TAXONOMY OF LANTANA SECT. LANTANA (VERBENACEAE): I. CORRECT APPLICATION OF LANTANA CAMARA AND ASSOCIATED NAMES

2006·24.7 MB·English

by  Roger W Sanders

TAXONOMY
LANTANA LANTANA
OF
SECT.
CORRECT
(VERBENACEAE): APPLICATION OF
I.
LANTANA CAMARA AND NAMES
ASSOCIATED
Roger W.Sanders
Botanical Research Institute of Texas
509 Pecan
Street
Fort WorthJexas 76102-4060,
U.S.A.
[email protected]
ABSTRACT
The previous lectocypification of Lantana camara L. is evaluated by examination of characters of
the lectotype, review of other original material and documentation of current usage of the name.
The current usage analyzed by surveying pertinent luerature and by samphng specimens an-
is for
between and
notations 1753 the present in relation to critical characters ol those speciiriens. Current
usage oi L. camara includes a widely cultivated and naturalized cultigcn species oi hybrid origin
name
that taxonomically distinct from camar'a. To determine the correct of the cultigen,
is L. all
names in Lantana sect. Lantana are reviewed, typified necessary and possible, and taxonomically
if
No name
disposed. available applies to the cultigen, which newly described dsLantana snigocamara
is
R.W. Sanders. Origin of the named hybrids in the camara complex hypothesized. Two new com-
L. is
R.W
binations are made, Lantana camara subsp. aculeata Sanders and Lantana nhea Vent, subsp.
IL.)
mutabilis (WJ. Hook.) R.W. Sanders. Nineteen lectotypifications and seven epitypifications are made.
RESUMEN
La lectotipificacion previa de Lantana camara L. se evalua mediante examen dc los caracteres del
lectotipo, revision de otro material original, y documentacion del uso actual del nombre. El use ac-
tual se analizo revisando la bibliografia adecuada y por muestreo de especimenes entre 1753 y el
presente por comentarios en relacion a caracteres criticos de esos especimenes. El uso actual de
L.
camara una ampliamente
incluye especie culti\ada naruralizada de origcn hibrido
y cjue es
taxonomicamente diferente de camara. Para determinar nombre correctode planta cultivada
L. el la
se revisaron todoslos nombre de Lantana sect, Lantana, se tipificaron cuando fue necesario posible,
y
taxonomicamente decidido. No hay nombres disponibles para planta cultivada, describe
y la c]ue se
como R.W
Lantana strigocamara Sanders. Se hace una hipotesisdel origen de los hibridos nombrados
en complejo camara. Se hacen dos combmaciones nucvas, Lantana camara subsp. aculeata
el L. (L.)
RAV. Sanders y Lantana nhea Vent, subsp. mutabilis (WJ. look.) R.W. Sanders. Se realizan diez y nue\'e
I
lectotipificaciones y siete epitipificaciones,
name
have gone under Linnaean Lantana camara
Plants that the are well
L.
known, not only as hardy summer ornamentals worldwide but also as natural-
humid
ized weeds having devastating economic impact in tropical areas of the
&
Old World (Wolf son Solomons 1964; Eiow^ard 1970; Stirton 1977; Swarbrick
Day The and
et 1995; et 2003). systematics of these cultivated naturalized
al. al.
made
plants not well understood, even though considerable has been
effort
is
to elucidate their biological control (Day 2003). This confusion arises in
et
al.
SIDA22(1):381 -421.2006
BRILORG/SIDA
382 22(1)
part from the convoluted history of exploration, cultivation, hybridization, and
artificial selection that began at least 60 years before the pubhcation ot Linnaeus'
Species Plantarum (1753). Records that have been compiled (Howard 1969; Stir ton
1977) suggest that, during the eighteenth century, fanciers hybridized different
vvald species and infraspecific taxa of LantanaL. sect Lantana from Mexico, the
shown names
West Indies, and Brazil. As in this paper, the early validly published
on garden-grown and com-
were based material (almost entirely so until 1817
much
monly which
so into the 1850s), of consisted of hybrid combinations.
As herbarium specimens Lanta na became
of wild-collected species of
sect.
many
available after the early 1800s, of the available, poorly distinguished
names Adding was
were frequently misapplied to them. to this confusion the
introduction of cultivated hybrids into neotropical regions where indigenous
taxa occur. Due to the propensity of lantanas to undergo polyploidy and the
& &
odd Ahuja Khoshoo
polyploid (Natarajan
partial fertility of levels 1957;
&
Mahal
1967; Spies 1983, 1984; Spies Stirton 1982a, Sanders 1987a, even
b, b),
c;
more complex hybrids formed between indigenous taxa and the escaped hy-
brid cultigens (Sanders 1987a, 1989a). Thus, the limits of natural variation
b,
c,
have been obscured, hampering the ability of taxonomists to develop effective
classifications for the group.
Schauer (1847), Briquet (1895), and Troncoso (1974) developed current sec-
Camara
Laniana Lantana Cham.)
tional concepts. Species of sect. (-sect. are
by predominantly narrow somewhat
characterized floral bracts that are in-
among
conspicuous the tubular corolla bases, by usually yellow or orange pig-
known
mented
corollas (white-flowered populations in several species [unpubl.
obser see also discussion below concerning purplish pigments in the section),
v];
and by blackish drupes. Each drupe contains a characteristically inflated com-
pound endocarp that resembles a horse skull in which the seed chambers are
s
Some
in the position of the eye sockets. of the species of the other major section,
Lantana CallioreasChdim., might be confused with those Lantana.
sect. of sect.
Generally species of Callioreas are described as involucrate, usually hav-
sect.
ing ovate to reniform conspicuously imbricate Iloral bracts and purplish co-
However, needed placement— drupes
rollas. fruits are to insure correct sectional
are usually wdiite or purplish and endocarps are subglobose, bilobed (seed
chambers each hemisphere), noninflated, and ornamented.
reticulately
lill
Schauer (1847) published the only worldwide revision of Lantana in De
more work and
Candolle's Prodromus. All recent has been limited regional
to
arden floras. The foremost student ol Verbenaceae in the twentieth century,
Harold N. Moldenke, never produced a revision of Lantana, but he did describe
number new
and
a of species mlraspecific taxa. Horticulturalists, ecologists,
much
and some taxonomists have submerged Laniana
floristic effectively of
Lanta na camara, wide spectrum
sect. into L. treating this of variation as a single
SANDERSJAXONOMY OF LANTANA SECT. LANTANA 383
Hortorium Schemske
Kuntze Troncoso 1965, 1974; Bailey 1976;
species, 1891;
(e.g.,
CuUen
Huxley 2000).
1976, et al. 1992; et al.
wild
Sanders (1987a, 1989a, b) undertook a study of the variation of
b,
c,
and populations Lantana Lantana in Florida and parts of
naturalized of sect.
West By chromosome numbers and meiotic behavior with
the Indies. correlating
wild
niorphology, he was able to distinguish the natural (often diploid) taxa
from naturalized and spontaneous hybrids and to develop morphological
the
and
taxonomic Sanders found that plant architecture, leaf bract size
criteria.
development and
and indument lorescence/infructescence
shape, features, inf
and pigment and dosage
fruiting bract persistence, floral classes effects
size,
and
correlate well with cytology, geographic distributions, species delimitations.
On and
the other hand, he argued that development of prickles specific corolla
many
colors, characters relied upon by other authors, vary too widely within
among supported by
and be conclusion recently
taxa to effective criteria, a
Day Further support comes from
molecular studies (Scott et al. 1997; et al. 2003).
the work of Isidro Mendez of Cuba who took up the study of Cuban and West
S.
Mendez
Indian Verbenaceae (1992, 1993, 2002). explicitly accepted the taxo-
nomic proposed by Sanders Lantana Lantana.
criteria for sect.
The natural taxa Lantana into three separate phenetic groups
of sect. fall
on
(to be validated in a subsequent paper; cladistic status not yet determined)
the basis of trichome structure on the abaxial surface of the leaf blades, as well
The
"pilose-morph"
shape, adaxial and venation of the leaf blades.
as luster,
by spreading
species (Pilose Group, Fig. are characterized soft, filiform, (erect
1)
or curly) hairs usually densely disposed on all veins and intervening tissue. Most
rounded
narrowly
of these have leaf blades that are broadly ovate to deltate, to
cordate the base and acuminate to obtuse at the apex, dull above (epidermis
at
The
and more pinninerved. "strigose-morph" species
poorly or less
reflective),
Group, have the abaxial hairs developed as conspicuous strigae,
(Strigose Fig. 2)
and and
antrorsely geniculate, are scattered re-
the hairs are stout, conical,
i.e.,
midrib, and secondary and sometimes higher order veins,
stricted to the tertiary,
but not the intervening Leaf blades are usually ovate-elliptic, often
to tissue.
tapering base and apex, lustrous above, and more or less tnplinerved. The
at
Group, abaxially bear well-separated, Ion
"setose-morph" species (Setose Fig. 3)
more
flexible to subrigid, spreading, setiform trichomes that are or less restricted
hke
the midrib and secondary and tertiary veins. Otherwise they are the pi-
to
though groups have adaxial
lose-morph Thus, even three strigae,
species. all
the Stigose Group the only one with strigae on both surfaces.
is
Despite Mendez's (2002) acceptance of Sanders' criteria, he disagreed with
L L
name camara
camara. Because the
Sanders over the application of the is
and one two
proposed type of the genus (Jar vis 1992; Jarvis et al. 1993) of the
names
names most other the
oldest in the secton, the correct application of in
384
BRIT.ORG/SIDA
22(1)
J.
Fig ^ff/ifflf?flramaraJectotype (LINN 783.4), representing Pilose Group, a. inflorescence, b, representative leaf blade,
= mm;c&d =
c.adaxiai leaf surface, d.abaxial leaf surface. Scale bars:a&b mm.
5
l
section hinge upon the correct apphcation of camara. Asserting that Sanders
L.
had committed two Mendez
errors, stated:
Sanders (1989|alJ attributed this binomial camam] ro a phcnotype ver)' different from prevailing
|L.
among
opinion the authors later to Linnaeus and not in agreement with the lectotype selected by
Moldenke
Moldeiike
(1Q83).
lSt
However
until the present study, analysis the lectotype and subsequent ap-
oi
name
plication of the have not been straightforward. The characters
critical
were not visible on the lectotype since of leaves were mounted adaxial
all its
side up. Sanders (unpubl.) was not able to determine w^hether the lectotype
matched
the pilose-tnorph or the strigose-morph plants, especially given the
quality of the microfiche and xerographic images available the time. How-
at
because Moldenke had
ever, H. N. selected the type, Sanders (unpubl.) relied
primarily on the numerous annotations Dr Moldenke App.
of (Table b see
B,
Moldenke
also 1980a) to develop a concept of the species and apply the name.
Sanders' concept was further reinforced by annotations of CD. Adams, and the
SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 385
s-
(Sotorfer BM), representing StrigoseGroup.ajnflorescen
2i(7/7fffnfl5ffl&r/t/i7jectotype
Fig. 5./??.,
= =
1cm;b8fC mm.
leaves.b.adaxialleafsurface.c.abaxialleafsurface.Scalebars:a
1
A
treatments Lantana The FloweringPlants Jamaica (Adams 1972) and
of in
of
&
Flora Tropical Florida (Long Lakela 1971), as well as on identified culti-
of
name
vated material received under that from the U.S. National Arboretum,
Longwood and commercial
Gardens, reputable nurseries, hi cases, the plants
all
v^ere characterized by leaves dominated by strigose-morph hairs, cordate-ovate
blades, and dull upper surfaces^ (here subsequently called the "Strigose-Cor-
date-DuU-" or "SCD cultigen"; Fig. In the West Indies, some wild-collected
4).
specimens bear codominant mixtures of both hair morphs. However, cytologi-
SCD
cal study confirmed these to be hybrids between the cultigen and native
taxa of the Pilose Group (Sanders 1987b). Thus, Sanders concluded that the lec-
name
camara
totype had the strigose-morph hairs, and he applied the L. spe-
SCD Many Group
cifically to the cultigen. of the described species in the Pilose
L
(including horrida, and arida) he lumped into a single species
L. tiliifolia, L.
name
and
applied the urticijolia Mill.
L.
Mendez appears to have studied more recent, higher quality images in con-
nection with a project to lectotypify Linnaean species of Verbenaceae of the
West Indies in collaboration with Steve Cafferty of the Linnaean Typification
&
Project (Mendez Cafferty 2001). Although he, too, was unable to see the abaxial
surfaces, he concluded on other grounds that the lectotype matched the pilose-
Adams included concept of L camera the SCD cultigen and Lscabrida.Jhefew specin^ens of Lscobrida
^ in his
that studied and bear his annotation as L.camora actually have lustrous upper leaf surfaces.
I
386 BRIT.ORG/SIDA
22(1)
J.
Fig ifl/)t(7/?^towfa(////?fof?2(?^99JEX), representing Setose Group.a. inflorescence, b.adaxial leaf sur^^
= mm;b&c =
mm.
leafsurface.Scalebars:a
5
1
morph, not the strigose-morph plants. He applied the name camara the
to
L.
which name
taxon Sanders applied the
to L. urticijolia.
name
Mendez's two assertions merit redress, then another, later-published
If
SCD
should be applied to the cultigen that Moldenke, Adams, and Sanders called
Lantana camara. Of course, current usage, regardless of usage by authors of the
early post-Linnaean period, must be estabhshed. current usage of camara
If L.
SCD
has clearly replaced the Linnaean concept with the concept of the culti-
gen, then conservation of the nomenclatural type could serve as an alternative
solution.
Therefore the purpose here to evaluate the lectotypification by review-
is 1)
by
ing publication, clarifying the characters of the lectotype, by understand-
its
ing current usage of camara, and by determmnig any variance between the
L.
type and current usage; 2) affirm the correct application of that name, 3) nec-
if
essary, determine which other name to be applied to the SCD cultigen, and 4)
is
known
dispose of all other names, to me, that are applicable to the complex.
Lantana camara
Typification of
Moldenke and Moldenke Lantana camara by
(1983) effectively lectotypified
LINN
citing 783.4 as "type'^ (ICBN, Art. Greuter al 2000). This
7.11, et
was and Mendez
lectotypification accepted by Sanders (1989b) (2002). In March,
BM, and Dr
2004, visited Charlesjarvis, long-time participant in the Linnaean
1
Typification Project, consented to break the low^er stem to permit study of the
two The
abaxial surface of the lowest leaves (Fig. type, indeed, of the pi-
1). is
lose-morph and by shape and an
verified leaf inflorescence structure as ele-
is
ment of the naturally occurring species distributed from the Bahamas and
Greater Antilles, through Mexico south northwestern South America. Thus
to
of Mendezs two assertions, the one that "Sanders (1989) attributed this bmo-
mial to a phenotype...not in agreement with the lectotype selected by Moldenke
and Moldenke
(1983)"
correct.
is
t>^
^y^
K documenting
Table Data history of annotation by H.N. Moidenke and oth
annotated L camara
as var. montziana. See text for further discussion.
Years Authorities N Pilose % Mixed % Strigose % Setose %
U^
1981-2005 Moldenke (incl. v. moritz) 62 15 24% 20 32% 27 44% 0%
Moldenke (excl, v. moritz) 47 0, 20 43% 27 0%
Other 45 0% 25 56 20 44% 0%
0.
b
moritz) 107 14% 45 42% 47 44% 0%
All (incl. 5
V.
b
0% 49%
All (excl. V. moritz) 92 45 47 51 0%
b b
1956-1980 Moldenke 8%
(incl. v. moritz) 153 13 56 37% 83 54% 1%
1
Moldenke (excl. moritz) 141 1% 56 40% 83 59%
v.
1
1 1
Other 60 0% 68% 32% 0%
41 19
6%
All (incl. V. moritz) 213 13 97 46% 102 48% 0%
1
0%
All (excl. V. moritz) 201 97 48 0, 02 51% 0%
1 b
1
1931-1955 Moldenke 16 2 13% 6 38% 44% 6%
7
1
Other 0% 0% 00%
1
1 1
All 17 2 12% 6 35% 8 47% 6%
1
1906-1930 All 16 0% 10 62% 6 38% 0%
1881-1905 0%
4 75% 25% 0%
All 3
1
1856-1880 0%
00% 0%
All 2
2
1831-1855
33% %
All 3 33 0, 33%
b
b
1 1
1
1753-1830
All 15 10 67 0, 2 13% 7% 3°
b b 2
1
^^
BRIT.ORG/SIDA
388 22(1)
bJ^
5fr/30f(7m(7ro,holotype(5onrfer5/^5(),FT6]. a jnflorescences and representative leaves.
Fig. 4,I(7nffl/)
(7
= mm;c&d = mm.
adaxial leaf surface, d. abaxial leaf surface. Scale bars:b 5
c. 1
Development of current usage
Linnaeus' concept of camara was developed from an array of cultivated and
L.
The synonyms and he
horticulturally selected plants. illustrations that cited
in the protologue are based on vouchers that are primarily elements of the Pi-
same
lose Group. All Hortus Cliffortianus (1737) specimens are either the spe-
LINN Lantana specimen cut-down
cies as 783.4 (319 1-B!, Linn. Herb, at S [a
Herb. CLff. specimen, C. Jarvis, pers. comm., dig. photo!]), are hybrids between
L
that taxon and the Strigose Group (319 Lantana or arc eleinents of ho rrida
1!),
SANDERS, TAXONOMY OF LANTANA SECT. LANTANA 389
Kunth
Lantana and 320 Lantana
(319 l-Q, 1-D!, la![the latter possibly hybrid-
ized with the Strigose Group]). Moreover, an unnumbered Herb. Chff sheet
is
(!)
.
an element Group
of the Strigose splendens Medik.).
(L.
Linnaeus One
did not see the vouchers for the syntype illustrations. of these
(icon in Plukenet, Phytographia 385. 169L [Voucher: 98:143
114, 4. top-left
1. f.
&
Group
specimen, BM-SL!]), belongs to the Setose hirsuta M. Martens
(L.
Galeotti). The Commelin syntype has no known voucher (C comm.),
Jarvis, pers.
illustrates only an inflorescence, and cannot be placed trichome-morph. Even
to
may much
so, the trichome characters not have been of concern to Linnaeus.
L
which
For example, in 1767, he did segregate mista, has distinctly hispid twigs,
which
as depicted in a Dillenius plate (see App. B), he cited. However, Linnaeus
new
did not use this character to differentiate the species, but rather used the
protracted leaf-blades, longer bracts, and capitula with mixed corolla colors.
C
According to Jarvis (pers. comm.), no other original material known.
is
how
To determine somewhat broad Linnaean concept was modified
this
by later botanists, present two lines of evidence. The a survey of sampled
I first is
determme
specimens
to annotation patterns by taxonomists (Table
Fig.
5).
1,
m
L
The specimens are all those annotated by H. N. Moldenke as camara LL/
TEX
Moldenke and BRIT/SMU,
(including the Herb.) a selection of those at K,
OXE
and few which obtained photographs BM, LINN, and
a for at Besides
I
Moldenke, annotating authorities include: Linnaeus, Medikus (implied by
cita-
tion, 1775), Schauer, Urban, Merrill, Hutchinson, R. Meikle, K. Morton, R.
J.
Fernandes, B. Verdcourt, G. Bromley and Atkins. Dr. Mendez and are excluded.
S.
1
Plants annotated as L. camara (or in a few cases as L. aculeata, see App. B)
were scored for whether trichomes on the abaxial surface were pilose-morph
only, strigose-morph only, setose-morph only, or codominant mixtures of
stri-
and
gose- pilose-morphs (including rare mixtures of strigose- and setose-mor-
when
Between Linnaean camara was
phs). the period, applied primarily
L. to
pilose-morph plants, and the late nineteenth century, specialists began to ap-
name
ply the primarily to the strigose-morph plants or those with mixed
tri-
chome
morphs. For those annotated specimens from the ncotropics, the mixed
SCD
trichome plants are presumed hybrids primarily between the naturalized
and
cultigen native species of the Pilose Group. In the paleotropics, the mi.xed-
trichome plants appeared to be either hybrids between escaped pilose-morph
1)
SCD
and
plants the cultigen, 2) escaped cultivars selected from hybrids between
L
camara and
various species of the Strigose Group, or naturalized plants
3)
consisting of complex spontaneous hybrids between plants of the preceding
two
categories.
As this annotation trend developed, the plants of the Pilose Group began
to be annotated with other later names. In particular, note the quotation of H.
Moldenke Howard
[apparently in personal correspondence] by
(1969):
BR1T.ORG/SIDA 22(1)
390
B0%
70%
60%
(II
O
50%
o
C
<
40%
o
30%
CD
20%
10%
0%
1753-1830 1B31-1905 1906-1930 1931-1955 1956-1980 981 -2005
1
Year
—
-
O
Pilose - Mixed Strigose - Setose
S
+ mrz - Mixed + mrz -"Strlgose mrz
Pilose -^
name
Fig. 5. Plot of portion of data from Table 1, showing historical pattern of specialists' application of the Lantana
camara to pilose-morph taxa (mostly L camara, some L horrida and their hybrids) vs. strigose-morph taxa (mostly L
strigocamara, L nivea, and L scabrida) vs. hybrids between the two groups (mixed morph plants) vs. setose-morph
plants (mostly L birsuta]. "+ mrz" indicates the inclusion of specimens annotated as L camara ^ar.moritziana.
You cannot depend on the accuracy of identification on the labels of plant specimens marked "Lan-
tana camara' in herbaria. have found that a large percentage of such material is actually L.
1
moritziana.L. ^landulosissima, L. scoria, L. ho}'nda,L.arida,L.gluiinosa,Q^c. etc.
Moldenke and specimens
In other words, authorities others) attributed of
(e.g..,
camara to other species names referable to the Pilose Group. Surprisingly, in
L.
the mid 1970s, Moldenke accepted Lopez-Palacios' (1974) reduction in rank
Dr.
L
of moritziana asa variety of Lcflmara. that anomaly excluded from the
If is
L
camara pilose-morph by
analysis, then the application of to strictly plants
taxonomic specialists negligible over the last 50 years.
is
The second line of evidence a survey of the literature (App. A). Adequate
is
descriptions of leaf indument are scarce prior to the 1850s, but those by Medikus
(1775), Sprengel (1825), and Schauer (1847, 1851) all indicate a prevailing accep
The
tance of plants with the abaxial surfaces soft-hairy first publication di-
verging and treating strigose-morph plants as ca mara by Otto and Dietrich
L. is
(1841). This reflected a growing confusion as to what L. camara really By the
is.