8599 (SI).
´ Rojas 7902 (SI). Paraguarı́: de
Itapúa: Villa Encarnacion,
Paraguarı́ a Escobar, Múlgura 3743 (SI). San Pedro: Alto
Paraguay, Primavera, Woolston 794 (NY). U.S.A. Texas: Jim
Hogg Co., NE Hebronville, Lundell 11958 (SI). URUGUAY. Artigas: Tomás Gomensoro, Marchesi 10078 (SI).
Minas: Arroyo Aguas Blancas, Crespo 26457 (SI). Paysandú: Arroyo Negro, Rosengurtt 2245 (SI). Rivera: rte. 29,
pasando Minas de Corrales, Denham 333 (SI). Tacuarembó: Cabrera 32345 (SI).
na (Jujuy, Salta). This variety is also known, but less
frequently, from the Argentinean provinces of
Catamarca, Chaco, Formosa, La Rioja, Misiones,
´ Aloysia gratissima var.
San Juan, and Tucuman.
schulziana is also present in southern Bolivia and
Paraguay.
12d. Aloysia gratissima (Gillies & Hook. ex Hook.)
Tronc. var. schulziana (Moldenke) Botta, Darwiniana 22: 87. 1979. Basionym: Aloysia
schulziana Moldenke, Lilloa 5: 381. 1940.
TYPE: Argentina. Salta: Colonia San Bernardo,
Feb. 1936, A. G. Schulz 1447 (holotype, NY
[barcode] NY00103887 not seen, NY image!;
isotypes, CTES [bc] CTES0013830 not seen,
CTES image!, SI [bc] SI003394!) Figure 5N–Q.
Aloysia meyeri Moldenke, Lilloa 5: 378. 1940. TYPE:
Argentina. Tucumán: Trancas, San Pedro de Colalao,
4 ene. 1940, T. Meyer 3092 (holotype, NY [barcode]
NY00103881 not seen, NY image!; isotypes, LIL [bc]
LIL001365 not seen, LIL image!, SI [bc] SI003389!).
Aloysia looseri Moldenke, Lilloa 5: 377. 1940. Lippia looseri
(Moldenke) Looser, Revista Univ. (Santiago) 26(2):
141. 1941. TYPE: Chile. Santiago, 15 Dec. 1925,
[cult.] G. Looser 4008 (holotype, NY [barcode]
NY00103879 not seen, NY image; isotypes, SI!,
SGO not seen).
Aloysia lomaplatae Ravenna, Onira 10(19): 60. 2006.
TYPE: Paraguay. ‘‘In scopulosis ad Loma Plata, civit.
Boquerón,’’ 13 Sep. 2001, P. Ravenna 5030 (holotype, FCQ not seen).
Leaves elliptic, with an acute apex and serrate
margin; blades with pinnate venation conspicuous on
abaxial surface. Florescences lax, longer than 5 cm.
Distribution and habitat. The greatest number of
specimens attributed to Aloysia gratissima var.
schulziana were collected from northwestern Argenti-
Discussion. Aloysia gratissima var. schulziana is
distinguished by its leaves with serrate margins and
conspicuous pinnate venation on the abaxial blade
surface, with the margins entire or slightly serrate
toward apex. Only the midvein is impressed in the
other three varieties.
Moldenke (1940) mentioned that type material of
Aloysia looseri was a specimen cultivated in Chile.
The species name is herein considered a synonym of
A. gratissima var. schulziana. Chile is not considered
as part of the natural distributional range for A.
gratissima var. schulziana. Elsewhere, this variety is
sometimes cultivated for ornamental purposes, as in
the suburbs of Buenos Aires.
Ravenna (2006: 60) stated in the protologue of
Aloysia lomaplatae Ravenna that the species can be
distinguished by its ‘‘relatively large, somewhat
scabrous, conspicuously lobulate-dentate blades.’’
These are characters that define the variety, and
consequently the species name is here considered a
synonym of A. gratissima var. schulziana.
Selected specimens examined. ARGENTINA. Catamar˜ Troncoso 1824 (SI).
ca: Ancasti, entre Ancasti e Icano,
´
Chaco: 18 de Mayo, Colonia Benıtez,
Schulz 8699 (SI).
Córdoba: Quilino, Cordo 77-d-45 (SI). Corrientes: Ituzaingó, camino a San Carlos, Krapovickas 17970 (SI). Formosa:
Matacos, Ingenario Juarez, 3 km al S del pueblo, sobre la
˜
rte. que va al Bermejo, a Belgrano, Arenas 2330 (SI); Patino,
Las Lomitas, Maranta 1104 (BA). Jujuy: Santa Bárbara,
Cabrera 31018 (S); entre San Pedro y Santa Clara, Cabrera
14633 (SI). La Rioja: Independencia, rte. 79, Pensiero
7426 (SI). Misiones: Posadas, Bertoni 761 (NY). Salta:
Anta. San Javier (8 km al Sud de I.V. González), 1988,
Saravia Toledo 1750 (SI). San Juan: Valle Fértil, Ruta
prov. 42, entre San Agustı́n de Valle Fértil y Las Tumanas,
´
a 23 km del primero, Biurrum 4002 (SI). San Martı́n: Rıo
Seco, Krapovickas 30899 (SI). Santiago del Estero: Choya,
Sayago 2649 (SI). Tarija: Villa Montes, Abra de Tapecua,
Krapovickas 19210 (SI). Tucumán: Leales, La Florida,
Krapovickas 17337 (SI). BOLIVIA. Potosı́: Nor Chichas,
Aripalca, 10 Mar. 1993, Torrico 107 (SI). Tarija: Villa
Montes, Abra de Tapecua, 25 May 1971, Krapovickas 19210
(SI). PARAGUAY. Boquerón: s. loc., Vanni 2410 (SI).
13. Aloysia hatschbachii Moldenke, Phytologia
18(6): 341. 1967. TYPE: Brazil. Paraná: Pien,
8 Mar. 1967, G. Hatschbach 16101 (holotype,
NY [barcode] NY00103876 not seen, NY
image!; isotypes, MO-2217903 not seen, MO
image!, TEX-LL [bc] LL00374938 not seen,
TEX-LL image!, TEX-LL [bc] LL00374939 not
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Figure 6. Aloysia hatschbachii Moldenke. —A. Floriferous branch, general aspect, with axillary florescences (homothetic
pleiobotrya). —B. Detail of branch, showing elliptic leaves with serrulate margins in upper half. —C. Intact flower with hispid
calyx subtended by strigose floral bract (at left). —D. Close-up of abaxial leaf surface, glabrous with lightly strigose midvein.
—E. Close-up of adaxial leaf surface, glabrous with glandular papillae. A, illustrated from Hatschbach 16101 (isotype, SI); B–E
from Hatschbach 51897 (SI).
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seen, TEX-LL image!, UC [bc] UC1414074 not
seen, UC image!, US [bc] US01049795 not seen,
US image!, SI [bc] SI041045!, SI [bc]
SI003385!, SI [bc] SI003384!, WIS [bc]
WIS0255184 not seen, WIS image!). Figure 6.
Shrubs 1–2 m tall; stems glabrous. Leaves
opposite, subcoriaceous, briefly petiolate, petiole 2–
4 mm; blades elliptic, 2–3 3 1–1.5 cm, apex
subobtuse to obtuse, base acute, decurrent, margins
entire toward base, serrulate in upper half, adaxially
glabrous, with glandular papillae, abaxially glabrous,
scarcely strigose over abaxial midvein, with venation
conspicuous, reddish brown, impressed on abaxial
surface. Florescences axillary, solitary, lax, 3–8 cm;
peduncles 1–4 cm; flowers white to lilac; pedicels 1
mm; floral bracts reduced, narrowly elliptic, 1–1.2
mm, strigose, apex acute. Flower with the calyx 2–3
mm, hispid, with 4 brief teeth, unequal, triangular;
corolla tube 3–4 mm, externally glabrate, with villous
fauce. Cluse 1.5 3 1 mm, glabrous.
Distribution and habitat. Aloysia hatschbachii is
endemic to Brazil (Paraná), where it has been noted
to grow in rocky soils.
Discussion. Aloysia hatschbachii shares with A.
salviifolia and varieties of A. gratissima the presence
of leaf margins with some degree of serration or
dentition, but they are always basally entire. Aloysia
hatschbachii is readily recognized by its conspicuous,
reddish brown, impressed pinnate venation on the
abaxial surface of the leaf blades.
Selected specimen examined. BRAZIL. Paraná: Pien,
Hatschbach 51897 (SI, US).
14. Aloysia herrerae Moldenke, Phytologia 2: 10.
1941. TYPE: Peru. Urubamba valley, July 1927,
F. L. Herrera 1534 (holotype, F [barcode]
F0043367F not seen, F image!; isotype, NY
[bc] NY00103877 not seen, NY image!, SI!).
Figure 7.
Aloysia ayacuchensis Moldenke, Phytologia 6(6): 256, 323.
1958, syn. nov. TYPE: Peru. Ayacucho, J. J. Soukup
4187 (holotype, NY [barcode] NY00103866 not seen,
NY image!).
Shrubs 1–2.3 m tall; stems glabrous. Leaves
opposite, sometimes ternate, subsessile or briefly
petiolate, petioles 1–5 mm; blades subcoriaceous,
narrowly elliptic, 2( 5.5) 3 0.4( 1.5) cm, apex
subobtuse, base obtuse, margins entire, adaxially
glabrous or with scattered scabrid hairs, especially
along midvein, abaxially scabrid with glandular
trichomes and prominent midvein. Florescences
terminal and axillary (heterothetic pleiobotrya),
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dense, 1–3 cm; peduncles 1 cm; flowers white,
cream with pink to violet tube; pedicels 0.5 mm;
floral bracts reduced, widely ovate, 2–2.5 mm, apex
acuminate, with scarce scabrous pubescence. Flower
with the calyx 2–3 mm, with underlayer of sessile
glandular trichomes, and scarce scabrous pubescence, calyx with 4 brief acuminate teeth; corolla
tube 3–5 mm, strigose on both surfaces. Cluses 1.5 3
1 mm, glabrous.
Distribution and habitat. Aloysia herrerae grows
in Peru and Bolivia and has been collected from
hillsides, rocky slopes, or open sunny areas. It has
been found at elevations up to 2800 m.
Discussion. Aloysia herrerae is an aromatic
plant, the leaves of which are used to treat
headaches. The species is distinguished by its
terminal florescences, which are also present in A.
citrodora and A. fiebrigii, with these three taxa
sharing heterothetic pleiobotyra. However, A. herrerae is differentiated by its opposite leaves, in
contrast to the verticillate leaves seen in A. citrodora
and A. fiebrigii. Terminal florescences are also
present in A. velutina and A. arequipensis, but these
two species lack entire leaf margins, while the leaf
margin is consistently entire in A. herrerae. Leaves
are serrulate in the upper portion of the blade in A.
arequipensis but crenate in A. velutina.
The type specimen of Aloysia ayacuchensis was
observed to have both terminal and axillary florescences and opposite leaves. These are both characters that define A. herrerae, so the species name is
here considered a synonym of the latter.
Selected specimens examined. BOLIVIA. Chuquisaca:
˜
Zudanez,
5 km from Candelaria toward Icla, Wood 14658
˜ de Chancos, Sander(K). PERU. Ancash: Huaraz, banos
man 4612 (F, US). Ayacucho: Ayacucho, Ochoa 710 (NY,
SI); s. loc., Soukup 5467 (US). Cajamarca: San Miguel,
Niepos, Llatas Quiroz 1513 (SI). Cuzco: Calca, C. Vargas
248 (NY, SI); Urubamba, Chincheros, Davis 1757 (F, K,
NY); Urubamba, Huallabamba, Olmstead 2009-30 (SI,
WTU); Huayoccari to Yabncocha, Núñez 7018 (F).
Huancavelica: Tayacaja, betw. Izcuchaca & Acostambo,
Hutchison 4199 (NY, SI, US).
15. Aloysia oblanceolata Moldenke, Phytologia 3:
108. 1949. TYPE: Paraguay. San Bernardino,
1915, T. Rojas 53a (holotype, NY [barcode]
NY00103883 not seen, NY image!; isotypes,
MVM not seen, SI [bc] SI003407!). Figure 8.
Aloysia gratissima (Gillies & Hook. ex Hook.) Tronc. var.
oblanceolata Moldenke, Phytologia 15: 462. 1968,
syn. nov. TYPE: Brazil. Rio Grande do Sul: Gloria, SE
of Porto Alegre, 2 Oct. 1948, A. L. Moldenke 19684
(holotype, NY [barcode] NY00103875 not seen, NY
image!).
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Figure 7. Aloysia herrerae Moldenke. —A. Fertile branch, with florescences in both axillary and terminal positions
(heterothetic pleiobotrya). —B. Intact flower subtended by widely ovate floral bract. —C. Close-up of floral bract surface, with
scattered scabrid hairs and ciliate margins and underlayer of sessile glandular trichomes. —D. Close-up of external calyx
surface, scabrid, with underlayer of glandular trichomes. —E. Close-up of adaxial leaf surface, with scattered scabrid hairs,
especially along midvein. —F. Close-up of abaxial leaf surface, scabrid with glandular trichomes and prominent midvein. A–F,
illustrated from Olmstead 2009-30 (SI).
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Figure 8. Aloysia oblanceolata Moldenke. —A. Fertile branch, general aspect. —B. Flower with hispid calyx and
subtending strigose bract (left). —C. Floral bract with ovate, acute apex. —D. Adaxial glabrous leaf surface. —E. Abaxial leaf
surface, with prominent midvein and revolute margins. A–E, illustrated from Arenas 148 (SI).
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Shrubs 1–2 m tall; stems glabrous. Leaves opposite,
generally clustered into fascicles of 4 to 8 leaves,
blades sessile, obovate to oblanceolate, 2( 3) 3 0.4–
0.6 cm, apex obtuse to round, base acute, attenuate,
margins entire, revolute to subrevolute, coriaceous,
adaxial surface glabrous, with conspicuous midvein
both adaxially and abaxially. Florescences axillary,
solitary, dense, 2–5 cm; peduncles 1–2 cm long;
flowers white; pedicels 0.5 mm; floral bracts widely
ovate, 1–1.5 mm, apex acute, strigose. Flower with the
calyx 2–3 mm, hispid, with 4 teeth, unequal,
triangular; corolla tube 3–4 mm, glabrate externally,
glabrous inside. Cluses 1.5 3 1 mm, glabrous.
and reticulate venation prominent on abaxial surface.
Florescences axillary, solitary, dense, 2.5–10 cm,
elongating to 20 cm in fructification; peduncles 1–2
cm; flowers in whorls of ca. 5, white to pinkish white;
pedicels 0.5 mm, floral bracts ovate, 2.5–4 mm, apex
acute, strigose. Flowers with the calyx 3 mm, hispid,
with long hairs on nerves, with 4 teeth, unequal,
triangular; corolla tube 3–4 mm, externally glabrous,
with villous fauce. Fruit undivided, drupaceous,
obovoid, covered by the acrescent calyx, 2–2.5 3
1.5–2 mm, with two 1-seeded locules.
Distribution and habitat. Aloysia oblanceolata
grows in Paraguay, Bolivia, and Brazil, and is
collected from shrubby forests or rocky soils.
Discussion. Aloysia oblanceolata is similar to
varieties of A. gratissima, sharing a leaf morphology
with blades that can be elliptic with a subobtuse
apex, acute base, and entire margin. Aloysia
oblanceolata may be distinguished by its fasciculate
leaves (vs. not fasciculate in A. gratissima) and its
revolute to subrevolute blade margins (vs. blade
margins not revolute in A. gratissima).
Upon examination, the type material of Aloysia
gratissima var. oblanceolata, later described by
Moldenke in 1968, is undistinguishable from A.
oblanceolata, also described by Moldenke in 1949.
Thus, this varietal name is here considered a
synonym of A. oblanceolata.
Selected specimens examined. BOLIVIA. Cochabamba:
Quillacolllo, 22 km hacia Oruro, Beck 874 (SI). Santa Cruz:
Florida, 13 km W Samaipata, Ferrucci et al. 2659 (SI).
BRAZIL. Paraná: Guarapuava, rio Cavernoso, Hatschbach
9339 (SI). Rio Grande do Sul: Rambo 49976 (SI).
PARAGUAY. Central: Yparacay, Hassler 11497 (NY).
Cordillera: Caacupé, Soria 2098 (NY); Itaugua, cantera,
Arenas 148 (SI).
16. Aloysia ovatifolia Moldenke, Lilloa 5: 379.
1940. Xeroaloysia ovatifolia (Moldenke) Tronc.,
Darwiniana 12: 51. 1960. TYPE: Argentina.
Córdoba: San Javier, La Barranca, 6 Feb. 1939,
A. Castellanos s.n. (holotype, NY [barcode]
NY00103884 not seen, NY image!; isotypes,
BA!, CORD [bc] CORD00003824 not seen,
CORD image!).
Shrubs small, 0.5–0.75( 1.2) m tall; stems densely
puberulent when young, glabrescent in age. Leaves
opposite, sometimes clustered into fascicles, petioles
0.2–1 cm; blades ovate to deltate, 1.5–5 3 1–3 cm,
apex subobtuse, base truncate, margins coarsely
crenate, adaxially hispid, abaxially strigose, pinnate
Iconography. Troncoso (1960: 52, fig. 1, 53, fig.
2, 55, fig. 3).
Distribution and habitat. Aloysia ovatifolia is
endemic to northwestern Argentina where it is found
on rocky slopes, in sunny, sandy, dry soils.
Discussion. Aloysia ovatifolia is unique in Aloysia
in having fruits undivided and drupaceous, being
schizocarpic in the rest of the taxa. It also can be
differentiated by its blades ovate to deltate, with
subobtuse apex, truncate base, and margins coarsely
crenate.
Selected specimens examined. ARGENTINA. Catamarca: Tinogasta, rte. 45, 14 km N Tinogasta, J. Hunziker
11998 (SI). Córdoba: Sobremonte, San Francisco del
˜
Chanar,
Burkart 29641 (SI). La Rioja: General Belgrano,
entre Nepe y Los Baldes, Riedel 226 (SI). Mendoza: San
Carlos, rte. 1001, 6 km S jct. rte. 40, Olmstead 2001-184
(SI). San Juan: Valle Fértil, ceros rodean Embalse, Roig
8424 (SI). San Luis: Belgrano, Estancia El Médano, 65 km
NO de San Luis, L. Anderson 3077 (SI). Santiago del
Estero: Guasayán, de Guampacha a Guasayán, Rotman 228
(SI). Tucumán: Trancas, ruta 9 al S de Choromoro, Burkart
30574 (SI).
17. Aloysia peruviana (Turcz.) Moldenke, Revista
Sudamer. Bot. 4: 15. 1937. Basionym: Lippia
peruviana Turcz., Bull. Soc. Imp. Naturalistes
Moscou 36(2): 200. 1863. TYPE: Peru. Punochuca, s.d., W. Matthews 585 (holotype, KW
[barcode] KW001001633 not seen, KW image!;
isotypes, BM [bc] BM000643629 not seen, BM
image!, BR-562372 not seen, BR [bc]
BR0000005623723 not seen, BR image!, K
[bc] K000470998 not seen, K image!). Figure 9.
Aloysia aloysioides Loes. ex Moldenke, Phytologia 2: 9–10.
1941. Lippia aloysioides Loes. ex Moldenke, nom.
inval. [cheironym]. Phytologia 2: 10. 1941. TYPE:
Peru. Lima, below Surco, Feb. 1909, A. Weberbauer
5206 (holotype, F [barcode] F0043368 not seen, F
image!; isotypes, NY [bc] NY00103865 not seen, NY
image!, S11-10554 not seen, S image!, SI [bc]
SI014007!, US [bc] US00118806 not seen, US
image!).
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Figure 9. Aloysia peruviana (Turcz.) Moldenke. —A. Floriferous branch, with axillary florescences (homothetic pleiobotrya).
—B. Intact flower in lateral view, with hispid calyx and subtending strigose floral bract (at left). —C. Intact leaf, abaxial surface,
evenly crenate margin and prominent venation. —D. Close-up of abaxial leaf surface, with hispid indument. —E. Close-up of
adaxial leaf surface with scattered strigose and hispid hairs. B, illustrated from Soukup 4872 (US); A, C–E, taken from Donovan
P978 (US).
Aloysia minthiosa Moldenke, Phytologia 2: 12. 1941, syn.
nov. TYPE: Peru. Ancash, Yaután, 9 Oct. 1922, J. F.
Macbride 2564 (holotype, F [barcode] F0042838F not
seen, F image!; isotypes, GH [bc] GH00299002 not
seen, GH image!, NY [bc] NY00103882 not seen, NY
image!, SI!).
Shrubs 1–2.5 m tall; stems lightly puberulent to
strigose. Leaves generally opposite; petioles 0.2–0.5
cm long; blades ovate to elliptic, 2–4 3 1–2 cm, apex
subobtuse, base rounded, margins evenly crenate,
adaxially scabrous, abaxially hispid or strigose,
venation pinnate and reticulate, conspicuous on
abaxial surface, membranaceous and plane texture.
Florescences axillary, solitary, dense, 8–10 cm;
peduncles 1–2 cm; flowers white or pale violet;
pedicels 0.5 mm; floral bracts narrowly elliptic, 2–3
mm, apex acute, strigose. Flower with the calyx 2–2.5
mm, hispid, hairs longer proximally, grading to
shorter distally, with 4 teeth, unequal, triangular;
corolla tube 6–7( 8) mm, externally glabrous, with
villous fauce. Cluses 2 3 1 mm, smooth and glabrous.
Distribution and habitat. Aloysia peruviana is
endemic to Peru, growing at elevations from 1200 to
2000 m, on rocky slopes.
Discussion. Lippia aloysioides Loes. is an invalid
name, only mentioned in the protologue of Aloysia
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aloysioides where Moldenke (1941: 10) stated ‘‘The
cheironym, Lippia aloysioides Loes., appears on the
label.’’ A cheironym is an unpublished name and
invalid.
Aloysia peruviana has perfumed, sweet-smelling
flowers, with showy blooms and long corolla tubes 6–
7(8) mm. There are also other species in Aloysia with
long corollas. Aloysia castellanosii would be distinguished by its oblong leaf blades, these being ovate to
elliptic in A. peruviana. Aloysia crenata and A.
citrodora differ in their ternate leaves, these being
opposite in A. peruviana. Aloysia salsoloides, A.
deserticola, and A. tarapacana also may have long
corollas, with lengths of 5–7 mm, 4–6 mm, and 5–6
mm, respectively. However, all three are plants with
spiny branches and reduced leaf blades. Two varieties
of A. scorodonioides also have long corollas, 5–6 cm in
both A. scorodonioides var. mathewsii and A. scorodonioides var. scorodonioides. However, both varieties may be distinguished by rugose leaf textures, in
contrast to a membranaceous texture in A. peruviana.
The type material of Aloysia minthiosa, described
by Moldenke in 1941 from Peru, indicates its
similarity to A. peruviana, with both sharing leaf
and florescence morphology and also long corolla
tubes. The name A. minthiosa is synonymized herein,
to A. peruviana.
Siedo (2010) designated a superfluous lectotype for
Lippia peruviana from BM, stating that all original
material had been destroyed. This has proved
inaccurate and the holotype has been confirmed to
exist at KW in Kiev, where material studied by
Turczaninow is housed (Natalia Shiyan, curator at
KW, 2013, pers. comm.).
cm, apex acute, base truncate, margins entire,
sometimes subrevolute, sclerophyllous, scabrous to
strigose on both surfaces, sometimes glabrate,
venation plane. Florescences axillary, solitary, dense,
4–13 cm; peduncles 1–3 cm. Flowers lilac; floral
bracts sublinear, 2–3.5 mm, apex acuminate, strigose. Flower with the calyx 2–4 mm, hispid, 4toothed, the teeth unequal, triangular; corolla tube 4–
5 mm, externally finely pulverulent, internally with
villous fauce. Cluses 1.5 3 1 mm, glabrous.
Selected specimens examined. PERU. Ancash: Km. 265
on rd. from Conococha to coast, Donovan P978 (US). Lima:
˜ 301 (SI); Camino a
Km. 70 valle del Rimac, Velarde Nunez
Huarachiri, Soukup 4872 (US); Surco, Soukup 3741 (F);
Santa Eulalia, 27 km from Chosica, Gentry 44821 (MO);
Santa Eulalia rd., N of Chosica, Gentry 36089 (F); above
Chosica, betw. Lima & Matucana, Ferreyra 755 (NY, SI);
Huarochiri, 7.5 km above Santa Eulalia rd., Olmstead
2009-45 (SI, WTU); Canta, Ferreyra 12952 (SI); Huarochiri, N Chosica, Ferreyra 759 (F).
Distribution and habitat. Aloysia polygalifolia is
endemic to Brazil from Paraná, Rio Grande do Sul,
and Santa Catarina, and is found in disturbed and dry
areas.
Discussion. Aloysia polygalifolia is similar to A.
cordata because both have ternate leaves adpressed
to the stem. Both have been collected from the state
of Paraná, although A. cordata has been noted from
more mesic conditions. However, A. cordata has
cordate leaf blades that are glabrate and only
minutely scaberulous, whereas leaf blades range
from scabrous to strigose, with a truncate base, in A.
polygalifolia. Exceptionally, leaves can be glabrate
(Thode et al. 398, ICN) as in A. cordata, but the
blades are never cordate in A. polygalifolia. Aloysia
polygalifolia is also similar to A. brasiliensis (see
discussion under A. brasiliensis).
Selected specimens examined. BRAZIL. Paraná: Palmas, Morro da Baliza, Hatschbach 30734 (NY); Palmas, 10
km NO Palmas, Hatschbach 28171 (NY). Rio Grande do
Sul: Guaı́ba, faz. Sao Maximiano, BR-116, Thode 398 (ICN,
SI). Santa Catarina: Ponte Serrada, Fachinal dos Guedes,
Smith & Reitz 12478 (SI); Agua Doce, 5 km S of turn to rd.
´ Smith & Klein 13029
E to Palmas, Smith 15683 (SI); Iranı,
(SI); Agua Doce, campos de Palmas, Smith & Klein 13577
(SI).
18. Aloysia polygalifolia Cham., Linnaea 7: 236.
1832, as ‘‘polygalaefolia.’’ Lippia polygalaefolia
(Cham.) Steud., Nomencl. Bot. [Steudel] 2(2):
54. 1841. TYPE: Brazil. s. loc., s.d., F. Sellow
s.n. (lectotype, designated by Siedo [2010: 201],
G [barcode] G00386462 not seen; isolectotype,
E [bc] E00373271 not seen, E image!). Figure
10.
19. Aloysia polystachya (Griseb.) Moldenke, Lilloa
5: 380. 1940. Basionym: Lippia polystachya
Griseb., Abh. Königl. Ges. Wiss. Göttingen 19:
242. 1874. TYPE: Argentina. Córdoba: Las
˜ Jan. 1871, P. G. Lorentz 130
Mollas, Las Penas,
(holotype, GOET [barcode] GOET008519 not
seen, GOET image!; isotypes, [B , B as F neg.
17535!], CORD [bc] CORD00006124 not seen,
CORD image!, GOET [bc] GOET008520 not
seen, GOET image!, K [bc] K000470937 not
seen, K image!, US [bc] US01049797 not seen,
US image!, SI [bc] SI004434!, SI [bc]
SI003577!, VT [bc] UVMVT026110 not seen,
VT image!).
Shrubs 1–3.5 m tall. Leaves ternate, adpressed to
stem; blades sessile, ovate to elliptic, 0.5–2 3 0.3–1
Shrubs 0.5–1.5 m tall; stems glabrous at maturity,
with short internodes. Leaves alternate, rarely
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Figure 10. Aloysia polygalifolia Cham. —A. Floriferous branches, general aspect, with axillary florescences (homothetic
pleiobotrya). —B. Intact flower with strigose floral bract subtending (at right) the hispid calyx. —C. Intact leaf, adaxial surface,
venation plane. —D. Close-up of adaxial leaf surface, with strigose and scabrous indument. —E. Close-up of abaxial leaf
surface, with strigose and scabrid indument. A, illustrated from Smith & Klein 13577 (SI); B–E from Smith & Klein 13029 (SI).
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opposite, petioles 0.1–0.3 cm; blades elliptic, 1–5 3
0.3–1 cm, apex acute to subobtuse, base acute,
attenuate, margin entire, adaxially scabrous, abaxially densely strigose. Florescences axillary, solitary
or sometimes fasciculate, dense, short, 0.5–3 cm;
peduncles 0.5–1 cm; flowers white, small; floral
bracts reduced, obovate, 1–1.5 mm, hispid. Flowers
with the calyx 1–1.5 mm, puberulous, with 4 teeth,
brief, unequal, triangular; corolla tube 1.5–2 mm,
externally puberulous, with villous fauce. Cluses 1 3
0.5 mm, glabrous.
1(4): 167. 1935, nom. illeg. superfl. Aloysia sellowii
(Briq.) Moldenke, Revista Sudamer. Bot. 4: 15. 1937.
Aloysia gratissima var. sellowii (Briq.) Botta, Darwiniana 22: 85. 1979. TYPE: Uruguay. Montevideo, F.
Sellow 1744 (holotype, G [barcode] G00386443 not
seen, G image!; isotypes, [B , B as F neg. 24670!], W
not seen, W fragm. at SI 66318!).
Aloysia lycioides var. revoluta Moldenke, Phytologia 3: 108.
1949. Aloysia gratissima var. revoluta (Moldenke)
Moldenke, Phytologia 9: 500. 1964. TYPE: Uruguay.
s. loc., s. coll. [no label, probably collected by J.
Arachavaleta s.n.] (holotype, MVM not seen; isotype,
NY [barcode] NY00103880 not seen, NY image!).
Iconography. Botta (1979: 106, fig. 13).
Distribution and habitat. Aloysia polystachya has
been observed to grow in central and northwestern
Argentina, and Paraguay, on sandy soils.
Discussion. Aloysia polystachya is an aromatic
plant, cultivated for medicinal properties (Arambarri
et al., 2009). Siedo (2010) lectotypified this taxon,
but this was superfluous, because the holotype has
been confirmed to exist and is housed at GOET.
Aloysia polystachya is easily distinguished from
other congeners since it has developed alternate
leaves to 5 cm long. Only one other species, A.
salsoloides, also has alternate leaves, but the leaf
blades are reduced, always shorter than 0.5 cm.
Selected specimens examined. ARGENTINA. Catamarca: Belén, cult., Arenas 257 (SI). Córdoba: Punilla, Sierra
Chica, Falda O, San Salvador, al N del Cerro Uritorco,
Quebrada de Ochoa, A. Hunziker 8951 (SI). Formosa:
˜ Las Lomitas, Barrio La Bomba, Arenas 3465 (SI). La
Patino,
´ Ulapes, Stuckert 17046 (SI).
Rioja: General San Martın,
Salta: Anta, Luna 147 (NY). San Juan: Valle Fertil, San
Agustı́n, Pedersen 11796 (NY, SI). BOLIVIA. Chuquisaca:
Azero, Estación Exp. zootecnica ‘‘El Salvador,’’ Krapovickas
31277 (SI). PARAGUAY. Central: Itá, Arenas 1916
(BACP). Pte. Hayes: Misión San Leonardo de Escalante,
June 1981, Sturzenegger s.n. (BACP 2453).
20. Aloysia pulchra (Briq.) Moldenke, Phytologia 1:
95. 1934. Basionym: Lippia pulchra Briq., Ark.
Bot. 2(10): 18. 1904. TYPE: Brazil. Rio Grande
do Sul: Porto Alegre, 1892, A. F. Regnell 579
(holotype, G [barcode] G00386457 not seen, G
image!; isotypes, F [bc] F0092929F not seen, F
image!, G [bc] G00386456 not seen, G image!,
NY [bc] NY00137807 not seen, NY image!, RB
not seen, SI!, S11-10470 not seen, S image!, SI
[bc] SI003403!, US [bc] US01049794 not seen,
US image!).
Lippia sellowii Briq., Annuaire Conserv. Jard. Bot. Genève
4: 21. 1900, replacement name for Lippia affinis Briq.,
Bull. Herb. Boissier 4: 339. 1896, nom. illeg. superfl.,
non Lippia affinis Schauer ex DC., Prod. [de Candolle]
11: 576. Aloysia uruguayensis Moldenke, Phytologia
Shrubs 1–3 m tall; stems glabrous. Leaves
opposite, with brief petioles 2–8 mm; blades elliptic
to obovate, 2–5 3 1–2 cm, apex obtuse, base acute,
attenuate, margin entire, sometimes slightly serrate
toward apex, coriaceous, adaxially scabrous, abaxially glabrate to slightly puberulous, venation pinnate, with lateral veins that converge in marginal
vein. Florescences axillary, solitary, lax, 5–8 cm;
peduncles 1–3 cm; flowers white; floral bracts ovate,
apex acute, 1–1.5 mm, lightly strigose, margin ciliate.
Flower with the calyx 2–3.5 mm, hispid in inferior
half, lightly hispid toward apex, with 4 teeth, brief,
unequal, triangular; corolla tube 4–5.5 mm, externally puberulous toward apex, with villous fauce.
Cluses 1 3 0.5 mm, glabrous.
Iconography. Botta (1979: 80, fig. 4).
Distribution and habitat. Aloysia pulchra grows
in the northeastern provinces of Argentina (Corrientes, Misiones), southern Brazil (Paraná, Rio
Grande do Sul), Paraguay, and Uruguay.
Discussion. Aloysia pulchra differs from A. gratissima in its habitat preference, with the latter found
in more xeric areas, and is a more xerophytic plant in
appearance. Morphologically, A. pulchra has elliptic
to obovate leaf blades, with entire margins and an
obtuse apex. While leaves range from elliptic to ovate
in A. gratissima, blade margins vary from entire to
dentate or serrate, with acute or subobtuse apices.
Both species are rather similar in general morphology, and subsequent studies are being performed in
order to check the validity of these two taxa.
When published, the name Lippia affinis Briq.
(Briquet, 1896) was superfluous and illegitimate
because of the priority of L. affinis Schauer (1847),
which affines with Lippia and not Aloysia. Briquet
(1900) later realized this homonymy and created L.
sellowii as a replacement name for the blocked L.
affinis Briq. However, Moldenke (1935) did not
realize Briquet had offered a replacement name and
created another replacement name, also superfluous,
as A. uruguayensis Moldenke.
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Aloysia gratissima var. sellowii (Briq.) Botta is here
considered a synonym of A. pulchra (Briq.) Moldenke,
following Botta’s (1979) observation about the close
relationship between these two taxa.
furrow on each blade lobe. However, A. riojana
differs from A. deserticola by its fruit with connate
commissural cluses. Plants are taller and more
graceful, to 1.7 m in A. riojana (vs. 1.5 m or less in
A. deserticola) with more slender, less spiny, and
longer branches.
Moldenke (1949: 106) mentioned a Berlin specimen identified in sched. as Acantholippia riojana by
Hieronymus, who was one of the two collectors of the
type. The Berlin sheet no longer exists and is the
reason why Múlgura (2012: 6) lectotypified this name
on a CORD specimen. Moldenke indicated in the
protologue that Hieronymus never published the
name, as in sched. and invalid, although Moldenke
acknowledged Hieronymus as the source for the later
validly published epithet, ascribing it as ‘‘Hieron. &
Moldenke.’’ The two could not have collaborated on
the epithet, given the difference in lifetimes (Hieronymus, 1846–1921; Moldenke, 1909–1996). The
epithet is better attributed as Hieron. ex Moldenke
(McNeill et al., 2012: Art. 46.5).
Selected specimens examined. ARGENTINA. Cor´
rientes: Capital, Riachuelo, Martınez
Crovetto 10801 (SI).
Misiones: Leandro Alem, Cerro Azul, Cabrera 28634 (SI);
San Javier, Arroyo Lorenzo, G. J. Schwarz 3803 (NY).
BRAZIL. Paraná: Palmeira, Papagaio, Smith 14930 (SI).
Rio Grande do Sul: entre Chapada y Lajeado Grande,
Brescia & Marchesi 4269 (SI). Santa Catarina: Isla do
Francés, Stienstra s.n. (SI). PARAGUAY. Alto Paraná: in
regione fluminis, Fiebrig 5904 (SI). Paraguarı́: Cerro
´ Zardini 10036 (SI). URUGUAY. Rivera: bajada
Mbatovı,
de Pena, Del Puerto 6053 (SI). Tacuarembó: Valle Edén,
Dematteis 1531 (SI).
21. Aloysia riojana (Hieron. ex Moldenke) Lu-Irving
& N. O’Leary, Syst. Bot. 39(2): 653. 2014.
Basionym: Acantholippia riojana Hieron. ex
Moldenke, Phytologia 3(3): 106. 1949. TYPE:
Argentina. La Rioja, Vinchina, 5 Mar. 1879, G.
Hieronymus & G. Niederlein 292 (lectotype,
designated by Múlgura et al. [2012: 6], CORD
[barcode] CORD00003809 not seen, CORD
image!;
isolectotypes,
CORD
[bc]
CORD00003810 not seen, CORD image!, G
[bc] G00366265 not seen, G image!).
Shrubs 1–1.7 m tall; branches spiny, stems hispid
to glabrate at maturity. Leaves small, opposite,
adpressed to the stem, sessile, squamiform, imbricate; blades rhomboidal in outline, 1.5 3 1.5–2 mm,
light green or yellow-green, 3-lobed with a large
apical lobe and 1 lateral smaller lobe on either side,
6 thickened texture, apex subobtuse, base rounded,
margin entire, revolute, adaxially scabrous, abaxially
with a conspicuous hirsute furrow on each blade lobe.
Florescences terminal, solitary, dense, 12–15 mm;
flowers lilac; floral bracts ovate or obovate, apex
acute or obtuse, 3–3.5 mm, slightly strigose. Flowers
with the calyx 3.5 mm, hispid, with 4 teeth, brief,
unequal, triangular; corolla tube 4–6 mm, with
villous fauce. Cluse 2–3 3 0.5 mm, orbicular in
cross section, commissural faces connate.
Iconography. Botta (1980: 520, fig. 2).
Distribution and habitat. Aloysia riojana is
endemic to Argentina (La Rioja, San Juan) where it
is frequent on sandy soils along riversides. Collections have been made at elevations up to 1800 m.s.m.
Discussion. Aloysia riojana is similar to A.
deserticola, both shrubs with spiny branches and
small, sessile leaves that are squamiform, adpressed
to stems with blades 3-lobed, with a conspicuous
Selected specimens examined. ARGENTINA. La Rioja:
´ Bermejo, c. Villa Unión,
Coronel Felipe Varela, cauce rıo
Biurrun 7705 (SI); Vinchina, en médano, Hunziker 2037
(SI). San Juan: Valle Fértil, ruta 510, cauce La Guardia, 2
km E de Baldecito, Biurrun 7696 (SI).
22. Aloysia salsoloides (Griseb.) Lu-Irving & N.
O’Leary, Syst. Bot. 39(2): 653. 2014. Basionym:
Acantholippia salsoloides Griseb., Abh. Königl.
Ges. Wiss. Göttingen 19: 244. 1874. Lippia
salsoloides (Griseb.) Briq., Nat. Pflanzenfam.
[Engler & Prantl] 4(3a): 152. 1897. TYPE:
Argentina. Catamarca: ‘‘ubi fruticeta praecipue
constituit in planitie Laguna blanca, alt.
10000,’’ P. G. Lorentz 457 (holotype, GOET
[barcode] GOET007269 not seen, GOET image!;
isotypes, [B , B as F neg. 17540!]; CORD [bc]
CORD00006133 not seen, CORD image!, SI
[bc] SI003359!).
Acantholippia hastulata Griseb., Symb. Fl. Argent., 279.
1879. Lippia hastulata (Griseb.) Hieron., Bol. Acad.
Nac. Ci. 4: 407. 1882. TYPE: Argentina, P. G. Lorentz
& G. H. Hieronymus 713 (holotype, CORD [barcode]
CORD00006139 not seen, CORD image!; isotypes,
[B , B as F neg. 17511!]; GOET [bc] GOET008868
not seen, GOET image!, GOET [bc] GOET008867 not
seen, GOET image!, SI [bc] SI003357!).
Shrubs 0.35–1.5 m tall; branches spiny; stems
hispid, glabrate at maturity. Leaves alternate, adpressed to stem, sessile; blades reduced, ovate, 1.5–
4.5 3 2 mm, 5-lobed with 1 large apical and 2 lateral
small lobes on either side, 6 thickened texture, apex
subobtuse base rounded, margin entire, revolute,
adaxially scabrous, abaxially hirsute, venation prom-
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inent, reticulate. Florescences terminal, solitary,
dense, 10–15 mm; flowers white; floral bracts ovate
or obovate, apex acute or obtuse, 3–4 mm, slightly
strigose. Flower with the calyx 3.5–4 mm, hispid,
with 4 brief teeth, unequal, triangular; corolla tube 5–
7 mm, with villous fauce. Cluse 2–3 3 0.5 mm,
glabrous.
hispid. Flower with the calyx 2–3.5 mm, hispid, with
underlayer of subsessile glandular trichomes, with 4
teeth, unequal, triangular; corolla tube 3–3.5 mm,
externally and internally glabrous. Cluses 1.5 3 1
mm, glabrous.
Iconography. Troncoso (1974: 347, fig. 12, k–n);
Botta (1980: 524, fig. 4); Caro (1982: 9, fig. 1 and 13,
fig. 2, sub Aloysia hastulata).
Distribution and habitat. Aloysia salsoloides
grows in northwestern Argentina and southern
Bolivia, noted as found in salty soils and at elevations
to 3200 m.
Discussion. Aloysia salsoloides shares with A.
deserticola, A. riojana, and A. tarapacana a shrubby
habit with spiny branches and reduced leaf blades,
with the leaves sessile and adpressed to stems.
However, A. salsoloides is distinguished from these
three by the alternate arrangement of its leaves that
are 5-lobed and not squamiform nor imbricate.
Leaves are opposite, squamiform, and densely
imbricate, with blades either entire or 3-lobed in A.
deserticola, A. riojana, and A. tarapacana.
Selected specimens examined. ARGENTINA. Catamarca: Belén, Laguna Blanca, Cabrera 32474 (SI). Jujuy:
Humahuaca, Zuloaga 9170 (SI). Salta: Rosario de Lerma,
57 km pasando Pte. Integración Argentina-Chilena,
Cialdella 407 (SI). BOLIVIA. Potosı́: Uyuni, Hicken 17
(SI).
23. Aloysia salviifolia (Hook. & Arn.) Moldenke,
Lilloa 5: 381. 1940. Basionym: Verbena salviifolia Hook. & Arn., Bot. Beechey Voy. 1: 42.
1830, as ‘‘salviaefolia.’’ Lippia chilensis Schauer, Prodr. [de Candolle] 11: 573. 1847,
replacement name, non Lippia salviifolia Cham.,
Linnaea 7: 227. 1832. Aloysia chilensis (Schauer) Moldenke, Revista Sudamer. Bot. 4: 15.
1937. TYPE: Chile. ‘‘Hab. Coquimbo,’’ s.d., s.
coll. (neotype, designated here, K [barcode]
K000470996 not seen, K image!). Figure 11.
Shrubs 1–2 m tall; stems glabrous. Leaves
opposite, rarely alternate, sessile, elliptic, 1–2( 4)
3 1( 1.5) cm, apex acute to subobtuse, base obtuse
to subtruncate, margin basally entire, irregularly
serrate toward apex or from mid-blade distally,
abaxial and adaxial surfaces strigose, venation
pinnate, reticulate, abaxially prominent. Florescences
axillary, solitary, dense, 5–10 cm; peduncles 1–2 cm;
flowers white; pedicels 0.5 mm; floral bracts widely
ovate, large, 4–4.5 mm, apex acute, acuminate,
Distribution and habitat. Aloysia salviifolia is
found in Chile (Regions III, IV), and one collection
exists from Argentina.
Discussion. Aloysia salviifolia is distinguished by
its leaf blade base obtuse to subtruncate, and margin
basally entire and irregularly serrate toward apex or
from mid-blade distally, and venation pinnate,
reticulate, and abaxially prominent. It is also
differentiated by its large floral bracts, to 4–4.5
mm, surpassing the calyx, a feature not frequent in
the rest of Aloysia.
Schauer (1847) considered that Verbena salviifolia
Hook. & Arn. should be under Lippia, but the
epithet’s use in Lippia was blocked by the prior name
L. salviifolia Cham., so that Schauer proposed L.
chilensis as the replacement name. When Moldenke
(1940) later transferred V. salviifolia to Aloysia, the
original epithet was not blocked by priority and was
available for use within Aloysia.
Noltie (2010: 181) stated that no original material
has been found from Verbena salviifolia at the E and
K herbaria. Noltie mentioned that ‘‘Arnott’s MS
description has been mistakenly attached to a sheet
annotated ‘Lippia chilensis Schauer,’ bearing mixed
specimens of Lobb 453 (K [barcode] K000470996)
and a Gillies s.n. collection from Mendoza (K
[barcode] K000470995), neither of which are not
original material.’’ Consequently, a neotype is here
selected, with the more representative Lobb 453
material from Chile here chosen.
Selected specimens examined. ARGENTINA. Mendoza:
s. loc., Gillies s.n. (K). CHILE. s. loc., Bridges 1346 (SI).
Region III: Atacama, San Felix, Ricardi 23967 (SI).
Region IV: Coquimbo, Quebrada de Rivadavia, Werdermann 103 (SI); Paihuano, Pfister 8318 (SI).
24. Aloysia scorodonioides (Kunth) Cham., Linnaea
7: 234. 1832. Basionym: Lippia scorodonioides
Kunth, Nov. Gen. Sp. [quarto ed.] 2: 269. 1818.
TYPE: Ecuador. ‘‘in regno Quitensis,’’ s.d., A.
Bonpland 2192 (holotype, P [barcode]
P00307136 not seen, P image; isotype, SI [bc]
SI003396!). Figure 12.
Shrubs 0.5–3( 4) m tall; stems canescent when
young, puberulous to strigose on mature stems.
Leaves opposite, petioles 0.5–1 cm, sometimes
sessile; blades elliptic, ovate, or orbicular, 1–5( 7)
3 1–3( 4) cm, apex obtuse to subobtuse, base
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Figure 11. Aloysia salviifolia (Hook. & Arn.) Moldenke. —A. Floriferous branch with axillary florescences (homothetic
pleiobotrya). —B. Entire leaf, abaxial surface, margin basally entire, irregularly serrate toward apex, prominent pinnate,
reticulate venation. —C. Intact flower with hispid floral bract subtending, calyx hispid, with underlayer of subsessile glandular
trichomes. —D. Corolla dissected open to reveal the androecium and gynoecium. —E. Close-up of abaxial leaf surface, with
strigose and scabrid indument. —F. Close-up of adaxial leaf surface, with strigose indument. From Bridges 1346 (SI).
acute, cordate, or rounded, margin evenly crenate,
rugose and creased texture, sometimes membranaceous, adaxially scabrous, with conspicuous reticulate venation, abaxially hispid or tomentose. Florescences axillary, solitary, lax or dense, 1–9( 12) cm;
peduncles 1–4 cm; flowers white, purple, or pink,
small; floral bracts linear to narrowly ovate,
subulate, 1–3 mm, strigose or scabrous. Flower with
the calyx 2–3 mm, densely hispid, sometimes with
long hairs in lower half, grading to shorter hairs
toward apex, with 4 unequal teeth; corolla tube 4.5–
6 mm, externally puberulous, with villous fauce;
ovary glabrous, sometimes slightly pilose. Cluses 2
3 1 mm, glabrate.
TAXONOMIC KEY TO THE VARIETIES OF ALOYSIA SCORODONIOIDES
Shrubs 0.5–2 m tall; leaves 1–2.5( 4) 3 1–
1.5( 2) cm, blades elliptic to ovate; florescences
dense, 1–3( 4) cm long, straight; peduncles 1–3
cm long; corollas 4.5–5.5 mm long . . . . . . . . . . . .
. . . . . . . . . . 24a. A. scorodonioides (Kunth) Cham. var.
hypoleuca (Briq.) Moldenke
1 0 . Shrubs 2–3( 4) m tall; leaves 2–5( 7) 3 2–
3( 4), blades ovate to orbicular; florescences lax
or dense, 4.5–12 cm long, curved, peduncles 2–4
cm long; corollas 5–6 mm long . . . . . . . . . . . . . . . . . . . 2
1.
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2.
Florescences lax, 8–12 cm long, peduncles
2–4 cm long . . . . . . . . . . . . 24b. A. scorodonioides
(Kunth) Cham. var. mathewsii (Briq.) Moldenke
2 0 . Florescences dense, 4.5–8 cm long, peduncles 2 cm long . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . 24c. A. scorodonioides var. scorodonioides
24a. Aloysia scorodonioides var. hypoleuca
(Briq.) Moldenke, Phytologia 36(5): 437. 1977.
Basionym: Lippia scorodonioides var. hypoleuca
Briq., Bull. Herb. Boissier 4: 338. 1896. TYPE:
Peru. s. loc., s.d., J. Dombey 259 (lectotype,
designated by Siedo [2010: 203], G [barcode]
0386453 not seen, G image!, G as F neg.
24668!). Figure 12J–L.
Aloysia scorodonioides var. parvifolia Moldenke, Phytologia
36(5): 437. 1977. TYPE: Bolivia. Near La Paz, Oct.
1885, H. H. Rusby 920 (holotype, NY [barcode]
NY00103890 not seen, NY image!; isotypes, BM [bc]
BM000643654 not seen, BM image!, MO-116725 not
seen, MO image!, NY [2, bc] 103888, 103889, not
seen, NY images!, US [2, bc] 01013795, 1323004 not
seen, US images!, WIS [bc] WIS0256108 not seen,
WIS image!).
Aloysia depressa Ravenna, Onira 11(4): 15. 2007, syn. nov.
TYPE: Bolivia. La Paz, Feb.–Mar. 1933, M. DoelloJurado s.n. (holotype, BA-9827!).
Aloysia axillaris J. R. I. Wood, Kew Bull. 64: 521. 2009,
˜ del
syn. nov. TYPE: Bolivia. Potosı́, Torotoro, Canon
Vergel, 2591 m, 3 Jan. 2000, J. R. I. Wood, M.
˜ 21305 (holotype, K [2, barcode]
Mercado & T. Ortuno
K000738254 not seen, K image!, K [barcode]
K000738253 not seen, K image!; isotypes, BOLV
not seen, LPB [bc] LPB0000838 not seen, LPB
image!).
Shrubs 0.5–2 m tall; stems puberulous. Leaves 1–
2.5( 4) 3 1–1.5( 2) cm; blades elliptic or ovate.
Florescences dense, 1–3( 4) cm, straight; peduncles
1–3 cm. Flowers pink to purple, corolla tubes 4.5–5.5
mm.
Distribution and habitat. Aloysia scorodonioides
var. hypoleuca has a narrow distribution being found
only in Peru and Bolivia growing on open, dry
hillsides in loose gravel and shallow soils, at
elevations to 3070 m.
Discussion. In Cuzco, Peru, near Calca, there are
some specimens with lax florescences with flowers
spaced apart 3 mm, e.g., Olmstead 2009-40 (WTU),
Vargas 160 (SI), and Cook 247 (US).
In his 2009 protologue, Wood stated that
Aloysia axillaris was distinguished by two different
inflorescence forms, one with solitary axillary
flowers without floral bracts and the second with
axillary spikes typical for the genus. These plants
were observed to be restricted to two areas in
Bolivia, from Potosı́ and Cochabamba. The author
noted the possibility of a hybrid origin for A.
axillaris. The analysis of the type material and the
description of the plant indicate that this corresponds to A. scorodonioides var. hypoleuca, with
both sharing small leaf blades, shorter than 4 cm,
with elliptic to ovate shapes. Aloysia axillaris
differs principally in the presence of solitary
flowers that lack floral bracts; this may be a
localized somatic mutation in two known populations in Bolivia, which is here interpreted as of
teratological or hybrid origin.
The study of the type material of Aloysia depressa
Ravenna indicates that this is the same as A.
scorodonioides var. hypoleuca. Both have the distinctive leaves of this variety (small elliptic to ovate
blades) and dense florescences shorter than 4 cm.
Selected specimens examined. BOLIVIA. Cochabamba:
˜ Wood 20173 (BOLV, K, LPB). La
Quillacolo, La Cabana,
Paz: Murillo, Mecapaca, Beck 3530 (SI). PERU. Cajamarca: Obrajillo, 1838–1842, Wilkes s.n. (US). Cuzco: Calca, 2
km from Pisac, Olmstead 2009-40 (WTU). Huancavelica:
Tayacaja, entre Izcuchaca y Acostambo, Hutchison 4201
(US). Lima: Huarochirı́, c. Matucana, Ferreyra 7021 (SI);
Matucana, Macbride 133 (US); Km. 86, Lima–Oraya,
Ferreyra 7013 (NY, SI).
24b. Aloysia scorodonioides var. mathewsii (Briq.)
Moldenke, Phytologia 1: 95. 1934. Basionym:
Lippia scorodonioides var. mathewsii Briq., Bull.
Herb. Boissier 4: 339. 1896. TYPE: Peru. s.
loc., W. Mathews 3160 (holotype, G [barcode]
G00386446 not seen, G image! as F neg.
24669!; isotypes, G [bc] G0386464 not seen,
G image!, K [bc] K000545990 not seen, K
image!). Figure 12M.
Aloysia scorodonioides var. lopez-palacii Moldenke, Phytologia 36(5): 437. 1977. TYPE: Ecuador. Pichincha:
Quito, 4 Feb. 1977, S. López-Palacios 4249 (holotype,
TEX-LL [barcode] LL00374940 not seen, TEX-LL
image!).
Shrubs 2–3( 4) m tall, stems canescent when
young, strigose at maturity. Leaves 2–5( 7) 3 2–
3( 4) cm, blades ovate to orbicular, with rugose
texture. Florescences lax, long, flowers not densely
disposed, 8–12 cm, curved; peduncles 2–4 cm.
Flowers white or pink, with corolla tubes 5–6 cm.
Distribution and habitat. Aloysia scorodonioides
var. mathewsii grows in Ecuador, Peru, Bolivia, and
northern Argentina. This variety has been collected
from elevations of 900–2600 m.
Discussion. Aloysia scorodonioides var. mathewsii is
distinguished by its long florescences, 8–12 cm, with
flowers laxly disposed. This contrasts with the dense
florescences 4.5–8 cm long in the autonymic variety,
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and the shorter florescences less than 4 cm in A.
scorodonioides var. hypoleuca.
sheet from F has a label, by Moldenke, that reads:
‘‘Aloysia leptophylla Loes.’’ In the protologue,
Moldenke (1941: 11) attributed the name as ‘‘Loes.
& Moldenke.’’ Given the differences in life spans
(Loesener, 1865–1941; Moldenke, 1909–1996), it is
unlikely Moldenke collaborated with Loesener, but
rather credited Loesener for the unpublished name
(nom. ined.). There is no further mention of Loesener
by Moldenke and the ascription of the plant name
should be Loes. ex Moldenke (McNeill et al., 2012:
Art. 46.5).
Selected specimens examined. ARGENTINA. Chaco:
Quitilipi, borde bosque, Schulz 2985 (SI). Jujuy: Santa
Barbara, ruta Prov. 6, de Santa Clara a Abra de los
Morteros, 12 km de Santa Clara, Zuloaga 11490 (SI). Salta:
Anta, San Javier, Saravia Toledo 1764 (SI). BOLIVIA.
Chuquisaca: Luis Calvo, Boyuibe, Beck 9428 (SI).
ECUADOR. Pichincha: Tumbaco, Asplund 6533 (US).
´
PERU. Apurimac:
Abarcay, Vargas 594 (SI). Cajamarca:
Jaén, Km. 127, rd. Bagua to Pucara, Lu-Irving 9-62 (SI,
WTU); Pucará, 127 km E Olmos, Hutchison 3520 (US).
Cuzco: Calca, Cook 247 (US).
24c. Aloysia scorodonioides (Kunth) Cham. var.
scorodonioides. Figure 12A–I.
Lippia scorodonioides var. detonsa Briq., Bull. Herb.
Boissier 4: 339. 1896. Aloysia scorodonioides var.
detonsa (Briq.) Moldenke, Phytologia 1: 95. 1934.
TYPE: Colombia. ‘‘In montibus Columbiae,’’ s.d., K.
T. Hartweg 1349 (holotype, BR [barcode]
BR000000550590 not seen, BR image!; isotype,
OXF not seen).
Aloysia leptophylla Loes ex Moldenke, Phytologia 2: 11.
1941, syn. nov. TYPE: Peru. s. loc., 1909–1914, A.
Weberbauer 5374 (holotype, F [barcode] V0042839F!
not seen, F image!; isotypes, NY [bc] NY00103878
not seen, NY image!, SI!).
Aloysia scorodonioides var. orbicularis Moldenke, Phytologia
3: 406. 1951. TYPE: Colombia. Yuaco, near Pasto,
˜ s.d., G. K. Wilhelm Hermann Karsten s.n.
Narino,
(holotype, W [barcode] W0032435 not seen, W
image!).
Aloysia boliviensis Moldenke, Phytologia 53(7): 460. 1983.
TYPE: Bolivia. La Paz, Murillo, Mecapaca, 28 Mar.
1982, J. C. Solomon 7410 (holotype, TEX-LL
[barcode] LL00374935 not seen, TEX-LL image!;
isotypes, LPB [bc] LPB0000841 not seen, LPB
image!, MO-3006114 not seen, MO image!, SI [bc]
SI003375!, U [bc] U0006999 not seen, U image!).
Shrubs 2–3( 4) m tall, stems canescent when
young, strigose at maturity. Leaves 2–5( 7) 3 2–
3( 4) cm, blades ovate to orbicular, rugose texture.
Florescences dense, 4.5–8 cm, curved; peduncles 2
cm long. Flowers white or pink, corolla tubes 5–6 cm.
Distribution and habitat. Aloysia scorodonioides
var. scorodonioides is found in Colombia, Ecuador,
Peru, Bolivia, Paraguay, and northern Argentina. It
has been collected from dry, brushy slopes, from open
xeric, calcareous soils, and at elevations of 800–2800
m. Flowers are said to be very fragrant.
Discussion. The study of the type material of
Aloysia leptophylla shows that this species corresponds to A. scorodonioides var. scorodonioides,
sharing dense florescences 4.5–8 cm long.
There has been confusion about the attribution of
the author of Aloysia leptophylla, since the holotype
Selected specimens examined. ARGENTINA. Jujuy: El
Carmen, Pampa Blanca, Cabrera 29961 (SI). Salta: Anta,
sector II, Saravia Toledo 1591 (SI). Tucumán: Burruyacu,
˜
Canada
Alegre, Stuckert 21287 (SI); s. loc., Venturi 849 (SI).
BOLIVIA. La Paz: Loayasa, Beck 6039 (SI); Illimani, Nov.
1911, Buchtien 3240 (NY, SI). Santa Cruz: Cordillera,
Charagua, Cabrera 33665 (SI). ECUADOR. Esmeraldas:
Lita–San Lorenzo rd., Gentry 70190 (MO). Loja: near
Saraguro, Hart 1481 (US). Pichincha: Guaillabamba valley
at Quito, Haught 3155 (US). Tungurahua: Ambato,
Pachano 120 (US). PARAGUAY. s. loc., 1896, Drake s.n.
˜
(SI). PERU. Cajamarca: Celendin, valley rio Maranon,
LuIrving 9-32 (SI, WTU). Cuzco: Anta, Vargas 3692 (SI).
Lima: along rio Chillón, Pennell 14438 (US).
25. Aloysia tarapacana (Botta) Lu-Irving & N.
O’Leary. Syst. Bot. 39(2): 653. 2014. Basionym:
Acantholippia tarapacana Botta, Hickenia
´
1(35): 197. 1979. TYPE: Chile. Tarapaca,
Arica, Puquios, 3750 m.s.m., 16 Sep. 1955,
M. Ricardi 3363 (holotype, SI [barcode]
SI003356!; isotypes, CONC not seen, LP [bc]
LP006687 not seen, LP image!).
Shrubs 0.5–1 m tall, stems cylindrical, with spiny
branches, hispid when young, glabrate at maturity.
Leaves opposite, adpressed to the stem, sessile,
squamiform, imbricate; blades reduced, rhomboidal,
2 3 1 mm, dark green, texture 6 thickened, apex
subobtuse, base rounded, margin entire, revolute,
adaxially scabrous, abaxially hirsute. Florescences
terminal, solitary, dense, 6–12 mm; flowers blue;
floral bracts ovate or elliptic, apex acute or obtuse,
4.5 mm, slightly strigose. Flower with the calyx 4
mm, hispid, with 4 brief teeth, unequal, triangular;
corolla tube 5–6 mm, with villous fauce. Cluse 2 3
0.5 mm.
Iconography. Botta (1980: 522, fig. 3).
Distribution and habitat. Aloysia tarapacana is
endemic to Chile (Region I), collected at elevations
from 3000 to 3390 m.
Discussion. Aloysia tarapacana is similar to A.
deserticola and A. riojana, with a shrubby habit and
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Figure 12. Aloysia scorodonioides (Kunth) Cham. A–I. Aloysia scorodonioides var. scorodonioides. —A. Floriferous branch
with axillary florescences (homothetic pleiobotrya). —B. Basal leaf, orbicular blade. —C. Apical leaf, ovate blade. —D. Intact
flower with hispid calyx. —E. Corolla dissected open to reveal the androecium and gynoecium. —F. Dissected calyx and floral
bract, scheme. —G. Close-up of abaxial leaf surface, with hispid indument. —H. Close-up of adaxial leaf surface, with scabrid
indument. —I. Cluse, lateral view. J–L. Aloysia scorodonioides var. hypoleuca (Briq.) Moldenke. —J. Floriferous branch with
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spiny branches. In all three species, the leaf blades
are in opposite position, reduced and squamiform,
and densely imbricate along the stems. Aloysia
tarapacana may be distinguished by its leaf blades
with entire margins and dark green in color. Leaves
are 3-lobed, with a conspicuous furrow on each blade
lobe, and light green or yellow in color in both A.
deserticola and A. riojana.
Distribution and habitat. Aloysia trifida is known
from Chile (Regions III and IV) and also grows in
Argentina (La Rioja, San Juan). Carmona and
´ r (1995) established that leaf anatomical
Ancıbo
characters indicated that this species was probably
a facultative halophyte. This species has been
collected from elevations up to 2500 m.
Selected specimens examined. CHILE. Region I: Arica,
Antes de Zapahuina, H. Escobar 233 (SI); Parinacota entre
Zapahuira y Putre, Cocucci et al. 3277 (SI).
26. Aloysia trifida (Gay) Lu-Irving & N. O’Leary.
Syst. Bot. 39(2): 653. 2014. Basionym: Lippia
trifida Gay, Fl. Chil. 5: 29. 1849. Acantholippia
trifida (Gay) Moldenke, Lilloa 5(2): 371. 1940.
TYPE: Chile. Copiapó, Feb. 1843, C. Gay s.n.
(holotype, G [barcode] G00366065 not seen, G
image!).
Lippia fonckii Phil., Anales Univ. Chile 90: 620. 1895, syn.
nov. Aloysia fonckii (Phil.) Moldenke, Phytologia 2:
50. 1941. TYPE: Chile. Coquimbo, ‘‘prope La
Higuera in litorali prov. Coquimbo,’’ s.d., F. S. Fonck
s.n. (holotype, SGO not seen, SGO image!; isotypes,
SGO not seen, image!, SI [barcode] SI003514!).
Aloysia reichei Moldenke, Lilloa 5: 380. 1940, as ‘‘reichii,’’
syn. nov. TYPE: Chile. Huanta, cordillera de
Coquimbo, Jan. 1904, K. F. Reiche 19 (holotype,
SGO-4203 not seen, SGO-4203 image!; isotype, SI
[barcode] SI003393!).
Aloysia reichei var. trilobata Moldenke, Phytologia 2: 309.
1947. TYPE: Chile. Coquimbo, Elqui, Rio Turbio, 19
Oct. 1940, R. Wagenknecht 4238 (holotype, NY
[barcode] NY103886 not seen, NY image!; isotypes,
SGO-4204 not seen, SGO-image!, SI [bc] SI003408!).
Shrubs 1–2 m tall; stems cylindrical, hispid when
young, glabrate at maturity. Leaves opposite, sessile,
blade entire, ovate to elliptic, or 3-parted, 3.5–4.5 3
2–3.5 mm, somewhat thickened texture, apex acute to
obtuse, base attenuate, margin entire, scabrous on
both surfaces. Florescences axillary, solitary, dense,
1.5–3 cm; flowers white; floral bracts ovate, apex
acute or obtuse, 3–3.5 mm, slightly strigose. Flowers
with the calyx 2–2.5 mm, hispid, with 4 teeth, brief,
unequal, triangular; corolla tube 4–5 mm, with
villous fauce; superior pair of stamens with glandular
appendices to anther connectives. Cluse 2–3 3 0.5
mm.
Iconography. Botta (1980: 518, fig. 6); Caro (1982:
27, fig. 5).
Discussion. Aloysia trifida is the only taxon in
Aloysia known to have glandular appendices on the
anther connectives of the superior pair of stamens.
This species is also distinguished by its leaf blades
entire, ovate to elliptic, but sometimes 3-parted.
The type material examined for Aloysia reichei
has 3-parted leaves, as well as the distinctive
glandular appendices on the anther connectives of
the upper stamen pair. These are both characters
exclusively observed in A. trifida and why A.
reichei is synonymized to the species. The type
specimen of Lippia fonckii also shares these
characters, and this species name is also synonymized to A. trifida.
Selected specimens examined. ARGENTINA. La Rioja:
Coronel Felipe Varela, ruta 40, entre Villa Unión y
˜
Sanogasta,
5 km E Puerto Alegre, Biurrun et al. 7706
˜ Kiesling et al.
(SI). San Juan: Calingasta, camp. Castano,
9130 (SI). CHILE. Region III: Atacama, Copiapó, ruta a
San Francisco, 61 km E interseccion ruta Paipote y Inca de
Oro, Taylor & Pool 11607 (MO, SI). Region IV: Coquimbo,
La Laguna, Jiles 5079 (SI).
27. Aloysia velutina Siedo, Lundellia 15: 44, fig. 4.
2012. TYPE. Peru. Cajamarca: Mpio. Cajamarca, sobre el km 156 de la carr. Pacasmayo–
Cajamarca, bosque espinoso, 2000 m, 5 Apr.
1982, I. Sanchez Vega 2763 (holotype, F
[barcode] F0093715F not seen, F image!;
isotypes, MO not seen, SI [bc] SI080019!).
Figure 13.
Shrubs 1–1.5 m tall; stems glabrous. Leaves
opposite, with an occasional third leaf per node and
ternate, petioles 0.3–1 cm long; blades ovate to
elliptic, 3–6 3 1–5 cm, apex acute to subobtuse, base
rounded to truncate, margin slightly crenate along
entire blade, adaxially velutinous, abaxially incanous. Florescences terminal and axillary, dense, 8–
15( 25) cm; peduncles 1–3 cm; flowers violet or
cream-white, sometimes with lavender tubes and
white lobes; pedicels 0.5 mm; floral bracts ovate,
axillary florescences (homothetic pleiobotrya). —K. Leaf, elliptic blade. —L. Intact flower with strigose floral bract subtending.
M. Aloysia scorodonioides var. matthewsii (Briq.) Moldenke. —M. Floriferous branch with axillary florescences (homothetic
pleiobotrya). A–F, I are illustrated from Venturi 849 (SI); G, H, from Haught 3155 (US); J–L, from Macbride 133 (US); M, from
Saravia Toledo 1764 (SI).
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Figure 13. Aloysia velutina Siedo. —A. Floriferous branch, with both axillary and terminal florescences (heterothetic
pleiobotrya). —B. Flower with hispid calyx and floral bract. —C. Leaf adaxial surface. —D. Close-up of abaxial leaf surface,
with incanous indument. —E. Close-up of adaxial leaf surface, with scabrid indument. A, B from Sagástegui 14147 (SI); C–E
from Llatas Quiroz 1953 (SI).
small, 2–2.5 mm, apex acute, acuminate, strigose, or
hispid. Flower with the calyx 2–2.5 mm, hispid, with
4 teeth, unequal, triangular; corolla tube 3–3.5 mm,
externally puberulous, with villous fauce. Cluses 1.5
3 1 mm, glabrous.
Distribution and habitat. Aloysia velutina is
endemic and known only from woodland habitats in
Cajamarca, Peru.
Discusssion. Aloysia velutina has violet or creamwhite flowers, sometimes with lavender tubes and
white lobes, and sometimes three leaves per node
(e.g., Olmstead 2009-8, WTU). The species is quite
similar in leaf and florescence appearance to A.
scorodonioides var. scorodonioides. However, A.
velutina has both axillary and terminal florescences,
whereas florescences appear only axillary in A.
scorodonioides, even if this trait is not always easy
to observe. The inflorescence morphology in A.
velutina (presence of both terminal and axillary
florescences) groups A. velutina with A. arequipensis,
A. citrodora, A. fiebrigii, and A. herrerae. However, A.
velutina is easily distinguished by its opposite leaves
that are ovate to elliptic, which contrasts with the
ternate leaves of A. citrodora and the verticillate
leaves, in whorls of three, with narrower blades
(linear to elliptic) for A. fiebrigii. The shallowly
crenate margins of the leaves of A. velutina easily
distinguish this species from A. herrerae and A.
arequipensis. Leaves seem longer in A. velutina too
(3–6 3 1–5 cm) versus 1–2 3 0.5–1.2 cm in A.
arequipensis and 2( 5.5) 3 0.4( 1.5) cm in A.
herrerae. Leaf margins are entire in the former, and
margins are finely serrate in the apical two thirds to
one half of the blade in the latter.
Selected specimens examined. PERU. Cajamarca: Contumazá, ca. 2 km from Contumazá, Dillon 4544 (SI). Cuzco:
Anta, Canyon rio Apurimac, 0.5 km from Pte. Cunyac,
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Olmstead 2009-8 (WTU). La Libertad: Otuzco, alrededor
San Ignacio, Sagástegui 14147 (SI). Lambayeque: Ferreafe, Incahuasi, Llatas Quiroz 1953 (SI).
10 .
28. Aloysia virgata (Ruiz & Pav.) Pers., Syn. Pl. 2(1):
139. 1806. Basionym: Verbena virgata Ruiz &
Pav., Fl. Peruv. 1: 20. 1798. Aloysia virgata (Ruiz
& Pav.) Juss., Ann. Mus. Natl. Hist. Nat. 7: 73.
1806, nom. inval. Zappania virgata (Ruiz & Pav.)
Poir., Encycl. [J. Lamarck et al.] 8: 845. 1808, as
‘‘Zapania virgata.’’ Priva virgata (Ruiz & Pav.)
Spreng., Syst. Veg. [Sprengel] 2: 753. 1825. TYPE:
[Peru. Pasco:] ‘‘Pozuzo,’’ 1827, J. A. Pavón s.n.
[Herb. Pavón 36] (holotype, P [barcode]
P000713759 not seen, P image; isotypes, G [bc]
G00236936 not seen, G image!, G [bc] G00236923
not seen, G image, G [bc] G00386451 not seen, G
image!, G [bc] G00386450 not seen, G image!,
MPU [bc] MPU011501 not seen, MPU image!, P
[bc] P000713760 not seen, P image!, SI [P fragm.,
photo]!).
Shrubs 2–7 m tall, stems hirsute, glabrate at
maturity. Leaves opposite; petioles 3–10 mm; blades
elliptic to ovate, 4–9( 15) 3 1.5–4 cm, apex acute to
subobtuse, base acute, subobtuse, or truncate, margin
evenly minutely serrate or crenate, sometimes almost
entire in appearance, membranaceous to coriaceous,
adaxially strigose, abaxially hirsute, with prominent
venation. Florescences axillary, solitary or 2 to 5,
sometimes 7, per leaf axil, lax, subpendulous, 10–20
cm; peduncles 1–3 cm, sometimes branched; flowers
white or cream, small; floral bracts linear to narrowly
elliptic, 2–3 mm, hirsute. Flower with the calyx 2.5–
3.5 mm, densely hispid, with long hairs and
subsessile glandular trichomes, with 4 teeth, unequal, acute, subulate, the teeth equaling or
exceeding the calyx tube in length; corolla tube
2.3–3.5 mm, externally puberulous toward apex, with
villous fauce. Cluses 1 3 0.5 mm, glabrate.
Iconography. Botta (1979: 97, fig. 9).
Discussion. Leaves of Aloysia virgata are said to be
used for therapeutical matters, such as antifungal
treatments (Arambarri et al., 2008, 2009).
Aloysia virgata (Ruiz & Pav.) Juss. is an invalid
combination (McNeill et al., 2012: Art. 33.1); Jussieu
assigned Verbena virgata Ruiz & Pav. to Aloysia, but
did not make the formal combination.
TAXONOMIC KEY
1.
TO THE
VARIETIES
OF
ALOYSIA
VIRGATA
Leaf apex acute, blades with acute to subobtuse
base, high blade length/width ratios; ovary
glabrous . . . . . . . . . . . . . 28b. A. virgata (Ruiz & Pav.)
Juss. var. virgata
Leaf apex subobtuse, blades with obtuse to
truncate base, low blade length/width ratios;
ovary pilose . . . . . . . . . 28a. A. virgata (Ruiz & Pav.)
Juss. var. platyphylla (Briq.) Moldenke
28a. Aloysia virgata (Ruiz & Pav.) Pers. var. platyphylla (Briq.) Moldenke, Phytologia 2: 408. 1948.
Basionym: Lippia virgata var. platyphylla Briq.,
Annuaire Conserv. Jard. Bot. Genève 7–8: 304.
1904. TYPE: Paraguay. Paraguarı́, Mar. 1881–
1884, B. Balansa 3116 (lectotype, designated by
Siedo [2010: 203], F-876782!; isolectotypes, BM
[barcode] BM000098764 not seen, BM image!,
G [bc] G00166263 not seen, G image!,
GH [bc] GH00312629 not seen GH image!,
K [bc] K000471000 not seen, K image!, K [bc]
K000487001notseen,Kimage!,P [bc]P02851828
not seen, P image!, SI [bc] SI003609!).
Lippia virgata var. elliptica Briq., Annuaire Conserv. Jard.
Bot. Genève 7–8: 304. 1904. Aloysia virgata var.
elliptica (Briq.) Moldenke, Phytologia 1: 441. 1940.
TYPE: Paraguay. L’Assomption, 15 abr. 1874, B.
Balansa 1016 p.p. (holotype, G [barcode] G00166264
not seen, G image!; isotypes, G [bc] G00381090 not
seen, G image!, G [bc] G00381089 not seen, G
image!, BR [bc] BR0000005505258 not seen, BR
image!, P [bc] P02851832 not seen, P image!, SI!,
S11-10533 not seen, S image!).
Aloysia naviculata Ravenna, Onira 11(4): 16–17. 2007.
TYPE: Paraguay. Dpto. Cordillera: Colonia Ojopoı´ E
de Piribebuy, 3 June 1985, P. Arenas 2912 (holotype,
BA-90077!).
Shrubs with ovate leaf blades, almost the same
length as width, with subobtuse apex, and obtuse to
truncate base. Pilose ovary.
Distribution and habitat. Aloysia virgata var.
platyphylla grows in Bolivia, Paraguay, and Argentina. In this last country, this variety exhibits a wider
distribution than the type variety. Aloysia virgata var.
platyphylla has been collected from woods and
thickets on dry or wet ground, in forest patches or
xerophytic areas.
Discussion. In the new combination Aloysia virgata
var. elliptica (Briq.) Moldenke, Moldenke repeated
the same taxonomic action, not once but four times
(Moldenke, 1940: 441; 1942: 310; 1947: 363; 1948:
408), with the last three names constituting later
isonyms, which are disregarded by the Code (McNeill
et al., 2012: Art. 6, Note 2). To add confusion to this,
Moldenke (1942: 310) committed a typographic error
between varieties platyphylla and elliptica, which he
later corrected (1947: 363). There are three sheets at
G collected by Balansa, number 1016, with a
collection date of 1874, from Paraguay, that correspond to A. virgata var. elliptica. There is no
604
Annals of the
Missouri Botanical Garden
indication of pro parte on these three sheets, although
the pro parte designation could be traced to three
additional sheets at G for the collection Balansa
1016, each noted as ‘‘Isotypus, Malabaila carvifolia
Boiss. & Balansa,’’ with ‘‘validated determination’’ to
Peucedanum palimbioides Boiss., Apiaceae, from
Turkey [G SIB #s 271723/1, 2, 3]. Thus the pro
parte designation for the type of Lippia virgata var.
elliptica Briq. referred to Balansa’s collection number
1016 mixed with another taxon in a different family,
rather than any confusion on Balansa’s part for
varieties in L. virgata. Consequently, a mixed
collection for Balansa 1016 exists at G that consists
of at least six sheets for Balansa 1016, three for L.
virgata var. elliptica and three for Malabaila
carvifolia (Apiaceae), with all six sheets indicated
as type material.
G image!, K [bc] K000545989 not seen, K image!; P
[bc] P00753761 not seen, P image!, P [bc] P00753760
not seen, P image!, P [bc] P02851825 not seen, P
image!, MPU [bc] MPU012512 not seen, MPU image!,
UC [bc] UC935077 not seen, UC image!).
Aloysia virgata var. argutedentata Moldenke, Phytologia
55(4): 232. 1984. TYPE: Argentina. Santiago del
Estero, C. Pellegrini, cerro del Remote, 14 Jan. 1928,
S. Venturi 5764 (holotype, US [barcode] US00118883
not seen, US image!; isotypes, A [bc] A00354571 not
seen, A image!, F [bc] F0092407F not seen, F image!,
GH [bc] GH00354572 not seen, GH image!, NY [bc]
NY01365328 not seen, NY image!, S11-10466 not
seen, S image!, SI [bc] SI3404!).
Selected specimens examined. ARGENTINA. Chaco: 18
de Mayo, Colonia Benı́tez, Venturi 7897 (SI). Corrientes:
´ Rapidos del Apipe,
´ Cabrera 28959 (SI); s. loc.,
Ituzaingo,
Burkart 6923 (SI). Entre Rios: Diamante, Puerto, Bacigalupo 1659 (SI). Formosa: Pirané, ferrocarril, Krapovickas
1136 (SI). Jujuy: El Carmen, Pampa Blanca, Kiesling 1654
(SI). Misiones: Capital, Posadas, F. M. Rodrı́guez 97 (SI).
Salta: Metán, El Tunel, Saravia Toledo 1929 (SI). Santiago
del Estero: Carlos Pellegrini, co. Del Remate, Venturi 5860
(SI). Tucumán: Burruyacú, Alto de Medina, Venturi 2692
(SI). BOLIVIA. Chuquisaca: El Salvador, El Huare, Saravia
Toledo 10339 (SI). Santa Cruz: Cordillera, Camiri, Ferrucci
2706 (SI). Tarija: Villa Monetes, Qda. de Tampinta,
Krapovickas 19383 (SI). PARAGUAY. Alto Paraná:
Fiebrig 6151 (SI). Boquerón: Colonia Menno, rio Verde,
Vanni 1858 (SI). Caaguazú: rte. 2, Km. 122, Zardini 10633
(SI). Central: Yaguarón, Krapovickas 12255 (SI). Chaco:
Carnachini, Rojas 7216 (SI). Cordillera: San Bernardino,
Hassler 263 (SI). Guairá: Colonia Independencia, serranı́a
Ybytumesú, Schinini 14 (SI). Misiones: San Ignacio,
Burkart 18241 (SI). Nueva Asunción: 60 km W Est. La
Patria, Nicora 9758 (SI). Paraguarı́: Cerro Mbatovı́, Zardini
4451 (SI). San Pedro: Villa Primavera, Woolston 793 (SI).
28b. Aloysia virgata (Ruiz & Pav.) Pers. var.
virgata.
Aloysia urticoides Cham., Linnaea 7: 238. 1832. Lippia
urticoides (Cham.) Steud., Nomencl. Bot. 2: 54. 1841.
TYPE: Brazil. s. loc., s.d., F. Sellow s.n. (lectotype,
designated by Siedo [2010: 201], G [barcode]
G00208721 not seen, G image!; isolectotypes,
G [bc] G00386463 not seen, G image! GH!, HAL
[bc] HAL0098258 not seen, HAL image!,
K [bc] K000487003 not seen, K image!, NY [bc]
NY0136532 not seen, NY image!, P [bc] P00713758
not seen, P image!, US not seen, US image!).
Lippia urticoides (Cham.) Steud. var. laxa Chodat, Bull.
Herb. Boissier, ser. 2, 2: 819. 1902. Lippia virgata
var. laxa (Chodat) Briq., Annuaire Conserv. Jard. Bot.
Genève 7–8: 304. 1904. Aloysia virgata var. laxa
(Chodat) Moldenke, Phytologia 1: 95. 1934. TYPE:
Paraguay. Sierra de Mbaracayu, Oct. 1898–1899, E.
Hassler 5206 (holotype, G [barcode] G00306073 not
seen, G image!; isotypes, G [bc] G00400306 not seen,
Shrubs with elliptic leaf blades, longer than wide,
with acute apex, and acute to subobtuse base.
Glabrous ovary.
Distribution and habitat. Aloysia virgata var.
virgata grows in Bolivia, Paraguay, Peru, southern
Brazil, and northern Argentina. It has been observed
in open fields and sometimes in sandy soils.
Discussion. Aloysia virgata is distinguished by its
lax, subpendulous florescences, solitary or two to five,
sometimes seven, per leaf axil. It is similar to A.
peruviana and A. scorodonioides. However, it is
contrasted by its leaf margins minutely serrate or
crenate and shorter corollas, less than 4 mm,
contrasted with leaf margins notoriously crenate or
dentate, and corollas more than 4 mm long in these
two last taxa.
There is a certain phenotype of Aloysia virgata var.
virgata found in southern Brazil (Mato Grosso do Sul,
Paraná, Santa Catarina) and northeastern Argentina
(Misiones), with large leaves to 15 cm long, with
narrowly elliptic blades and an acute apex, and long
florescences to 15–20 cm in fructification. Generally,
there are five to 12 florescences per node, many times
reiterated in successive nodes, which makes the plant
look like a feather duster. Siedo (2006) referred to
this form as A. virgata var. urticoides (Cham.) Siedo,
based upon A. urticoides Cham.; however, Siedo’s
variety was never validly published and is an ined.
name. In the present treatment we consider this to fall
within the range of morphological variation of A.
virgata s. str.
Selected specimens examined. ARGENTINA. Catamarca:
˜ y Ancasti, Biurrum 8133 (SI).
Ruta Prov. 2, entre Icano
Chaco: Ruta 11, 54 millas N de Resistencia, Cordo 77-A-96
(SI). Corrientes: San Miguel, 12 km. N de San Miguel, Ruta
17, Ahumada 2375 (SI). Formosa: Ing. Juárez, Burkart
20287 (SI). Jujuy: San Pedro, Sierra de Zapla, Burkart
11993 (SI). Misiones: El Dorado, ruta 17, a 15 km de El
˜ Dique Cabra
Dorado, Cabrera 28951 (SI). Salta: La Vina,
Corral, Cabrera 29734 (SI). Tucumán: Tafı́ Viejo, Tapia,
Rodriguez 545 (SI). BOLIVIA. Beni: San Borja, Beck 12734
(SI). Chuquisaca: Siles, Beck 9364 (SI). La Paz: Nor/Sud
Volume 101, Number 3
2016
O’Leary et al.
605
Revision of Aloysia (Verbenaceae) in South
America
Yungas, Puente Villa, Beck 4791 (SI). Santa Cruz:
´ Michel 115 (SI). BRAZIL. s.
Cordillera, Alto Parapetı,
loc., Saint Hilaire s.n. (MVM). Bahia: s. loc., Blanchet 1330
(SI). Mato Grosso do Sul: Bela Vista, Schinini 1993 (SI).
Minas Gerais: s. loc., Claussen 6087 (SI). Paraná: Perola
D’Oeste, Hatschbach 22629 (SI). Rio de Janeiro: entre
Macuco & Santa Marı́a Magdalena, Santos 2043 (SI). Santa
Catarina: Cambará, aguas de Chapecó, Klein 5604 (SI). Sao
˜
Paulo: Porto Feliz, Morello 49 (SI). PARAGUAY. Alto
Paraná: Pto. Bestosi, Rojas 7987 (SI). Caaguazu: ruta 2,
´ ‘‘Ybytu Silla,’’
Zardini 10579 (SI). Cordillera: Tobatı,
Zardini 27301 (SI). Guaira: Ybytyruzú, Cerro Polilla,
Zardini 13918 (SI). Paraguarı́: Chololó, Eskuche 6247 (SI).
PERU. Cuzco: Santa Ana, Cook 1484 (US). Junı́n: Pte.
Herreria, Schunke Vigo 6202 (US). San Martı́n: Puente
Colombia, entre Tarapoto y Juanjui, Ferreyra 17541 (US).
Hickey, L. J. 1974. Clasificación de la arquitectura de las
hojas de dicotiledóneas. Bol. Soc. Argent. Bot. 16: 1–26.
Lawrence, G. H. M. 1951. Taxonomy of Vascular Plants.
The Macmillan Company, New York.
López Palacios, S. 1977. Verbenaceae, Aloysia. Pp. 180–
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International Botanical Congress. IBC2011 Abstracts
Book: 210. Melbourne.
Lu-Irving, P. & R. G. Olmstead. 2013. Investigating the
evolution of Lantaneae (Verbenaceae) using multiple
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McNeill, J., F. R. Barrrie, W. R. Buck, V. Demoulin, W.
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K. Marhold, J. Prado, W. F. Prud’homme van Reine, G.
F. Smith, J. H. Wiersema & N. J. Turland. 2012.
International Code of Botanical Nomenclature (Melbourne Code). Reg. Veg. 154. Koeltz Scientific Books,
Königstein.
Moldenke, H. N. 1935. Nomenclatural and taxonomic notes.
Phytologia 1: 167–171.
Moldenke, H. N. 1940. The flora of extra-tropical South
America. Lilloa 5: 353–440.
Moldenke, H. N, 1941. Novelties in the Eriocaulaceae and
Verbenaceae. Phytologia 2: 6–32.
Moldenke, H. N. 1942. The Known Geographic Distribution
of the Members of the Verbenaceae and Avicenniaceae.
Edwards Brothers, Inc., New York.
Moldenke, H. N. 1947. Notes on new and noteworthy plants
III. Phytologia 2: 363–372.
Moldenke, H. N. 1948. Notes on new and noteworthy plants
IV. Phytologia 2: 408–428.
Moldenke, H. N. 1949. Notes on new and noteworthy plants
VIII. Phytologia 3: 106–122.
Moldenke, H. N. 1961. Material toward a monograph of the
genus Acantholippia. Phytologia 7: 326–338.
Moldenke, H. N. 1968. Notes on new and noteworthy plants
XLIX. Phytologia 15: 462–463.
Moldenke, H. N. 1982. Notes on new and noteworthy plants
CLV. Phytologia 50: 308–310.
Moldenke, H. N. & A. L. Moldenke. 1983. Verbenaceae,
Aloysia. Pp. 231–235 in M. D. Dassanayake & F. A.
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DOUBTFUL TAXON
Aloysia dodsoniorum Moldenke, Phytologia 50: 308.
1982. TYPE: Ecuador. Guayanas: Capeira,
Gauyaquil to Daule, 15 Sep. 1981, C. H. Dodson
& P. M. Dodson 11224 (holotype, TEX-LL
[barcode] LL00374937 not seen, TEX-LL image!).
No further material, apart from the type specimen,
could be found for this taxon. The type material
resembles a Lippia.
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Appendix 1. Accepted species and varieties of Aloysia.
1. Aloysia arequipensis Siedo
2. Aloysia brasiliensis Moldenke
3. Aloysia castellanosii Moldenke
4. Aloysia catamarcensis Moldenke
5. Aloysia chamaedryfolia Cham.
6. Aloysia citrodora Paláu
7. Aloysia cordata Siedo
8. Aloysia crenata Moldenke
9. Aloysia deserticola (Phil.) Lu-Irving & N. O’Leary
10. Aloysia dusenii Moldenke
11. Aloysia fiebrigii (Hayek) Moldenke
12a. Aloysia gratissima (Gillies & Hook. ex Hook.) Tronc.
var. angustifolia (Tronc.) Botta
12b. Aloysia gratissima var. chacoensis (Moldenke) Botta
12c. Aloysia gratissima (Gillies & Hook. ex Hook.) Tronc.
var. gratissima
12d. Aloysia gratissima var. schulziana (Moldenke) Botta
13. Aloysia hatschbachii Moldenke
14. Aloysia herrerae Moldenke
15. Aloysia oblanceolata Moldenke
16. Aloysia ovatifolia Moldenke
17. Aloysia peruviana (Turcz.) Moldenke
18. Aloysia polygalifolia Cham.
19. Aloysia polystachya (Griseb.) Moldenke
20. Aloysia pulchra (Briq.) Moldenke
21. Aloysia riojana (Hieron. ex Moldenke) Lu-Irving & N.
O’Leary
22. Aloysia salsoloides (Griseb.) Lu-Irving & N. O’Leary
23. Aloysia salviifolia (Hook. & Arn.) Moldenke
24a. Aloysia scorodonioides (Kunth) Cham. var. hypoleuca
(Briq.) Moldenke
24b. Aloysia scorodonioides var. mathewsii (Briq.) Moldenke
24c. Aloysia scorodonioides var. scorodonioides
25. Aloysia tarapacana (Botta) Lu-Irving & N. O’Leary
26. Aloysia trifida (Gay) Lu-Irving & N. O’Leary
27. Aloysia velutina Siedo
28a. Aloysia virgata (Ruiz & Pav.) Pers. var. platyphylla
(Briq.) Moldenke
28b. Aloysia virgata var. virgata
Appendix 2. Index to collectors.
Collections are listed alphabetically by collector’s last name.
The number in parentheses, after the collector’s number,
corresponds to the species number in the species list
(Appendix 1).
Ahumada, O. 8 (12c), 986 (28a), 2375 (28b), 4458 (28b),
4665 (12c), 9002 (12c). Alberti, F. R. 619 (12c), 1232 (12c).
Aliscioni, S. S. 692 (12a). Alonso, E. 548 (12b). Arenas, P.
148 (15), 257 (19), 276 (6), 351 (6), 370 (15), 428 (22), 444
(6), 664 (12b), 927 (11), 1023 (6), 1085 (12a), 1879 (12b),
1916 (19), 1918 (6), 2330 (12d), 3266 (19), 3465 (19), 20262
(19). Arganara
˜ ´s, J. L. 61 (12b). Asplund, E. 6533 (24b).
Bacigalupo, N. 1659 (28a). Baer, G. A. 2 (12c), 31794
(16). Báez, J. R. 3 (12c), s.n. (12b). Balegno, B. 971 (12a),
1068 (12a), 1410 (12a), 1584 (12c). Barkley, F. A. 214 (12c),
s.n. (12c). Barrionuevo, A. s.n. (12a). Bartlett, H. H. 19200
(12c), 19543 (12c), 19687 (12d), 19942 (12c), 20004 (12c),
20076 (12c), 20513 (3), 20548 (9), 20573 (3), 20602 (12c).
Beck, S. 463 (12c), 874 (15), 3530 (24a), 4791 (28b), 6039
(24c), 9364 (28b), 9428 (24b), 12734 (28b), 14132 (22),
21555 (9), 26988 (11). Bertoni, M. 761 (12d), 3617 (20),
98421 (20). Biganzoli, F. 1414 (20). Biloni, A. 16624 (28b).
Biloni, J. S. 6255 (16), 6671 (22). Biurrun, F. 650 (12c), 718
Volume 101, Number 3
2016
O’Leary et al.
607
Revision of Aloysia (Verbenaceae) in South
America
(16), 974 (4), 1214 (12c), 1250 (16), 1302 (16), 2569 (12d),
2774 (6), 4002 (12d), 4080 (19), 4250 (19), 4272 (16), 4688
(6), 4700 (4), 4701 (16), 4740 (26), 4963 (9), 4996 (16), 5042
(3), 5639 (9), 5819 (3), 5980 (21), 5998 (4), 6370 (22), 6392
(12d), 7693 (21), 7696 (21), 7705 (21), 7706 (26), 7710 (21),
8133 (28b). Blanchet, J. 1330 (28b). Bocco, M. E. 821 (12c).
Boelcke, O. 1249 (12c). Boffa, P. s.n. (12b). Bonifacino, M.
1953 (5). Botta, S. M. 116 (12c), 273 (12c), 364 (22), 685 (4),
695 (3). Brescia, R. 4269 (20). Bridges, E. 1346 (23).
Brizuela, A. 90 (19), 181 (16), 386 (12c), 389 (19), 437 (12c),
550 (12c), 882 (12c), 1082 (19), 1180 (12c). Brown, A. 1610
(12d). Bruch, C. s.n. (12c). Buchtien, O. 3240 (24c).
Buratovich, F. 219 (12d), 941 (12c). Burkart, A. 3083
(12c), 4140 (12c), 6923 (28a), 6928 (12c), 8486 (12c), 11966
(22), 11993 (28b), 12536 (21), 12540 (21), 12542 (6), 12543
(16), 12545 (16), 12549 (4), 13195 (12d), 13856 (12c), 13858
(12c), 13968 (16), 14201 (20), 14209 (20), 15281 (20), 15834
(12c), 18002 (12c), 18241 (28a), 20226 (19), 20287 (28b),
21361 (12a), 22066 (3), 23796 (12c), 23800 (12a), 23805
(12a), 26347 (12c), 26348 (12c), 26594 (12c), 27051 (12c),
27876 (12c), 28067 (12a), 28074 (12c), 29450 (12c), 29465
(12c), 29641 (16), 30574 (16), 30593 (24c), 30595 (12d),
30596 (28b), 30606 (12d), 30608 (12c).
Cabezas, V. 23192 (12c). Cabrera, A. L. 3046 (12c), 4368
(12c), 7209 (12c), 7716 (22), 8192 (12c), 9016 (22), 12183
(12c), 13252 (22), 14241 (24c), 14633 (12d), 15711 (24b),
16695 (6), 16783 (4), 16877 (6), 18475 (12c), 19245 (12c),
20325 (12c), 20988 (24b), 21092 (12d) 22438 (22), 23330
(28a), 24637 (6), 24648 (12c), 27399 (22), 27962 (24b),
28063 (16), 28331 (20), 28485 (20), 28634 (20), 28743 (20),
28870 (28b), 28951 (28b), 28959 (28a), 28973 (12c), 28989
(20), 29017 (12c), 29029 (12c), 29062 (20), 29104 (28a),
29106 (8), 29117 (20), 29187 (28b), 29529 (12c), 29554
(12c), 29588 (16), 29603 (12c), 29616 (12c), 29632 (12c),
29664 (12a), 29670 (12c), 29727 (24b), 29729 (24c), 29731
(12c), 29734 (28b), 29736 (12d), 29745 (24c), 29746 (12c),
29828 (12c), 29961 (24c), 30346 (12d), 31018 (12d), 31725
(22), 31735 (22), 31792 (9), 32301 (28a), 32345 (12c), 32474
(22), 32746 (28a), 33665 (24c), 34018 (28a), 34101 (24c).
Calderón, C. E. 991 (12a), 1243 (12c), 1386 (12c). Cano, E.
724 (12c), 1971 (12c). Cantino, P. 327 (12c), 557 (12c), 559
(12c), 693 (3), 734 (16). Carette, E. 3046 (12c), 3877 (12c).
Castellanos, A. 585 (12a), 623 (12a), 11669 (19), 19062
(12c), 19615 (12a), 28/327 (6), 28/331 (16), 33887 (19),
33892 (12a), 33894 (16). Castillon, L. 943 (6). Cerón 12387
(12c). Cerrate, E. 1282 (1). Cialdella, A. 218 (3), 407 (22).
Claussen, P. 6087 (28b). Cocucci, A. 3277 (25), 3366 (11).
Cook 247 (24b), 1484 (28b). Cordini, R. I. 60 (12b). Cordo,
H. 77-A-33 (12c), 77-A-49 (12c), 77-A-50 (12a), 77-A-93
(12a), 77-A-94 (12c), 77-A-96 (28b), 77-A-101 (12b), 77-A102 (28b), 77-A-160 (12c), 77-B-11 (12c), 77-B-66 (28b), 77B-70 (12c), 77-B-82 (12c), 77-B-83 (19), 77-B-88 (12c), 77C-21 (12c), 77-C-33 (12c), 77-D-45 (12d), 77-D-46 (12c), 77D-47 (12c), 78-A-39 (28a), 78-A-41 (12a), 78-A-42 (12b),
78-A-43 (12d), 78-A-44 (12c), 78-A-45 (12d), 78-A-46 (12d),
78-A-48 (24c), 82-A-24 (22), 89-A-93 (12c). Correa, A. 209
(12c), 238 (12c), 4318 (4), 18073 (12c). Correa, J. B. 26
(12c). Corzo, R. 777 (19). Costa, M. 6 (7). Cozzo, D. s.n.
(12a). Crespo, S. 26457 (12c). Cristóbal, C. L. 1649 (12c).
Cuezzo, A. R. 971 (19), 1656 (3), 9362 (12c), 9498 (12c).
Davis, E. 1757 (14). Dawson, G. 3340 (6), 3432 (16). De
la Sota, A. V. 198 (12b), 261 (12c), 447 (12c), 458 (12c), 580
(12c), 4236 (12c). De la Vega, R. 26 (6), 44 (6). Deginani, N.
1375 (20). Del Castillo, A. 451 (12c). Del Puerto 6053 (20).
Dematteis, M. 1531 (20). Denham, S. 333 (12c). Descole, H.
R. 3330 (28b). Devoto, F. 1142 (24c), 1566 (12c), 2208
(12c), 2257 (12c), 3427 (24c). Dier, L. 148 (22). Dillon, M.
4544 (27), 6018 (9). Donadı́o, S. 177 (19). Donovan P978
(17). Drake, J. s.n. (24c). Dunn, D. 20552 (12c).
Ekman, E. L. 1999 (20), 2000 (20), 2004 (5). Escobar, H.
233 (25). Eskuche, U. 6247 (28b). Eukontes, J. 534 (20).
Eyerdam, W. 22363 (12c), 23408 (12c).
Fabris, H. A. 2729 (12c), 2994 (12b), 3081 (12c), 3546
(12c), 6036 (22), 6336 (22), 7954 (12c), 7989 (28b), 8121 (6).
Ferreyra, R. 755 (17), 759 (17), 7013 (24a), 7021 (24a),
12952 (17), 17541 (28b). Ferrucci, M. 2659 (15), 2706 (28a).
Fiebrig, K. 3040 (11), 5904 (20), 6151 (28a). Fortunato, R.
1269 (12c), 2339 (12c), 5040 (12c), 5091 (16), 5950 (12c),
6628 (12c). Forzza, R. C. 1977 (12c), 1978 (6). Frenguelli, J.
19 (12c). Fries, R. 746 (22).
Garaventa 7092 (12c). Garcı́a, E. 1209 (9). Garcı́a, P. 814
(12a). Gentry, H. 36089 (17), 44821 (17), 70190 (24c).
Gerold, H. 161 (6). Giardelli, M. L. 25 (12a), 418 (12a),
19542 (12c). Giberti, G. 825 (22). Golbach, R. 9 (12c).
Gomez 28/770 (19). Guaglianone, R. 220 (20), 599 (12c),
956 (20) 2071 (12c), 2787 (12c).
Haene, E. 93 (3), 2121 (9). Hagelund, K. 132 (12d),
10590 (5). Hart, C. 1481 (24c). Hassler 53 (12c), 263 (28a),
2635 (12c), 11497 (15). Hatschbach, G. 9339 (15), 14905
(2), 20792 (7), 22546 (10), 22629 (28b), 26325 (8), 28171
(18), 28366 (10), 30734 (18), 51897 (13). Haught, O. 3155
(24c). Hayward, K. 2067 (4). Herrera, E. 194 (19). Herter,
G. 158 (5). Hicken, C. 17 (22), 113 (12c), 3519 (22), 3528
(12b), 3531 (12c), 3537 (12c). Hieronymus, G. 82 (19), 547
(21), 755 (6). Huajardo, E. D. 2559 (12c), s.n. (3). Hunziker,
A. 2037 (21), 4729 (4), 4771 (19), 5069 (4), 7912 (16), 8055
(11), 8945 (12c), 8951 (19), 13507 (19), 15321 (6), 17046 (6),
18369 (12c), 21887 (4), 22803 (6), 24629 (16). Hunziker, J.
1042 (12c), 1276 (12c), 2037 (21), 11998 (16), 12605 (6),
12908 (12c), 13117 (4). Hurrel, J. 6907 (12c). Hutchison, P.
3520 (24b), 4199 (14), 4201 (24a).
Irigoyen, J. 142 (12c), 222 (12c). Isern, J. 8014 (12a),
8336 (12a).
Jiles, G. 5079 (26). Job, M. M. 573 (12a), 835 (12a), 1072
(12b). Jörgensen, P. 1020 (12c), 1023 (6), 1736 (22), 2473
(12c), 2474 (28a). Juárez, F. 1314 (12c).
Kiesling, R. 1654 (28a), 3069 (12c), 3458 (12c), 3535
(22), 3578 (6), 3708 (12c), 3992 (22), 4346 (3), 4821 (3),
4837 (3), 4912 (12c), 4954 (12c), 5243 (22), 5269 (22), 5370
(12c), 5533 (12d), 5941 (3), 6313 (3), 6620 (6), 8847 (9),
9130 (26). Klein, E. 3489 (2), 5604 (28b). Krapovickas, A.
1136 (28a), 1523 (22), 1645 (28a) 2597 (12a), 3239 (12c),
6492 (12a), 11741 (28a), 12255 (28a), 13743 (28b), 13748
(12d), 17337 (12d), 17970 (12d), 18530 (12c), 18544 (12c),
19210 (12d), 19383 (28a), 20788 (20), 22064 (6), 25485 (20),
25760 (20) 26801 (12c), 26802 (28b), 26989 (12c), 27356
(12c), 27453 (20), 27999 (28b), 28868 (5), 30307 (12c),
30899 (12d), 31277 (19), 38344 (10). Kristensen, K. 1359
(9). Kuntze, O. s.n. (12c).
Lanfranchi 1076 (12c), 1097 (16). Lee Anderson 748
(12c), 1412 (16), 1521 (12c), 3077 (16). Legname, P. R.
6887 (12c), 9108 (12c). Leuenberger, B. 3981 (28a), 4773
(12c). Lillo, M. 3278 (12c), 6073 (16), 7183 (12c), 32305
(12a). Llatas Queiroz, S. 1513 (14), 1953 (27). López, E. 34
(12c). Lorentz, P. G. s.n. (12c). Lourteig, A. 1037 (12c),
2189 (10). Lu-Irving, P. 9-32 (24c), 9-62 (24b). Luna, P. E.
147 (19). Lundell, C. 11958 (12c).
Macbride, J. 133 (24a). Maldonado, R. 206 (12a), 860
(12d), 970 (12c). Malvarez, R. 286 (12c), 1367 (12a), 1431
(12c). Maranta, B. 1104 (12d). Marchesi, E. 10078 (12c).
Marquez 63 (6). Martı́nez, A. s.n. (12c). Martinez, E. 479
(6). Martı́nez, G. 94 (12c), 910 (12c). Martı́nez Crovetto, R.
4231 (12c), 8956 (20), 9474 (20), 9597 (20), 9935 (20),
10638 (12c), 10801 (20), 10824 (28b), 11363 (12c). Maturo,
608
Annals of the
Missouri Botanical Garden
H. 160 (28a). Medina, B. R. 240 (20). Melillo, A. C. 2339
(12b). Meyer, T. 2671 (12b), 3367 (12c), 3370 (3), 4248 (6),
4289 (12c), 4868 (12c), 5066 (12b), 8590 (12b), 9994 (12c),
10639 (12c), 10954 (12c), 11090 (12c), 11742 (20), 11965
(20), 12156 (12c), 16436 (12d), 22633 (22), 23456 (12c),
34400 (11). Michel, A. 115 (28b). Moldenke, H. 19729 (3).
Monetti, L. 1035 (12c). Montes, J. E. 12 (20), 534 (12c),
1031 (28b), 1410 (20) 1864 (20), 2125 (28b), 2299 (20), 2314
(28b), 3434 (28b), 14841 (20), 14911 (20), 15011 (28b),
15161 (20), 15455 (28b), 27627 (20), 27697 (20). Morel, J.
146 (12b) 913 (12b) 1257 (12c), 2049 (12c), 4518 (12c).
Morello, J. 49 (28b), 1966 (12c), 4013 (12a). Moretti, A.
1925 (6). Morrone, O. 635 (12c), 1749 (20), 3052 (12c),
3113 (12d), 4119 (12c), 4366 (12c), 4367 (6), 4634 (24c).
Múlgura, M. 481 (20), 767 (12c), 925 (28a), 1105 (12c),
1158 (12c), 2204 (28b), 3183 (28a), 3743 (12c), 3847 (28b),
4142 (6), 4223 (22).
Naranjo, C. 923 (12c). Nicora, E. 966 (12c), 1310 (12c),
1711 (12b), 2321 (12c), 2494 (19), 3276 (12c), 3288 (12c),
4640 (12c) 4726 (12c), 8260 (9), 8472 (9), 8483 (9), 8565 (9),
8612 (9) 9758 (28a) 17824 (16) 19558 (12a). Niederlein, G.
23907 (20). Novara, L. 505 (12d), 855 (12c), 1301 (12c),
1829 (12c), 1954 (12c), 2331 (19), 5703 (9), 6305 (12c),
7260 (12c), 7550 (6), 9311 (28a), 10106 (24c). Núne
˜ z, O.
7018 (14).
Ochoa, C. 710 (14). O’Donell, C. A. 3159 (12c), 4366
(12c), 5377 (12c). Okada, K. 2747 (12c). Olea, D. 78 (6), 99
(12d). Olmstead, R. 2001-184 (16), 2004-109 (12c), 2004125 (28b), 2004-129 (20), 2004-133 (28b), 2007-13 (6),
2007-13 (6), 2007-52 (22), 2007-68 (28a), 2007-82 (4),
2007-82 (4), 2009-30 (14), 2009-40 (24a), 2009-45 (17),
2009-8 (27), 2010-217 (10).
Pachano, A. 120 (24c). Parodi, L. R. 14244 (12c).
Pastore, F. 2031 (12c). Paula-Souza, J. 6989 (12c), 7623
(22), 7697 (22), 7796 (12d), 8122 (28a). Pedersen, T. M. 614
(12c), 874 (12c), 5564 (12c), 6428 (12c), 8261 (12c), 11796
(19), 15295 (4). Pennell, M. 14438 (24c). Pensiero, J. 1722
(12c), 4249 (12c), 5640 (12c), 5707 (12c), 7426 (12d), 7450
(12c). Perez Moreau, R. 13577 (12a). Pfsister 8318 (23).
Pierotti, S. 16 (12a). Pittier, H. 970 (6). Pozner, R. 144 (4).
Prado, D. E. 107 (12c). Prina, A. 2696 (12c). Pringle, C. s.n.
(12c). Pujalte, J. C. 133 (9).
Quarı́n, C. 2117 (20). Queiroz, L. P. 13358 (12d), 13465
(22).
Ragonese, A. 2604 (12a), 2623 (20), 2829 (12c), 3031
(12a), 3120 (12a), 7162 (28a), 9426 (12a), 9635 (12c), 9676
(12c), 23993 (12d). Rambo, B. 28141 (2), 49976 (15). Reca,
A. 22 (22), 33 (22). Rentzell, I. 18837 (12c), 19133 (12c),
19236 (12c). Renvoize, S. A. 2898 (12c), 3002 (20), 3156
(20), 3389 (12c), 3537 (28a). Ricardi, M. 23967 (23). Riedel,
L. 226 (16). Risso, J. L. 872 (6). Rivero, R. 44 (9). Rodrigo,
A. P. 2535 (12b). Rodriguez 6 (12c), 66 (20), 216 (12c), 545
(28b), 661 (12c), 891 (12c), 1196 (16), 1214 (6), 23848 (20),
30/2059 (20). Rodriguez, D. 1175 (12c), 1214 (6).
Rodriguez, F. M. (9), 97 (28a), 123 (20), SI 28218 (9). Roig,
F. A. 8166 (16), 8424 (16). Rojas, T. 2542 (12a), 5903 (8),
7216 (28a), 7701 (12b), 7902 (12c), 7987 (28b). Romanczuk, C. 20 (12c), 420 (28a), 741 (20). Rosengurtt, B. 2245
(12c), 4967 (5). Rotman, A. 189 (12c), 228 (16), 241 (12c),
304 (16), 518 (12c), 673 (12c). Rúgolo, Z. 1275 (12a). Ruiz
Huidobro s.n. (12c), 1102 (12c), 3134 (12a), 4565 (20), 4886
(20), 5284 (20), 5395 (20), 5457 (20), 5551 (20). Ruiz Leal,
A. 1102 (12c), 1220 (12c), 1505 (12c), 3877 (12c), 4476
(12c), 8795 (16), 9167 (16) 9848 (12c), 10439 (12c), 22090
(9), 22101 (3). Ruthsatz, B. 109 (22), 118 (22), 132 (22), 168
(22), 213 (9), 242 (22), 328 (22).
Sagástegui, A. 14147 (27). Saint Hilaire, G. s.n. (28b).
Sanderman, S. 4612 (14). Santos, A. 2043 (28b). Santos
Biloni, J. 6618 (22). Sanzin, R. 42 (12c). Saravia Toledo, C.
716 (12c), 743 (12c), 902 (12c) 1247-a (19), 1440 (12c), 1591
(24c), 1750 (12d), 1764 (24b), 1766 (12c), 1929 (28a), 1931
(12c), 1968 (12d), 10339 (28a), 11830 (12c), 12085 (12c).
Sayago, M. s.n. (6), 407 (16), 557 (6), 940 (12c), 1942 (6),
2397 (16), 2407 (16), 2553 (16), 2608 (16), 2649 (12d).
Scarpa, G. 701 (19). Schinini, A. 14 (28a), 1993 (28b), 6767
(6), 7525 (12c), 9688 (12c), 10366 (12c), 12770 (12c), 12822
(12c), 13902 (12c), 14019 (20), 14121 (20), 17016 (12c),
24130 (12c), 26306 (28a), 26811 (12c), 27599 (20), 34455
(28b). Schreiter, R. 7133 (6), 9475 (6), 11126 (22), 37997
(6), 37998 (6). Schulz, A. G. 2893 (24c), 2985 (24b), 8740
(28a), 10361 (28a). Schulz, C. L. 333 (12b), 556 (12d), 772
(12b), 1493 (12b), 6467 (12b), 6875 (20), 8317 (12c) 8699
(12d), 9099 (12c), 11465 (12c). Schunke Vigo 6202 (28b).
Schwarz, G. J. 763 (20), 1687 (20), 1924 (20), 1952 (20),
2312 (20), 3233 (20), 3334 (20), 3742 (20), 3803 (20), 4444
(20), 4561 (20), 4610 (20), 5439 (20), 5536 (20), 6398 (20).
Schwindt, E. 110 (20). Semper, J. s.n. (16), 116 (12c), 339
(12c). Sesmero 304 (20). Sielo 3278 (12c), 9790 (28b). Sigle
137 (12c). Silva 7037 (2). Slanis, A. C. 23 (12d), 40 (12c).
Sleumer, H. 3291 (22). Smith 9012 (2), 12478 (18), 13029
(18), 13577 (18), 14930 (20), 15683 (18). Solı́s Neffa, V. 875
(6). Solomon, J. 15755 (6). Soria 2098 (15). Soriano, A. 940
(12c), 944 (4), 1102 (12a). Soukup, J. 3741 (17), 4872 (17),
5467 (14). Soza, V. 1831 (22), 1834 (16). Steibel, P. 2334
(12c), 3178 (12c). Steinbach, J. 8248 (12c). Stienstra s.n.
(20). Stuckert, T. 1354 (19), 7004 (3), 12584 (24c), 17046
(19), 21287 (24c). Sturzenegger s.n. (19). Suero, A. s.n.
(12c).
Taylor, A. 11607 (26). Terribile, M. 376 (12b), 418 (6).
Thode, V. 157 (13), 398 (18). Torrico 107 (12d). Troiani, H.
558 (12c). Troncoso, N. s.n. (12c), 299 (12c), 1062 (12c),
1253 (12c), 1277 (12c), 1824 (12d), 1826 (12c), 1856 (12c),
1857 (12c), 1859 (12c), 1860 (12c), 1861 (6), 1893 (6), 1895
(6), 1896 (6), 1897 (6), 1898 (6), 1910 (12b), 1929 (6), 1930
(12c), 1931 (6), 1989 (12c), 2496 (12a), 20589 (12c).
Ulibarri, E. 332 (4), 333 (12c), 679 (9), 906 (16), 950 (4),
1481 (9).
Vanni, R. 1830 (12c), 1858 (28a), 2410 (12d). Varela 675
(12c). Vargas, C. 248 (14), 594 (24b), 3692 (24c), 12671 (1).
Vattuone, I. C. 71 (12c). Velarde Nune
˜ z, O. 301 (17).
Venturi, S. 834 (12c), 849 (24c), 2692 (28a), 3756 (19), 3963
(16), 4262 (6), 4885 (22), 5860 (28a), 7458 (28a), 7897 (28a),
8144 (22), 8300 (22), 10579 (24b). Vervoost, F. 498 (12c),
688 (22), 4318 (4), 4476 (22), 8642 (12c). Vignati, M. 977
(12c). Villafane,
˜ M. 342 (12c), 464 (12c), 558 (12c), 693
(12a), 751 (12a), 776 (12c).
Wall, E. s.n. (12c). Werdermann 103 (23). Werner, D.
789 (22). Wilkes s.n. (24a). Wood, J. 14658 (14). Woolston,
A. 793 (28a), 794 (12c). Wulff, A. 103 (12c).
Xifreda, C. 519 (20).
Zabala, S. 526 (19). Zardini, E. 4451 (28a), 8599 (12c),
10036 (20), 10579 (28b), 10633 (28a), 13918 (28b), 27301
(28b). Zuloaga, F. O. 2687 (12c), 3670 (12d), 3738 (12c),
3855 (12c), 5036 (28b), 5405 (28b), 6384 (4), 6703 (28b),
7891 (12c), 8632 (6), 9170 (22), 9257 (6), 9296 (22), 9362
(12c), 10188 (6), 10596 (12c), 11490 (24b), 12706 (3), 12845
(16), 12857 (6), 12912 (12c), 12972 (6).
Appendix 3. List of taxa newly synonymized in the present
work.
Aloysia ayacuchensis Moldenke [¼ A. herrerae Moldenke]
Volume 101, Number 3
2016
O’Leary et al.
609
Revision of Aloysia (Verbenaceae) in South
America
Aloysia axillaris J. R. I. Wood [¼ A. scorodonioides var.
hypoleuca (Briq.) Moldenke]
Aloysia depressa Ravenna [¼ A. scorodonioides var.
hypoleuca (Briq.) Moldenke]
Aloysia floribunda M. Martens & Galeotti [¼ A. gratissima
(Gillies & Hook) Tronc. var. gratissima]
Aloysia gratissima var. oblanceolata Moldenke [¼ A.
oblanceolata Moldenke]
Aloysia leptophylla Moldenke [¼ A. scorodonioides (Kunth.)
Cham. var. scorodonioides]
Aloysia minthiosa Moldenke [¼ A. peruviana (Turcz.)
Moldenke]
Aloysia reichei Moldenke [¼ A. trifida (Gay) Lu-Irving & N.
O’Leary]
Lippia fonckii Phil. [¼ A. trifida (Gay) Lu-Irving & N.
O’Leary]
Volume 101, Number 3, pp. 457–610 of ANNALS OF THE MISSOURI BOTANICAL GARDEN was published on
27 APRIL 2016.