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Acta Botanica Mexicana 104: 107-156 (2013)
107
ORIGINS OF MEXICAN MALPIGHIACEAE
William R. andeRson
University of Michigan Herbarium
Ann Arbor, Michigan 48108, USA
wra@umich.edu
ABSTRACT
The approximately 42 lineages of Malpighiaceae currently known in Mexico are
identied and briey described and discussed. All the Mexican lineages have their ultimate
roots in South America, although in some cases the connections are inferred only through
phylogeny and several Mexican genera probably originated in Mexico. All the lineages have
effective adaptations for dispersal except the genus Galphimia, but distributions outside
Mexico and a phylogenetic tree suggest that while many Malpighiaceae reached Mexico
through “stepping-stone” dispersal, some lineages probably arrived as the result of episodes
of long-distance dispersal from South America.
Key words: biogeography, Malpighiaceae, Mexico, phylogeny.
RESUMEN
Se identican y se describen y discuten brevemente los aproximadamente 42 linajes
de Malpighiaceae que se conocen hasta ahora para México. Todos los linajes mexicanos
tienen sus últimas raíces en Sudamérica, aunque en algunos casos las conexiones se ineren
únicamente mediante logenia y algunos géneros mexicanos probablemente se originaron
en México. Todos los linajes tienen adaptaciones efectivas para su dispersión excepto el
género Galphimia, pero las distribuciones fuera de México y un árbol logenético sugieren
que aunque muchas Malpighiaceae llegaron a México mediante el modelo de dispersión
de “piedras de paso”, algunos linajes probablemente arribaron desde Sudamérica mediante
dispersión a larga distancia.
Palabras clave: biogeografía, logenia, Malpighiaceae, México.
Acta Botanica Mexicana 104: 107-156 (2013)
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INTRODUCTION
In a recent review of the literature on the Mexican seasonally dry tropical flora,
Pérez-García et al. (2012, p. 177) stated, “[W]e conclude that treating TDF [tropical
dry forest] as a plant community that has a prominent South American component is
not warranted, and thus this plant formation cannot be considered as being part of the
Neotropical Realm.” They contrasted their conclusion with that of Rzedowski (1991),
who stated (p. 4), “An analysis of the geographical affinities of the phanerogamic flora
of Mexico indicates that its links with the south are about four times more important
that those with the north.” Pérez-García et al. also stated (p. 171) that “More phylogenies
of Mexican species and their sister groups occurring beyond the country’s borders are
needed in order to determine the ultimate sources and relative ages of the components
of this diverse biota . . .” That is the purpose of this paper—as a specialist in the sys-
tematics of the family Malpighiaceae, I propose to describe what is known about the
relationships of the present Mexican representatives of that family, with special empha-
sis on the number of lineages of Malpighiaceae in Mexico and the probable ultimate
origins of those lineages. In doing that I hope not only to contribute to the ongoing
controversy concerning the origins of the Mexican flora, but also to demonstrate what
a powerful tool phylogenetic systematics can be in interpreting biogeographical history.
The Malpighiaceae is a family of tropical and subtropical flowering plants,
comprising approximately 1300 species in 75 genera; about 80% of the genera and
90% of the species occur mostly or entirely in the New World, with the rest native to
the Old World (Anderson et al., 2012). In 1990 I suggested on the basis of morphol-
ogy that the family originated and diversified in South America, spreading eventu-
ally from there to Central America, North America, the West Indies, and the Old
World (principally Africa and Asia). Phylogenetic research published in the last 12
years has broadly supported my 1990 suggestion (Cameron et al., 2001; Davis et al.,
2001; Davis & Anderson, 2010). Today there are more genera and species of Malpi-
ghiaceae in South America than anywhere else (Anderson et al., 2012).
Some of the generic names used in this paper will be unfamiliar to people
acquainted with the Mexican flora. The new names and generic interpretations re-
sult from an ongoing reassessment of relationships in the family. Anyone puzzled
by such unfamiliar names can go to our website (Anderson et al., 2012), where a
nomenclatural database will answer questions about what happened to old familiar
names. The same website contains references to literature on Malpighiaceae (mostly
with pdfs or links to online sources of the original publications) and drawings and
photos of most of the genera mentioned in this paper.
Anderson: Origins of Mexican Malpighiaceae
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MEXICAN LINEAGES OF MALPIGHIACEAE
Here I list and describe briefly the lineages of Malpighiaceae that I can iden-
tify in the Mexican flora. When no source is cited, the reader should assume that
these statements represent my own unpublished observations and opinions.
Adelphia: four species in South America (W. Anderson, 2006), one of which is
also found to the north. Adelphia is embedded in the poorly resolved Hiraea clade,
which is mostly South American (but see also Hiraea and Psychopterys below).
The wholly South American genus Lophopterys is sister to the rest of the Hiraea
clade (Davis & Anderson, 2010), which suggests that the clade originated in South
America.
A. hiraea (Gaertn.) W. R. Anderson (Fig. 1): a vine, wind-dispersed by winged
samaras, extending from northwestern South America throughout Central Amer-
ica into Mexico, where it is collected occasionally in wet forests of Chiapas and
Tabasco; it also occurs in Jamaica.
Aspicarpa/Gaudichaudia (Figs. 2, 3): a large, diverse lineage, mostly Mexican ex-
cept for two species that extend into the United States and two species in Central
America, one of which extends into northwestern South America. Aspicarpa and
Gaudichaudia are embedded in the very large, strongly supported, mostly South
American Stigmaphyllon clade, which is represented in Mexico by six genera [As-
picarpa (including Gaudichaudia), Banisteriopsis, Bronwenia, Cottsia, Diplopterys,
and Stigmaphyllon] (Davis & Anderson, 2010).
In the traditional sense, Aspicarpa comprises several species in southern
South America and several species in North America, but the two groups do not
constitute a monophyletic group (Davis & Anderson, 2010), so this discussion will
ignore the South American species; the type is from Mexico so the name Aspicarpa
will remain with the North American species. There are approximately 3–6 species
of Aspicarpa s. str., growing throughout Mexico from Oaxaca to Sonora, Chihua-
hua, and Coahuila, with two of them occurring also in adjacent areas of the United
States (Arizona, New Mexico, and Texas). They grow in diverse habitats, from oak
and pine-oak forests to shrubby woodlands to desert scrub. They are erect shrublets
or more or less prostrate herbaceous stems from a perennial base that are not or only
very weakly twining. The fruit of Aspicarpa breaks apart into dry nutlets that are
Acta Botanica Mexicana 104: 107-156 (2013)
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Fig. 1. Adelphia hiraea. A. owering branch; B. node to show interpetiolar stipules and large
glands on petiole; C. apex of lamina to show marginal processes; D. circinate ower bud with
one large eccentric gland on one bracteole; E. ower, posterior petal uppermost (eccentric
bracteole gland is behind pedicel, below posterior petal); F. androecium laid out, abaxial
view, the stamen just above the letter “F” opposite posterior petal; G. samara, abaxial view.
Scale bar equivalents: A, 4 cm; B, C, 1.3 cm; D, 8 mm; E, 6 mm; F, 2.7 mm; G, 3 cm. Based
on: A–F, Foster 2313, MICH; G, Davidse et al. 20520, MICH.
unwinged but bear a dorsal crest and usually a lateral crest or rudimentary winglet.
Such mericarps have no obvious adaptation for dispersal (Fig. 2).
Anderson: Origins of Mexican Malpighiaceae
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Fig. 2. Aspicarpa and Gaudichaudia. A–E, A. brevipes (DC.) W. R. Anderson. A. owering
branch; B. base of leaf, abaxial view, and adjacent node with stipule; C. ower, posterior petal
uppermost; D. androecium and gynoecium, stamen at left opposite anterior sepal; E. mericarps,
adaxial view (above) and abaxial view (below). F–K, G. krusei W. R. Anderson. F. owering
branch; G. base of leaf, adaxial view, and adjacent node with stipule; H. ower, posterior petal
uppermost; I. ower with petals removed, lateral view; J. samara, adaxial view, still attached to
receptacle by carpophore; K. samara, abaxial view. Scale bar equivalents: A, 4 cm; B, 4 mm; C,
8 mm; D, 2 mm; E, 8 mm; F, 4 cm; G, 4 mm; H, 8 mm; I, 5 mm; J, K, 8 mm. Based on: A–E,
Pringle 4422, US; F–I, Anderson 12868, MICH; J, K, Koch & Fryxell 8264, MICH.
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Fig. 3. Gaudichaudia galeottiana (Nied.) Chodat. A. branch with chasmogamous owers
and fruits from cleistogamous owers; B, C. enlargement of surfaces of small young leaves,
adaxial side (B) and abaxial side (C); D. chasmogamous ower, posterior petal uppermost;
E. androecium and gynoecium of chasmogamous ower, the stamen at left opposite anterior
sepal; F. samara, adaxial view, still attached to cleistogamous ower by carpophore; G.
samara from cleistogamous ower, abaxial view. Scale bar equivalents: A, 4 cm; B, C, 4 mm;
D, 1 cm; E, 4 mm; F, G, 8 mm. Based on Anderson & Laskowski 4087, MICH.
Gaudichaudia comprises 15–40? species; the definition of species in this
group is made difficult by the widespread occurrence of hybridization and poly-
ploidy (W. Anderson, 1993). Most species are found in southern Mexico, but two
Anderson: Origins of Mexican Malpighiaceae
113
closely related species extend into Central America and one extends into Colombia
and Venezuela. They occur in the same assortment of habitats as Aspicarpa. A few
are erect shrublets, but most are twining herbaceous vines from a perennial base.
The fruit breaks apart into dry winged samaras, well adapted for wind-dispersal
(Figs. 2, 3).
The latest phylogeny of Malpighiaceae (Davis & Anderson, 2010) indicates
that Aspicarpa and Gaudichaudia together form a monophyletic group, but that nei-
ther is monophyletic without the other—the three sequenced species of Aspicarpa
are distributed among the six sequenced species of Gaudichaudia. Therefore, for
the purposes of this paper we should treat Aspicarpa/Gaudichaudia as a single large
and diverse Mexican lineage. On morphological grounds it seems safe to say that
the ancestor of the lineage was probably a plant with a lateral-winged samara that
reached western Mexico via long-distance dispersal from southern South America,
where Camarea, the sister genus, occurs. That ancestor (unlike anything that now
occurs in South America) presumably found open, seasonally dry habitats that it was
able to exploit and into which its descendants were able to expand.
Banisteriopsis: approximately 70 species, all South American (Gates, 1982; Ander-
son & Davis, 2006, 2007). Two species that are not closely related within the genus
(Gates, 1982) have reached Mexico; both are vines and wind-dispersed by winged
samaras. Banisteriopsis is a member of the Stigmaphyllon clade; see note above
under Aspicarpa.
B. elegans (Triana & Planch.) Sandwith: growing in mesic forests from north-
western South America throughout Central America to Guatemala, known in
Mexico from a single collection from a wet forest in Chiapas.
B. muricata (Cav.) Cuatrec. (Fig. 4): very widely distributed in South and
Central America, growing in diverse habitats (wet forests, dry forests, roadside
thickets); known in Mexico from numerous collections in Chiapas and several in
Oaxaca, mostly from tropical deciduous forests.
Bronwenia: ten species, all South American (Anderson & Davis, 2007); two of those
species, both woody vines that are sometimes shrubby and both wind-dispersed by
winged samaras, have reached Mexico. The genus is in the Stigmaphyllon clade (Da-
vis & Anderson, 2010), where it is sister to the rest of the clade; see also note above
under Aspicarpa.
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Fig. 4. Banisteriopsis muricata. A. leafy and owering branches; B. node to show stipules
and abaxial base of leaf; C. ower bud and umbel; D. ower, posterior petal uppermost; E.
posterior petal, lateral-abaxial view; F. portion of androecium, abaxial view, stamen opposite
posterior petal to left; G. gynoecium, anterior style in center; H. apex of style; I. samara; J.
nut of samara, abaxial view, to show carpophore. Scale bar equivalents: A, 3 cm; B, C, 6 mm;
D, 8.6 mm; E, 4.3 mm; F, 1.5 mm; G, 3 mm; H, 1 mm; I, 1.5 cm; J, 6 mm. Based on: A–I,
Schinini 14735, MICH; J, Maguire & Maguire 40235, MICH.
Anderson: Origins of Mexican Malpighiaceae
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B. acapulcensis (Rose) W. R. Anderson & C. Davis (Fig. 5): northwestern
South America, common in Central America, occasional in Oaxaca and Guerrero
in tropical deciduous forest.
B. cornifolia (Kunth) W. R. Anderson & C. Davis: northwestern South Ameri-
ca, common in Central America, occasional in Chiapas, Veracruz, and Oaxaca in
tropical deciduous forest and more mesic forest.
These two species are closely related, but unless one wants to suggest that
they diverged in Mexico (and given their rarity in Mexico that seems unlikely) they
must have reached Mexico independently and thus should be considered two Mexi-
can lineages.
Bunchosia (Fig. 6): approximately 75 species of trees and shrubs, bird-dispersed
by fleshy berries; South America, Central America, Mexico, and the West Indies
(Anderson et al., 2012). The latest phylogenetic tree for Malpighiaceae (Davis &
Anderson, 2010) does not establish where the genus originated, but it is noteworthy
that the sister genus, Thryallis, is limited to southeastern Brazil and adjacent Para-
guay and Bolivia (C. Anderson, 1995). There are at least 20 species of Bunchosia in
southern Mexico, over half of them endemic, the rest also in Central America and
two also in the West Indies; they occupy diverse habitats, from open dry shrubby
associations and tropical deciduous forests to mesic and wet forests. The uniform
morphology of Bunchosia does not permit me to recognize obvious groups at this
time, as in some other genera (e.g., Heteropterys, see below), and so it is impos-
sible to say whether the assemblage of Mexican species descended from a single
immigrant or from several immigrants. Consequently, I am forced for the present
to consider the Mexican species of Bunchosia a single lineage, but it will not be
surprising if future studies establish that Bunchosia in Mexico comprises more than
one lineage.
Byrsonima: approximately 130 species of trees, shrubs, and subshrubs, bird-dis-
persed by fleshy drupe-like fruits. Most species are South American, but there are a
few species in Central America, Mexico, and the Caribbean (including southernmost
Florida). Byrsonima is in the Byrsonima clade; its two sister genera, Blepharandra
and Diacidia, are wholly South American, as is the Acmanthera clade, sister to the
Byrsonima clade (Davis & Anderson, 2010). There are two species of Byrsonima in
Mexico; they are not closely related.
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Fig. 5. Bronwenia acapulcensis. A. branch with owers and fruits; B. node with interpetiolar
stipules; C. base of lamina, abaxial view; D. ower bud and portion of inorescence axis; E.
ower, posterior petal uppermost; F. posterior petal, abaxial view; G. lateral petal, abaxial
view; H. androecium laid out, abaxial view, stamen above letter “H” opposite posterior petal;
I. gynoecium; J. apex of style; K. samara with crest on side of nut; L. samara with winglet on
side of nut. Scale bar equivalents: A, 4 cm; B, 4 mm; C, D, 8 mm; E, 5.7 mm; F, G, 4 mm; H,
I, 2 mm; J, 1 mm; K, L, 2 cm. Based on: A, E–K, Burnham 977, MICH; B–D, Burnham 976,
MICH; L, Almeda 3983, MICH.
Anderson: Origins of Mexican Malpighiaceae
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Fig. 6. Bunchosia mcvaughii W. R. Anderson. A. owering branch; B. detached smaller leaf
from same plant; C. node with epipetiolar stipules; D. abaxial leaf surface to show gland; E.
ower, posterior petal uppermost; F. anther; G. gynoecium; H. fruit; I. cross-section of fruit.
Scale bar equivalents: A, B, 4 cm; C, 4 mm; D, 2 mm; E, 1 cm; F, 2 mm; G, 3.3 mm; H, I, 1
cm. Based on: A–G, McVaugh 25107, MICH; H, I, Pérez J. 1704, MICH.
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B. bucidifolia Standl. (Fig. 7): endemic to the Yucatán Peninsula of Mexico
(Campeche, Quintana Roo, Yucatán) and adjacent Belize and Guatemala, often in
tropical deciduous forest but also in more mesic habitats. The petals are white and
pink or red (probably white turning pink and then red in age), and the connectives
of the anthers do not exceed the locules at the apex; those characters suggest a
relationship with the Caribbean species B. lucida (Mill.) DC., and it is possible
that B. bucidifolia descended from a Caribbean ancestor, but similar characters
are not rare among the many South American species of this genus, so caution is
in order until we can obtain molecular sequences for B. bucidifolia. See also the
discussion of Eoglandulosa below, preceding the Conclusions.
B. crassifolia (L.) Kunth: a species or species-complex that is very widespread
in South America, Central America, southern Mexico, and the West Indies. Relat-
ed species are all South American; the taxonomy of the yellow-flowered species
of Byrsonima is exceedingly difficult and needs a great deal of work. In Mexico
B. crassifolia inhabits diverse habitats, from dry to mesic, and is often prominent
in shrubby savannas.
Calcicola (Fig. 8): two species of shrubs, wind-dispersed by winged samaras, en-
demic to central, seasonally dry Mexico from Coahuila to Oaxaca, growing in open
deciduous woods, thorn-scrub, or desert scrub on rocky or sandy limestone slopes
(Anderson & Davis, 2007). This genus is sister to Malpighia and a large group of Old
World genera, with Mascagnia sister to the whole clade (Davis & Anderson, 2010),
and its morphology reflects that position (Anderson et al., 2012). It is probably de-
scended from a mascagnioid ancestor that came originally from South America; see
discussion below under Malpighia and Mascagnia.
Callaeum (Fig. 9): 11 species of South America, Central America, and Mexico,
woody vines, all but one South American species wind-dispersed by winged samaras
(Johnson, 1986; Anderson et al., 2012). There are three South American species, one
species found only in northern Central America except for a single collection from
Chiapas [C. nicaraguense (Griseb.) Small], one species in southern and southeastern
Mexico and northern Central America [C. malpighioides (Turcz.) D. M. Johnson],
and six species endemic to Mexico and adjacent Texas. The sister to Callaeum is the
South American genus Alicia (Davis & Anderson, 2010), which suggests that the
genus may have originated in South America, but the resolution within Callaeum in
our phylogenetic tree is weak, so it is not possible to say much more about the origins
Anderson: Origins of Mexican Malpighiaceae
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Fig. 7. Byrsonima bucidifolia. A. owering branch; B. stipules on inner base of petiole; C.
abaxial leaf surface to show persistent hairs; D. ower buds; E. ower, lateral view, posterior
petal uppermost; F. stamens, adaxial view (left) and lateral view (right); G. gynoecium; H.
fruit. Scale bar equivalents: A, 4 cm; B, C, 4 mm; D, E, 8 mm; F, G, 2.7 mm; H, 1 cm. Based
on: A–C, Lundell & Lundell 7772, MICH; D–G, Lundell & Lundell 7448, MICH; H, Davidse
et al. 20622, MICH.
of the Mexican species at this time. They are best treated as a single lineage for now,
but the present structure of our phylogenetic tree suggests that Callaeum may have
reached Mexico twice. The widespread and common Mexican species occur most
often in tropical deciduous forests and other open, seasonally dry habitats, but also
in more mesic and even wet forests, especially in southeastern Mexico.
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Fig. 8. Calcicola parvifolia (A. Juss.) W. R. Anderson & C. Davis. A. owering branch;
B. node with interpetiolar stipules; C. edge of lamina with marginal gland, abaxial view
except where turned to show adaxial vesture; D. node and leaf (adaxial view) with axillary
inorescence; E. ower, posterior petal uppermost; F. gynoecium, anterior style in center;
G. samaras, abaxial view (left) and adaxial view (right); H. old ower after fall of samaras,
pyramidal torus in center, surrounded by lobed disc. Scale bar equivalents: A, 4 cm; B, C,
4 mm; D, E, 8 mm; F, 2 mm; G, 1 cm; H, 4 mm. Based on: A, D–F, Breckon et al. 2343,
MICH; B, C, Schoenwetter JSOX-91, MICH; G, García M. 3297, MICH; H, McVaugh
24000, MICH.
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Fig. 9. Callaeum septentrionale (A. Juss.) D. M. Johnson. A. owering branch; B. node with
epipetiolar stipules; C. base of lamina, abaxial view, showing marginal glands; D. ower bud
in umbel; E. ower, posterior petal uppermost; F. petals, abaxial view, posterior petal (left)
and lateral petal (right); G. androecium laid out, abaxial view, the smallest stamen opposite
posterior petal; H. gynoecium; I. distal portion of styles, adaxial view (left) and abaxial view
(right); J. samara, abaxial view; K. samara, abaxial view. Scale bar equivalents: A, 4 cm; B,
C, 4 mm; D, 8 mm; E, 1.3 cm; F, 8 mm; G, H, 2.7 mm; I, 1.3 mm; J, K, 2.7 cm. Based on:
A–I, Daniel 287, MICH; J, Anderson 13314, MICH; K, Anderson & Laskowski 4078, MICH.
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Carolus: six species, four endemic to South America, one endemic to Panama, and
one distributed from South America to Mexico. Carolus is embedded in a clade
comprising two Old World genera (Flabellariopsis and Hiptage) and two South
American genera (Dicella and Tricomaria); it is not morphologically similar to ei-
ther group (Davis & Anderson, 2010; Anderson et al., 2012). The evidence, such as
it is, suggests an origin in South America.
C. sinemariensis (Aubl.) W. R. Anderson (Fig. 10): a woody vine, in most
populations dispersed by winged samaras, in northern South America, Central
America, and at scattered localities across southern Mexico, mostly in mesic for-
ests or secondary associations, but sometimes also in tropical deciduous forest.
Christianella: five species, all vines and wind-dispersed by winged samaras (W.
Anderson, 2006); four are found only in South America, the fifth is found only in
Central America and Mexico. Christianella is sister to the other six genera of the
Christianella clade, all limited to South America and southern Central America ex-
cept for the African genus Flabellaria and the genus Callaeum, which is discussed
above.
C. mesoamericana (W. R. Anderson) W. R. Anderson (Fig. 11): known from
five collections from Panama, Costa Rica, and Guatemala and one from Chiapas;
the habitat, so far as it is known, is “mixed forest.”
Cottsia (Fig. 12): three species endemic to the deserts of northern Mexico and adja-
cent United States, all vines wind-dispersed by winged samaras (Anderson & Davis,
2007; Anderson et al., 2012). Cottsia is sister to the rest of the Aspicarpa clade [Ca-
marea, South American; Janusia, South American; Aspicarpa s. l., South American;
and Aspicarpa s. str./Gaudichaudia, Mexican], but no lineage is known that could
have been directly ancestral to Cottsia. Sister to the Aspicarpa clade is the Cor-
dobia clade (Mionandra, Cordobia, Gallardoa, and Peixotoa), all South American
(Anderson & Davis, 2007; Davis & Anderson, 2010; Anderson et al., 2012). It is
difficult to escape the conclusion that the ancestor of Cottsia must have travelled via
long-distance dispersal from southern South America to northern Mexico, quite in-
dependently of the ancestor of Aspicarpa/Gaudichaudia (Anderson & Davis, 2007).
Diplopterys: 31 species, all but one mostly South American with two of those reach-
ing Panama or Costa Rica (Gates, 1982; Anderson & Davis, 2006). Diplopterys is in
Anderson: Origins of Mexican Malpighiaceae
123
Fig. 10. Carolus sinemariensis. A. owering branch; B. abaxial margin of lamina; C. node
with stipules; D. ower, posterior petal uppermost; E. lateral petal, abaxial view; F. anthers,
adaxial view (left) and abaxial view (right); G. gynoecium; H. apex of style; I. samaras,
abaxial view (left) and adaxial view (right). Scale bar equivalents: A, 4 cm; B, C, 4 mm; D,
7 mm; E, 4 mm; F, 2 mm; G, 2.7 mm; H, 1.3 mm; I, 2 cm. Based on: A–H, Wilbur & Wilbur
2394, MICH; I, Cochrane et al. 12331, MICH.
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Fig. 11. Christianella mesoamericana. A. owering branch; B. node to show stipules; C.
abaxial base of lamina; D. ower bud; E. ower from above, posterior petal uppermost; F.
abaxial views of posterior petal (above) and lateral petal (below); G. androecium laid out,
abaxial view, the shortest stamen opposite posterior petal; H. gynoecium, anterior style in
center; I. apex of style; J. samaras, abaxial view (left) and adaxial view (right); K. hairs on
abaxial surface of samara wing. Scale bar equivalents: A, 4 cm; B, 2 mm; C, 4 mm; D, E, 8
mm; F, 5.7 mm; G, H, 4 mm; I, 1 mm; J, 2 cm; K, 2 mm. Based on: A, C–H, Sytsma & D’Arcy
3278, MICH; B, Matuda 18577, F; I–K, Standley 87301, F.
Anderson: Origins of Mexican Malpighiaceae
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Fig. 12. Cottsia gracilis (A. Gray) W. R. Anderson & C. Davis. A. owering branches; B.
base of leaf and adjacent stem to show stipule and marginal processes; C. detached, relatively
broad leaf; D. ower, posterior petal uppermost; E. androecium and gynoecium, with two
fertile posterior stamens and three anterior staminodes; F. fertile stamen, adaxial view; G.
anterior carpel; H. fruit; I. fruit base and torus with one samara attached by carpophore; J.
seed (left) and nut of samara in longitudinal section with seed removed (right); K. embryos.
Scale bar equivalents: A, 4 cm; B, 4 mm; C, 4 cm; D, 5.7 mm; E, 2.9 mm; F, G, 2 mm; H, 1
cm; I, 5.7 mm; J, K, 4 mm. Based on: A, B, Monson 8, CAS; C–G, Anderson 12552, MICH;
H–K, Anderson & Laskowski 3520, MICH.
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the Stigmaphyllon clade and is sister to Stigmaphyllon (Davis & Anderson, 2010).
The one species not found in South America is endemic to Mexico.
D. mexicana B. Gates: known only from the type, a woody vine collected in
primary wet forest in Veracruz. The fruit is unknown; the peculiar bracts and
bracteoles are of a type found otherwise in six species that grow only in South
America. Among those species two produce wind-dispersed winged samaras and
the other four have the wings greatly reduced and the mericarps probably dis-
persed by water. The ancestor of D. mexicana surely came from South America,
and it will be a surprise if the fruit, when found, does not break apart into wind-
dispersed samaras.
Echinopterys (Fig. 13): two species endemic to Mexico, growing in tropical decidu-
ous forest, desert scrub, and other seasonally dry habitats; shrubs or woody vines,
wind-dispersed by bristly fruits. Echinopterys is sister to the rest of the Bunchosia
subclade of the Bunchosia clade, while sister to that subclade is the Tristellateia sub-
clade, with Henleophytum and Heladena basal in this subclade (Davis & Anderson,
2010; Anderson et al., 2012). Henleophytum is a Cuban endemic that has bristly
fruits similar to those of Echinopterys. Both of those subclades are strongly sup-
ported, so unless bristly fruits are plesiomorphic in the clade, it is difficult to explain
their independent appearance in those two genera. Aside from Henleophytum, no
likely source for the ancestor of Echinopterys is known. However, it is interesting to
note that the sister of Henleophytum is Heladena, a genus of South America, so the
ultimate source of Echinopterys was probably South American. Echinopterys is one
of the most enigmatic and interesting genera of Malpighiaceae endemic to Mexico.
Galphimia (Figs. 14, 15): 26 species, 4 in southern South America and 22 in Mex-
ico, with one of the Mexican species extending into Texas and another extending
into Central America as far south as Nicaragua (C. Anderson, 2007). Galphimia is a
genus of perennial herbs, subshrubs, shrubs, and small trees, occurring throughout
Mexico in diverse habitats, both dry and mesic. The fruit breaks apart into small
dry cocci that are without wings or any other adaptation for dispersal. Some of the
Mexican species (e.g., G. glauca Cav.) have the petals persistent and drying below
the enlarging fruit, suggesting a possible aid to dispersal, but C. Anderson (pers.
comm.) reports that the cocci fall freely at maturity and there is no obvious tendency
for the flower to detach with the fruit intact, so she doubts that the dried corolla is
actually active in dispersal. Besides, such a dry, persistent corolla is not basal in
Anderson: Origins of Mexican Malpighiaceae
127
Fig. 13. Echinopterys. A–M, E. eglandulosa (A. Gray) Small. A. owering branch; B. base of
petiole, adaxial view, to show stipules; C. detached leaf; D. ower bud; E. ower, posterior
petal at upper right; F. posterior petal, abaxial view; G. posterior-lateral petal, abaxial view;
H. anterior-lateral petal, abaxial view; I. stamens, abaxial view (left) and adaxial view (right);
J. gynoecium; K. cross-section of ovary; L. coherent apices of styles; M. mericarp, abaxial
view. N, E. setosa Brandegee. Mature fruit, one mericarp removed, the other two mericarps
separating from torus. Scale bar equivalents: A, 4 cm; B, 2 mm; C, 4 cm; D, 6.7 mm; E–H,
8 mm; I, 2 mm; J, K, 2.7 mm; L, 1 mm; M, N, 4 mm. Based on: A, B, D, Feddema 2370,
MICH; C, E–M, McVaugh 25850, MICH; N, Muller 3281, MICH.
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128
Fig. 14. Galphimia vestita S. Watson. A. habit; B. node with stipules (and detached marginal
leaf gland); C. abaxial leaf surface; D. ower bud, anterior sepal bent down; E. ower; F.
petal, abaxial view; G. anthers, adaxial view (left) and abaxial view (right); H. fruit; I. cocci,
abaxial view (above) and adaxial view (below); J. coccus with half of abaxial wall removed.
Scale bar equivalents: A, 4 cm; B, 4 mm (2 mm); C, 4 mm; D, 5.7 mm; E, F, 5 mm; G, 2 mm;
H–J, 4 mm. Based on: A–D, White 3574 (Sonora), GH; E–J, Wiggins 7471 (Sonora), DS.
Anderson: Origins of Mexican Malpighiaceae
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Fig. 15. Galphimia and Verrucularia. A–F, G. gracilis Bartl. A. owering and fruiting
branch; B. ower, lateral view, posterior petal at upper right; C. stamens, abaxial view (left)
and adaxial view (right); D. gynoecium; E. intact fruit; F. coccus, adaxial view. G–K, V.
glaucophylla A. Juss. G. portion of inorescence; H. stamens, adaxial view (left) and abaxial
view (right); I. gynoecium; J. intact fruit; K. cocci, lateral view (above) and abaxial view
(below). Scale bar equivalents: A, 3 cm; B, 6 mm; C, D, 3 mm; E, 4.2 mm; F, 3 mm; G, 1 cm;
H, I, 1.3 mm; J, K, 4 mm. Based on: A–F, Fryxell & Anderson 3484, MICH; G–I, Harley et
al. 25901, MICH; J, K, Carvalho et al. 1036, MICH.
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130
the genus (Davis & Anderson, 2010), so at most the effectiveness of such dispersal
would be local within Mexico. The sister of Galphimia is Verrucularia (W. R. An-
derson, 1981; Davis & Anderson, 2010), a genus of two species in southeastern and
northern Brazil, which has a similar fruit, so it seems most likely that the ancestor of
Galphimia migrated to Mexico from South America but it is not at all obvious how
that ancestor travelled so far; the Central American species, G. speciosa C. E. An-
derson, is not basal in the clade (Davis & Anderson, 2010), so its distribution seems
most likely to be the result of back-migration from Mexico southward, not a remnant
from the northbound ancestor of Galphimia. The clade of four species in southern
South America, represented in our recent phylogeny (Davis & Anderson, 2010) by
G. brasiliensis (L.) A. Juss. and G. platyphylla Chodat, is sister to a Mexican spe-
cies, G. gracilis Bartl., and that clade is sister to the rest of Galphimia, so Galphimia
in Mexico may represent two lineages, but another way to interpret that tree is that
Galphimia in Mexico is a single lineage, one representative of which migrated to
southern South America and diversified to produce the four species currently there.
Gaudichaudia—See Aspicarpa/Gaudichaudia.
Heteropterys: approximately 150 species, mostly South American but also wide-
spread in Central America, Mexico, and the West Indies; one mostly Caribbean spe-
cies has reached West Africa (Anderson et al., 2012). Heteropterys reached Mexico
at least four times, with the Mexican species falling into three distinct groups that
were recognized by Niedenzu (1928) and our phylogenetic tree (Davis & Anderson,
2010).
H. brachiata (L.) DC., H. cotinifolia A. Juss., H. palmeri Rose, and H. pana-
mensis Cuatrec. & Croat (Fig. 16): these species represent a large South Ameri-
can group of pink- or pink and white-flowered species called ser. Rhodopetalis
by Niedenzu (1928), represented in our phylogenetic tree by H. brachiata and
H. rufula A. Juss. (Davis & Anderson, 2010). All four species are woody vines
(occasionally shrubby) wind-dispersed by winged samaras. Heteropterys brachi-
ata occurs from westernmost South America throughout Central America and is
common across central and southern Mexico, where it grows in diverse dry and
mesic habitats; H. cotinifolia is endemic to Mexico from Sonora to Chiapas, in
tropical deciduous forests; H. palmeri is endemic to Mexico except for one col-
lection from El Salvador, growing in tropical deciduous forests from Sonora and
Sinaloa across southern Mexico to Oaxaca; H. panamensis occurs in more or less
Anderson: Origins of Mexican Malpighiaceae
131
Fig. 16. Heteropterys cotinifolia. A. fruiting branch; B. epipetiolar stipule; C. gland on
petiole; D. gland on abaxial leaf surface; E. ower bud borne on pedicel and peduncle; F.
ower bud enlarged to show keeled petals; G. ower, lateral view, posterior petal uppermost;
H. petals, abaxial views, lateral petal (right) and posterior petal (left); I. androecium laid out,
abaxial view, stamen second from left opposite posterior petal; J. gynoecium, anterior style
in center; K. distal portion of style, lateral view; L. samara. Scale bar equivalents: A, 4 cm;
B, 4 mm; C–E, 8 mm; F, G, 5.7 mm; H, 5 mm; I, 4 mm; J, 2.7 mm; K, 1 mm; L, 2 cm. Based
on: A–D, L, Anderson & Laskowski 4330, MICH; E–K, Flores-Franco et al. 2692, MICH.
Acta Botanica Mexicana 104: 107-156 (2013)
132
mesic forests from Panama to Nicaragua with three collections from Chiapas.
These four species probably represent two lineages in Mexico. The two species
endemic to Mexico were most likely derived from H. brachiata after its arrival in
Mexico, while H. panamensis was probably derived from H. brachiata in Central
America and subsequently invaded Mexico, penetrating only as far as Chiapas.
H. laurifolia (L.) A. Juss. and H. lindeniana A. Juss. (Fig. 17): these belong to
subg. Parabanisteria, a mostly South American group of yellow-flowered spe-
cies with long, eventually revolute sepals; the subgenus is represented in our
phylogenetic tree by H. byrsonimifolia A. Juss. and H. leona (Cav.) Exell (Davis
& Anderson, 2010). H. laurifolia is common and widely distributed in western
South America, Central America, the West Indies, and across southern Mexico
from the Yucatán Peninsula to Sinaloa; it is a woody vine or large shrub some-
times described as a small tree, wind-dispersed by winged samaras, growing in
both seasonally dry and more mesic vegetation. Heteropterys lindeniana is a
shrub or small tree growing usually along streams or in mangrove swamps in the
Yucatán Peninsula of Mexico and Belize and in coastal areas of northern Central
America; it resembles H. laurifolia in most characters but its mericarp has been
modified from the wind-dispersed samara of H. laurifolia by the nearly complete
loss of the dorsal wing, almost certainly as an adaptation to dispersal by water. It
seems likely that the two species represent a single lineage, with H. lindeniana
derived from H. laurifolia in Mexico or adjacent Central America.
H. macrostachya A. Juss.: widely distributed in South America and in Central
America to Nicaragua; not reported from Honduras or Guatemala but rare in Be-
lize and Chiapas, in diverse habitats but most commonly in mesic or seasonally
dry forest, a woody vine (sometimes shrubby) wind-dispersed by winged sama-
ras. This species belongs to a group of yellow-flowered species with the leaves
abaxially densely sericeous, mostly limited to eastern and southern Brazil (ser.
Metallophyllis of Niedenzu, 1928); the group is represented in our phylogenetic
tree by six species, including H. macrostachya (Davis & Anderson, 2010).
Hiraea: at least 75 species (C. Anderson, pers. comm.), mostly South American
but with approximately ten species in Central America, four in Mexico, and one in
the Lesser Antilles, the Mesoamerican species all woody vines wind-dispersed by
winged samaras. The Mexican species fall into three groups, which could represent
either three or four Mexican lineages:
Anderson: Origins of Mexican Malpighiaceae
133
Fig. 17. Heteropterys. A–H, H. laurifolia. A. owering branch; B. abaxial leaf surface,
enlarged to show venation and glands; C. ower bud; D. ower, posterior petal uppermost; E.
posterior petal, adaxial view; F. anther, abaxial view; G. distal portion of style; H. fruit with
two samaras developed. I–L, H. lindeniana. I. adaxial leaf surface; J. abaxial leaf surface,
enlarged to show nely reticulate venation and gland; K. intact fruit from above, with all
three mericarps developed; L. mericarp, lateral view. Scale bar equivalents: A, 3 cm; B, 6
mm; C, 3.75 mm; D, E, 3 mm; F, G, 1 mm; H, 2 cm; I, J, 4 mm; K, L, 1 cm. Based on: A–C,
McVaugh 10176, MICH; D–G, Lott 1020, MICH; H, Téllez 10439, MICH; I, J, Gentle 1309,
MICH; K, L, Whitefoord 2546, MICH.
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134
H. barclayana Benth. and H. reclinata Jacq.: these are sister species with four-
flowered umbels, distinguished from each other by different leaf vesture; they
grow in both mesic and tropical deciduous forests. H. reclinata occurs in north-
western South America, throughout Central America, and in southern Mexico
from the Yucatán Peninsula and Veracruz to Jalisco; H. barclayana is found from
Nicaragua in Central America to Mexico, where it occurs from Chiapas and Ve-
racruz to Nayarit and Sinaloa. They seem most likely to have diverged in Central
America and invaded Mexico independently, in which case they represent two
Mexican lineages, but it is possible that they diverged in Mexico and should be
considered one Mexican lineage.
H. fagifolia (DC.) A. Juss.: this is another species with four-flowered umbels
but morphologically quite distinct from the H. reclinata complex (C. Anderson,
pers. comm.); it is widespread in South and Central America, and known in
southeastern Mexico from the Yucatán Peninsula, Chiapas, Oaxaca, and Vera-
cruz. In Mexico and Central America it grows mostly along rivers and in moist
forests.
H. smilacina Standl. (Fig. 18): this species has multiflowered umbels and be-
longs to a large South American complex; it is known from northwestern South
America (Ecuador and Colombia), Central America, and southeastern Mexico
(Tabasco, Chiapas, Oaxaca, and Veracruz), growing in moist forests.
Lasiocarpus (Figs. 19, 20): approximately four species endemic to tropical decidu-
ous woods in Mexico from Sinaloa to Chiapas; shrubs and trees, wind-dispersed by
bristly mericarps. Lasiocarpus is sister to the morphologically and ecologically simi-
lar genus Ptilochaeta of southern South America (Brazil, Paraguay, Argentina, and
Bolivia); nothing resembling those two genera or closely related to them is found
anywhere between them (Davis & Anderson, 2010). The 2010 phylogenetic tree
shows us that the sister to the Lasiocarpus/Ptilochaeta clade is the morphologically
very different Dinemagonum/Dinemandra clade of Chile. More compelling, per-
haps, is the fact that in Ptilochaeta the flowers are functionally bisexual as in most
Malpighiaceae (including Dinemagonum and Dinemandra), whereas in Lasiocarpus
the flowers are functionally unisexual and the species functionally dioecious (W. R.
Anderson, unpublished data), so there can be little doubt that Ptilochaeta is the less
derived of the two genera and the ancestor of Lasiocarpus must have migrated from
South America to Mexico.
Anderson: Origins of Mexican Malpighiaceae
135
Fig. 18. Hiraea smilacina. A. owering branch; B. petiole with stipules; C. abaxial margin of
lamina, showing hairs and marginal gland; D. parallel tertiary veins, abaxial view of lamina;
E. ower, posterior petal uppermost; F. androecium laid out, abaxial view, stamen above the
letter “F” opposite posterior petal; G. anther, adaxial view; H. gynoecium, anterior style in
center; I. distal portion of style; J. samaras, adaxial view (above) and abaxial view (below).
Scale bar equivalents: A, 4 cm; B, 2 cm; C, 4 mm; D, 1.3 cm; E, 5.7 mm; F, 4 mm; G, 2 mm;
H, 4 mm; I, 1 mm; J, 2.7 cm. Based on: A–I, Foster 2365, MICH; J, Foster 2372, DUKE.
Acta Botanica Mexicana 104: 107-156 (2013)
136
Fig. 19. Lasiocarpus ferrugineus H. S. Gentry. A. staminate owering branch; B. node to
show intrapetiolar stipules; C. staminate ower; D. inorescence in bud with imbricated
scales; E. pistillate ower; F. one carpel, abaxial view; G. fruiting branch; H. one mature
mericarp, adaxial view. Scale bar equivalents: A, 4 cm; B, 4 mm; C, 2.7 mm; D, 4 mm; E,
2.7 mm; F, 2 mm; G, 4 cm; H, 8 mm. Based on: A–C, McVaugh 25933, MICH; D, Anderson
& Anderson 6162, MICH; E, F, McVaugh 25862, MICH; G, H, Anderson & Anderson 6157,
MICH.
Anderson: Origins of Mexican Malpighiaceae
137
Fig. 20. Distribution of Lasiocarpus and Ptilochaeta.
Malpighia (Fig. 21): 50 or more species, shrubs and small trees, all but one bird-
dispersed by fleshy drupe-like fruits; about half the species grow in Mexico and
Central America and the other half in the West Indies, with almost no species
occurring in both areas, the one notable exception being the widespread M. gla-
bra L. In Mexico the species grow mostly in seasonally dry habitats like tropi-
cal deciduous woodland, but some grow also in mesic or even wet forests. The
species of Mexico and Central America that have been sequenced form a single
clade (Davis & Anderson, 2010). Of approximately 19 species in Mexico, 12 are
endemic and seven occur also in Central America; there are four Central Ameri-
can species that do not occur in Mexico. Four Mexican/Central American spe-
cies extend into northwestern South America. Where Malpighia originated is not
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138
Fig. 21. Malpighia rzedowskii W. R. Anderson. A. owering branch, with enlarged hairs from
abaxial surface of lamina (left circle) and adaxial surface (right circle); B. node with interpetiolar
stipules; C. umbel of ower buds, one of four removed; D. ower, lateral view, posterior petal
erect, one posterior-lateral petal removed; E. petals, above left an anterior-lateral petal, the
other two posterior-lateral petals; F. partial androecium laid out, abaxial view, the stamen at left
opposite posterior petal; G. gynoecium, anterior style in middle; H. apex of style; I. fruit, dried
but intact; J. one pyrene of fruit with esh removed, adaxial view (left) and abaxial view (right);
K. cross-section of one pyrene of fruit with esh removed. Scale bar equivalents: A, 4 cm (0.8
mm); B, 4 mm; C, 8 mm; D, 6.7 mm; E, 5 mm; F, G, 3.3 mm; H, 2 mm; I–K, 1.3 cm. Based on:
A–H, Rzedowski & McVaugh 1409, MICH; I–K, McVaugh 16037, MICH.
Anderson: Origins of Mexican Malpighiaceae
139
established by the 2010 phylogeny, but one possibility is that it evolved in dry
vegetation of central Mexico, home to the Mexican endemic Calcicola, which is
sister to Malpighia and to a large group of Old World genera (Davis & Anderson,
2010); see the discussion above under Calcicola. It is also of interest that M. al-
biflora (Cuatrec.) Cuatrec., a species of southern Central America and adjacent
Colombia, is sister to the rest of the Mesoamerican species (Davis & Anderson,
2010), because its fruit, while fleshy as in most species, breaks apart into sepa-
rate mericarps at maturity, as in Calcicola and Mascagnia. Finally, the perplex-
ing species Malpighia leticiana (W. R. Anderson) W. R. Anderson & C. Davis,
known only from its type, deserves special comment. Its immature fruit bears
coriaceous, probably succulent lateral wings, which originally led to its being
described in Mascagnia. The latest phylogeny (Davis & Anderson, 2010) shows it
to be embedded among the Mexican species of Malpighia, for which reason it was
reassigned to Malpighia. Its fruit presumably represents some kind of reversal to
an ancestral condition; further comment is not possible until it is better known.
See the discussion of this species and of the problem of distinguishing Malpighia
from Mascagnia in Anderson & Davis, 2005.
Mascagnia: approximately 40 species of woody vines wind-dispersed by winged sa-
maras, found in South America, Central America, and Mexico, with that range soon
to be enlarged to include the West Indies when the five species currently assigned
to Triopterys are absorbed into Mascagnia (Anderson & Davis, 2013 [in press]).
Mascagnia as a whole (including Triopterys) is strongly supported in our latest tree
(Davis & Anderson, 2010), but major branches within the genus are unsupported.
Nevertheless, it seems likely that the genus originated in South America, where its
closest sister Amorimia is endemic. Mascagnia in South America includes many
species with the petals pink, lavender, or white and a smaller number of species with
yellow petals; both colors are present in Mexico. The five Mexican species represent
three or possibly only two lineages.
M. lilacina (S. Watson) Nied. and M. polybotrya (A. Juss.) Nied. (Fig. 22):
endemic to Mexico, both with petals blue or lavender; they grow in seasonally
dry habitats, M. lilacina limited to northern Mexico (Coahuila, Durango, Nuevo
León, Tamaulipas, and San Luis Potosí) and M. polybotrya to western and south-
ern Mexico (from Michoacán to Oaxaca). The phylogeny in Davis & Anderson,
2010, shows these to be sister taxa, presumably derived from a common Mexican
ancestor, but the lack of resolution at higher nodes in that tree makes it impossible
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140
Fig. 22. Mascagnia lilacina. A. owering branch; B. node showing interpetiolar stipules; C.
abaxial base of lamina to show glands; D. ower bud; E. ower, posterior petal uppermost;
F. gynoecium, anterior style in center; G. samaras, abaxial view (left), adaxial view (right).
Scale bar equivalents: A, 4 cm; B, 2 mm; C, 5.7 mm; D, 8 mm; E, 5.7 mm; F, 4 mm; G, 2 cm.
Based on: A–F, Johnston 8514, GH; G, Johnston 9001, GH.
to say more about the source of that ancestor at this time (but see note below un-
der M. vacciniifolia and M. almedae). The ecology and distribution of M. lilacina
suggest that a similar plant may have been ancestral to Calcicola (see discussion
above).
Anderson: Origins of Mexican Malpighiaceae
141
M. tomentosa C. E. Anderson: C. Anderson (2001) described this yellow-flow-
ered species to accommodate plants distributed from Nicaragua to Chiapas, Oax-
aca, and Veracruz. Its sister species (Davis & Anderson, 2010) are South Ameri-
can, and M. tomentosa seems most likely to have originated in Central America
from an immigrant from South America and spread northward to southeastern
Mexico.
M. vacciniifolia Nied. and M. almedae W. R. Anderson: petals lavender or
pink; mostly found in mesic forests. M. vacciniifolia occurs from Costa Rica
to southeastern Mexico (Chiapas, Oaxaca, and Veracruz), while M. almedae is
known only from two collections in Chiapas; the latter is morphologically very
close to M. vacciniifolia and probably a local derivative of the more widespread
species. The phylogeny in Davis & Anderson, 2010, places M. vacciniifolia with-
out support as sister to the rest of Mascagnia, but that placement seems unlikely
to be correct, given that these species have triangular samaras without a dorsal
winglet and at least M. vacciniifolia climbs by means of adventitious roots, which
is unique in the family. When the phylogeny within Mascagnia is better resolved
it is possible that all four of the Mexican species with lavender or pink petals will
come together in a single lineage, presumably derived from some unknown im-
migrant from South America.
Psychopterys (Fig. 23): eight species of southern Mexico and adjacent Belize and
Guatemala, woody vines growing in matorral, tropical deciduous forests, and wet
forests, wind-dispersed by winged samaras (Anderson & Corso, 2007). This dis-
tinctive lineage is embedded in the mostly South American Hiraea clade (Davis &
Anderson, 2010), but its position in that group is not well resolved and it resembles
other genera of the clade only in the butterfly-shaped samaras. The species probably
all descended from a single immigrant from South America; nothing like it is known
from farther south in Central America.
Stigmaphyllon: 116 species, with a large group (subg. Stigmaphyllon) in the New
World (mostly in South America) and a much smaller group (subg. Ryssopterys) in
Asia (C. Anderson, 1997 & 2011). Stigmaphyllon reached Mexico six times.
S. bannisterioides (L.) C. E. Anderson (Fig. 24): shrubs or woody vines in
beach and mangrove vegetation along the Atlantic coast from Ceará, Brazil, to
Veracruz, Mexico; also in the West Indies and coastal West Africa. Unlike most
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142
Fig. 23. Psychopterys dipholiphylla (Small) W. R. Anderson & S. Corso. A. owering branch;
B. enlargement of abaxial surface of lamina; C. node and base of leaf; D. ower bud with
pedicel, bracteoles, bract, and portion of inorescence axis; E. enlarged ower bud; F. ower;
G. abaxial view of young anther (below) and twisted old anther (above); H. gynoecium;
I. distal portion of style and stigma; J. samara, adaxial view; K. samara, abaxial view; L.
samara with one lateral wing cut away, lateral view to show dorsal wing; M. embryo. Scale
bar equivalents: A, 4 cm; B, 2 mm; C, 8 mm; D, 5 mm; E, 3.3 mm; F, 5 mm; G, 2 mm; H, 5
mm; I, 1 mm, J–L, 2 cm; M, 8 mm. Based on: A–I, Anderson 13819, MICH; J, K, M, Hahn
s.n., P; L, Hinton 7523, NY.
Anderson: Origins of Mexican Malpighiaceae
143
Fig. 24. Stigmaphyllon bannisterioides. A. owering branch; B. base of lamina, abaxial view;
C. posterior petal; D. androecium laid out, adaxial view, the stamen second from left opposite
posterior petal; E. gynoecium, the anterior style in center; F, G. samaras, abaxial views
(above) and lateral views (below), illustrating variation in size and shape of dorsal wing and
lateral ornamentation; H. embryo. Scale bar equivalents: A, 4 cm; B, 4 mm; C, 1 cm; D, E,
4 mm; F–H, 1.3 cm. Based on: A, B, Crosby 42, MICH; C–E, Cremers 7812, MICH; F, H,
Feuillet 898, MICH; G, Zanoni et al. 20185, MICH.
species in the genus, this one is presumably dispersed by water, the wing of the
mericarp being reduced to a rudimentary dorsal crest (Fig. 24). Unfortunately
this species was not included in the last phylogeny of the family (Davis & Ander-
son, 2010), but its present distribution and lack of similarity to other species in
Mexico makes it most likely to have arrived in Mexico independently of the other
Mexican lineages (C. Anderson, pers. comm.).
S. ellipticum (Kunth) A. Juss.: vines wind-dispersed by winged samaras, very
common from northwestern South America throughout Central America to south-
eastern Mexico (the Yucatán Peninsula, Chiapas, Oaxaca, and Veracruz); often in
dry disturbed vegetation, but also growing in more mesic forests. The morphol-
ogy of this species is unlike other Mexican species and suggests an independent
arrival in Mexico (C. Anderson, pers. comm.).
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144
S. lindenianum A. Juss. and S. retusum Griseb.: both woody vines wind-dis-
persed by winged samaras. They are morphologically similar and probably sisters
(C. Anderson, 1997) and have similar but not identical ranges: S. lindenianum
occurs from northwestern Colombia throughout Central America to southeastern
Mexico (the Yucatán Peninsula, Chiapas, Oaxaca, and Veracruz); S. retusum oc-
curs in Central America from Costa Rica northward and penetrates into Mexico
as far as Puebla, Querétaro, and San Luis Potosí. Both grow in a wide variety of
habitats, from seasonally dry matorral to wet forests. Stigmaphyllon lindenianum
is embedded in a South American clade (Davis & Anderson, 2010), and S. retu-
sum can be expected to be placed in the same clade, so they must represent the
result of at least one immigration to Mexico; more likely they reached Mexico
independently and should be counted as two Mexican lineages.
S. pseudopuberum Nied.: vines wind-dispersed by winged samaras, endemic to
Chiapas and adjacent Guatemala, in montane rain forests and other mesic vegeta-
tion; it is morphologically similar to S. puberum (Rich.) A. Juss., a species wide-
spread in northern South America, the West Indies, and Central America reach-
ing Guatemala and Belize but not reported from Mexico (C. Anderson, 1997). It
seems likely that S. pseudopuberum originated in its present area from an immi-
grant that resembled S. puberum.
S. selerianum Nied. (Fig. 25): vines wind-dispersed by winged samaras, endem-
ic to tropical deciduous forests in Oaxaca and Chiapas; morphologically isolated,
unlike any species in Central America, presumably derived from an immigrant
from South America (C. Anderson, 1997 & pers. comm.).
Tetrapterys: In the broad sense still in use this is a genus of approximately 70 species
of woody vines (sometimes shrubby), wind-dispersed by winged samaras, centered
in South America but with species in Central America, Mexico, and the West Indies.
Unfortunately, Tetrapterys is clearly not monophyletic, as shown in the phylogeny
in Davis & Anderson, 2010. The Mexican species represent five lineages, which will
be discussed here in subgroups:
Tetrapterys s. str.: three species that clearly belong together in the same genus
but that arrived independently in Mexico:
T. discolor (G. Mey.) DC.: a variable but distinctive species or species
complex, widespread in South America, that presumably migrated north
Anderson: Origins of Mexican Malpighiaceae
145
Fig. 25. Stigmaphyllon selerianum. A. large leaf; B. owering branch; C. leaf; D. petiole
with glands; E. node with stipules; F. inorescence; G. ower bud; H. bract from below
inorescence, abaxial view; I. ower, posterior petal uppermost; J. androecium laid out,
adaxial view, third stamen from right opposite posterior petal; K. gynoecium, anterior style in
center; L. apex of anterior style, lateral view. Scale bar equivalents: A–C, 4 cm; D, E, 8 mm;
F, 1.3 cm; G, 1 cm; H, 4 mm; I, 1.3 cm; J, K, 4 mm; L, 2 mm. Based on: A, I–L, Anderson &
Anderson 5555, MICH; B–H, Reyes García 333, MICH.
Acta Botanica Mexicana 104: 107-156 (2013)
146
into Central America, where it occurs mostly in wet forests in most coun-
tries, and thence into southeastern Mexico (Tabasco, Chiapas, Oaxaca, and
Veracruz).
T. schiedeana Schltdl. & Cham. (Fig. 26): throughout Central America
from Costa Rica (with one collection from adjacent Panama) northward, very
common in Mexico south of Sinaloa and Tamaulipas; mostly in wet or mesic
forests but also in tropical deciduous forests. The relationships of this species
within Tetrapterys s. str. are not obvious, and the representation of the genus
in our 2010 phylogeny is too poor to help with that problem, so for now we
will have to be content with saying that its ancestor probably came from
South America, but whether the species originated in Mexico and migrated
south or originated in Central America and migrated north into Mexico is
unknown.
T. tinifolia Triana & Planch.: widespread in northern South America and
throughout Central America into southeastern Mexico (Chiapas, Oaxaca, and
Veracruz), in wet forests; sister to T. goudotiana Triana & Planch. of Central
America and Colombia and many similar species of South America.
Tetrapterys s. l. 1 (Fig. 27): six species that occur in Mexico and/or northern
Central America: T. argentea Bertol.: mesic forests of southeastern Mexico (Chi-
apas), Guatemala, and El Salvador; T. cotoneaster A. Juss.: endemic to tropical
deciduous forests of southern Mexico (Guerrero, Morelos, Oaxaca); T. hetero-
phylla (Griseb.) W. R. Anderson: tropical deciduous forests of southern Mexico
(Guerrero, Oaxaca, Chiapas), Guatemala, and El Salvador; T. mexicana Hook. &
Arn.: endemic to pine-oak forests and tropical deciduous forests of northwestern
and southern Mexico from Sinaloa to Oaxaca; and T. sp. nov. ined., endemic to
oak woodlands in east-central Mexico (Hidalgo, Querétaro, San Luis Potosí, and
Veracruz). Those five species, plus a sixth undescribed species that is endemic to
Honduras, form a clade that resembles Tetrapterys s. str. in some characters but
differs in others (especially the strictly marginal leaf glands). Unfortunately, none
of these species was included among those sequenced for our recent phylogeny
(Davis & Anderson, 2010), so the phylogenetic placement of this clade remains
to be determined, but for the purpose of this paper it is enough to say that this
clade is surely a single lineage in Mexico. Nothing like these plants occurs far-
ther south in Central America or in northern South America; the closest possible
Anderson: Origins of Mexican Malpighiaceae
147
Fig. 26. Tetrapterys schiedeana. A. owering branch; B. node with stipules and adaxial
view of leaf bases; C. marginal leaf gland, abaxial view; D. umbel; E. ower bud; F. ower,
posterior petal uppermost; G. stamens, adaxial view (left) and abaxial view (right); H.
gynoecium, anterior style in center; I. samaras, adaxial view (left) and abaxial view (right).
Scale bar equivalents: A, 4 cm; B, 4 mm; C, 2 mm; D, 1 cm; E, F, 4 mm; G, H, 2 mm; I, 1.3
cm. Based on: A–H, McVaugh 19046, MICH; I, McVaugh & Koelz 1305, MICH.
Acta Botanica Mexicana 104: 107-156 (2013)
148
Fig. 27. Tetrapterys mexicana. A. leafy branch; B. base of leaf, abaxial view, to show marginal
glands; C. node to show interpetiolar stipules; D. owering branch; E. ower, posterior petal
uppermost; F. anther, lateral view; G. gynoecium, anterior style to right; H. distal portion of
posterior-lateral style, adaxial view; I. distal portion of anterior style, lateral view (left) and
adaxial view (right); J. samara, abaxial view. Scale bar equivalents: A, 4 cm; B, 1.3 cm; C,
4 mm; D, 4 cm; E, 8 mm; F, 2 mm; G, 2.7 mm; H, I, 1.3 mm; J, 8 mm. Based on: A–C, J,
McVaugh 11931, MICH; D–I, Anderson & Anderson 6137, MICH.
Anderson: Origins of Mexican Malpighiaceae
149
relatives are probably T. buxifolia Cav. on Hispaniola and T. bracteolata Griseb.
in southeastern Brazil. Whatever the ancestor of this clade may have been, it
must have reached northern Central America or Mexico from some considerable
distance.
Tetrapterys s. l. 2 (Fig. 28): two sister species (T. arcana C. V. Morton and T.
seleriana Nied.) in diverse habitats (dry scrub to forest) from Nicaragua to the
Yucatán Peninsula, Oaxaca, and Veracruz. Those species belong to a clade that is
very strongly supported in our phylogeny (Davis & Anderson, 2010), where they
are represented by T. arcana; it is morphologically quite distinct from Tetrapterys
s. str. and all the other species are South American, so it seems clear that the
Mexican plants represent a single lineage derived from South American plants
that moved north into Central America; such species were presumably similar
to T. styloptera A. Juss., which is widespread in northern South America and
reaches in Central America to Nicaragua.
Summary of Mexican lineages of Malpighiaceae: Adelphia, 1; Aspicarpa/Gaudi-
chaudia, 1; Banisteriopsis, 2; Bronwenia, 2; Bunchosia, 1, possibly more; Byrsoni-
ma, 2; Calcicola, 1; Callaeum, 1, possibly 2; Carolus, 1; Christianella, 1; Cottsia, 1;
Diplopterys, 1; Echinopterys, 1; Galphimia, 1, possibly 2; Heteropterys, 4; Hiraea,
4, possibly 3; Lasiocarpus, 1; Malpighia, 1; Mascagnia, 3, possibly 2; Psychopterys,
1; Stigmaphyllon, 6; Tetrapterys, 5. TOTAL: 42, possibly several more or fewer.
ADAPTATIONS FOR DISPERSAL IN MEXICAN LINEAGES OF
MALPIGHIACEAE
Almost all the Malpighiaceae that reached Mexico had some kind of effective
adaptation for dispersal. The majority of the species in Mexico now have wind-
dispersed mericarps (winged or bristly), or probably evolved from wing-fruited an-
cestors by loss of the wing in Mexico (Aspicarpa spp.). Three genera have fleshy,
bird-dispersed fruits, with Bunchosia and Byrsonima undoubtedly dispersed from
South and Central America by means of such fruits; Malpighia, which seems to be
of Mexican origin, may be derived from an ancestor with a mascagnioid samara (see
the discussion above).
Two Mexican species grow near water and have mericarps with reduced
wings that are surely dispersed by water. One, Heteropterys lindeniana, seems
Acta Botanica Mexicana 104: 107-156 (2013)
150
Fig. 28. Tetrapterys seleriana. A. owering branch; B. marginal leaf gland, abaxial view; C.
minute stipule beside base of petiole; D. ower bud on pedicel subtended by peduncle with
subapical bracteoles, the peduncle in turn subtended by a bract; E. bracteoles with marginal
glands near base; F. ower, posterior petal uppermost; G. androecium laid out, adaxial view
above, abaxial view below, the stamen fth from left opposite posterior petal; H. gynoecium,
anterior style in center; I. distal portion of style; J. samaras, adaxial view (left) and abaxial
view (right); K. enlargement of dorsal wings on samara; L. embryo. Scale bar equivalents:
A, 4 cm; B, C, 4 mm; D, 8 mm; E, 4 mm; F, 6.7 mm; G, H, 2.7 mm; I, 0.8 mm; J, 1 cm; K, 4
mm; L, 2.2 mm. Based on: A–I, Cabrera C. 5549, MICH; J–L, Matuda 3099, MICH.
Anderson: Origins of Mexican Malpighiaceae
151
likely to have evolved from H. laurifolia in the Yucatán Peninsula or adjacent
Central America, while the other, Stigmaphyllon bannisterioides, is widespread
in the Caribbean region and probably followed the Atlantic coast from Brazil to
Mexico.
One Mexican species, Diplopterys mexicana, is unknown in fruit. Its fruit
seems most likely to be a winged samara, but it could also be adapted for dispersal
by water like several related species in South America.
That brings us to Galphimia, possibly the most intriguing clade of Malpighia-
ceae in Mexico. The cocci of this genus (like those of its sister, Verrucularia) have
no obvious adaptation for dispersal (Fig. 15), unless the persistent dried petals of
some Mexican species aid in local dispersal (see discussion above). No South Ameri-
can species of Galphimia now occurs north of the Amazon or west of Bolivia, al-
though one of the two species of Verrucularia does occur in Amazonas, Brazil, just
south of Venezuela (W. Anderson, 1981). Moreover, the only species of Galphimia
in Central America (G. speciosa) is not basal in the genus and is probably the result
of extension of the range of a Mexican species southward. The genus Galphimia is
clearly monophyletic (Davis & Anderson, 2010), and the strongly supported clade
of Galphimia+Verrucularia is well-supported as sister to the Lophanthera+Spachea
clade, a group of South America, Panama, and Costa Rica (Davis & Anderson,
2010). I see no way to escape the conclusion that the cocci of the ancestor of Galphi-
mia traveled by long-distance dispersal from South America to Mexico, in spite of
their unadorned exocarp, and there is no modern evidence that that dispersal was ac-
complished via a series of intermediate populations in northwestern South America
or Central America.
POSSIBLE SOURCES OF MALPIGHIACEAE NOW IN MEXICO
The evidence available indicates that the Malpighiaceae originated and un-
derwent their early diversification in South America, probably at a time when dis-
semination from South America to other continents was difficult (W. Anderson,
1990; Cameron et al., 2001; Davis et al., 2001; Davis & Anderson, 2010), so it is not
really surprising to find that the ultimate source for all the Mexican lineages de-
scribed above was South America. That statement, however, is an oversimplification
that could lead to misinterpretation. The reality seems to be that the approximately
42 lineages of Malpighiaceae now found in Mexico can be arranged on a gradient
from ones that represent nothing more than an extension of the range of a South
Acta Botanica Mexicana 104: 107-156 (2013)
152
American species north into Mexico (e.g., Heteropterys macrostachya) to ones with
roots presumably in South America but without any close relative in South America
at this time (e.g., Echinopterys). The paragraphs below put the Mexican lineages into
four groups that represent the two extremes of that gradient and two intermediate
stations on the gradient.
1. Seventeen of the Mexican lineages of Malpighiaceae represent a straight-
forward range extension of a South American species or group of species that is
embedded in a South American clade, leaving no basis for doubt that the Mexican
plants originated in South America. 11 of these lineages have penetrated no farther
than southeastern Mexico (defined here as all of Mexico from Veracruz and Oaxaca
eastward) while the other six are more widespread in Mexico.
Adelphia hiraea: South America to Tabasco and Chiapas
Banisteriopsis elegans: South America to Chiapas
Banisteriopsis muricata: South America to Chiapas and Oaxaca
Bronwenia acapulcensis: South America to Oaxaca and Guerrero
Bronwenia cornifolia: South America to southeastern Mexico
Byrsonima crassifolia: South America to southern Mexico
Carolus sinemariensis: South America to southern Mexico
Heteropterys brachiata group: South America to central and southern Mexico
Heteropterys laurifolia group: South America to southern Mexico
Heteropterys macrostachya: South America to Chiapas
Hiraea reclinata: South America to southern Mexico
Hiraea fagifolia: South America to southeastern Mexico
Hiraea smilacina: South America to southeastern Mexico
Stigmaphyllon bannisterioides: South America to Veracruz
Stigmaphyllon ellipticum: South America to southeastern Mexico
Tetrapterys discolor: South America to southeastern Mexico
Tetrapterys tinifolia: South America to southeastern Mexico
2. The following nine species or species groups are Central American and
Mexican lineages that do not or barely occur in South America but have unmistak-
able roots in South American clades of the same genus. Seven of these lineages
have reached only southeastern Mexico, while the other two are more widespread in
southern Mexico:
Christianella mesoamericana: Central America to Chiapas
Heteropterys panamensis: Central America to Chiapas
Anderson: Origins of Mexican Malpighiaceae
153
Hiraea barclayana: Central America to northwestern Mexico
Mascagnia tomentosa: Central America to southeastern Mexico
Mascagnia vacciniifolia group: Central America to southeastern Mexico
Stigmaphyllon lindenianum: Central America and adjacent Colombia to south-
eastern Mexico
Stigmaphyllon pseudopuberum: Guatemala and Chiapas
Stigmaphyllon retusum: Central America to southern Mexico
Tetrapterys seleriana group: Central America to southeastern Mexico
3. The following nine lineages are Mexican, some also in Central America;
they have congeners but no obvious close relatives in South America:
Bunchosia spp. group: Central America to Mexico, with one species also in
the West Indies
Byrsonima bucidifolia: northern Central America and the Yucatán Peninsula
of Mexico
Callaeum spp. group: Central America to Mexico
Diplopterys mexicana: Veracruz, Mexico
Galphimia spp. group: Mexico with one species also in Central America
Mascagnia lilacina group: southern to northeastern Mexico
Stigmaphyllon selerianum: southeastern Mexico
Tetrapterys mexicana group: northern Central America and Mexico
Tetrapterys schiedeana: Central America to southern Mexico
4. The following seven lineages seem likely to have evolved in Mexico, al-
though some of them have extended their ranges southward and one (Malpighia) is
widespread in the West Indies. The claim that their ultimate origin must have been
in South America is based on their phylogenetic relationships (Davis & Anderson,
2010), not on their present distribution. For each lineage see the discussion above.
Aspicarpa/Gaudichaudia: Mexico and adjacent United States and Central
America, with one derived species in northwestern South America
Calcicola: central Mexico
Cottsia: northern Mexico and adjacent United States
Echinopterys: Mexico
Lasiocarpus: southern Mexico
Malpighia: Mexico, Central America, northwestern South America, and the
West Indies
Psychopterys: southern Mexico and adjacent Belize and Guatemala
Acta Botanica Mexicana 104: 107-156 (2013)
154
THE FOSSIL GENUS EOGLANDULOSA
Reliably identifiable macrofossils of Malpighiaceae are few, but one fossil of
what certainly appears to be a flower of Malpighiaceae dates from the Eocene and
was described as Eoglandulosa warmanensis Taylor & Crepet (Taylor & Crepet,
1987). The fossil was found in Tennessee, USA, far north and east of Mexico; the
family now extends barely north of Mexico, with two species in Arizona and New
Mexico and six species in Texas; one Caribbean species is native to southernmost
Florida. Two aspects of Eoglandulosa are informative about its most likely rela-
tionships: 1) All five sepals bear paired abaxial glands, whereas most of the more
derived genera of the family have paired glands on the four lateral sepals and the
anterior sepal is eglandular (or all the sepal glands are much reduced or absent). 2)
The pollen is radially symmetrical and tricolporate, as in the genera near the base
of our phylogeny (Davis & Anderson, 2010; Anderson et al., 2012); more derived
genera have globally symmetrical pollen with very diverse numbers and distribu-
tions of colpi and apertures. Of the genera of Malpighiaceae now in Mexico, only
Byrsonima has that combination of calyx glands and pollen type. Of the two species
of Byrsonima currently in Mexico, B. bucidifolia is the one whose morphology sug-
gests a possible ancestry in the Caribbean, with one possible sister being B. lucida,
which is native in southernmost Florida today (see discussion of Byrsonima above).
Lacking information on the age of Byrsonima, we cannot say with any confidence
that Eoglandulosa was a Byrsonima or something related to Byrsonima, but it is
tempting to speculate that Eoglandulosa may indicate an early invasion of North
America by Byrsonima, perhaps travelling via the Caribbean. Beyond that, I cannot
see that Eoglandulosa sheds much light on the origins of the lineages of Malpighia-
ceae currently in Mexico.
CONCLUSIONS
The Malpighiaceae in Mexico today represent approximately 42 lineages, all
with their ultimate roots in South America, so this family, at least, gives strong sup-
port to Rzedowski’s generalization (1991) about the affinities of the Mexican flora
with plants of South America. Some of those lineages probably arrived recently,
diverging little in Mexico and, in many cases, not or hardly penetrating beyond the
relatively mesic vegetation of southeastern Mexico into the seasonally dry vegeta-
tion types that abound to the west and north. Other lineages (e.g., Aspicarpa/Gaudi-
Anderson: Origins of Mexican Malpighiaceae
155
chaudia and Galphimia) probably reached Mexico long ago, if one can judge from
their ecological success and phyletic diversification throughout much of Mexico.
Indeed, several lineages must have originated in Mexico from unknown ancestors
whose South American source can be inferred only from interpretation of a phy-
logenetic tree. Most of the Malpighiaceae that reached Mexico have (or originally
had) effective adaptations for dispersal, the one notable exception to that statement
being the genus Galphimia; that is as one would expect in a suite of plants that had to
migrate short to long distances to reach Mexico. Even so, one of the most interesting
conclusions I can draw from the data presented above is that in a number of cases
there is no evidence that the ancestors now in Mexico reached that country by means
of gradual extension of terrestrial ranges (“stepping-stone” dispersal). The evidence
now available indicates that there have been a number of long-distance dispersal
events from South America to Mexico; for discussion of the importance of long-
distance dispersal in Neotropical plant lineages, see Christenhusz & Chase, 2013.
ACKNOWLEDGEMENTS
The beautiful drawings in this paper are all from the talented pen of Karin
Douthit. I am indebted to Christiane Anderson and Charles Davis for many years
of collaboration and exchange of information and ideas. Socorro González Elizondo
very kindly helped by translating my abstract from English into Spanish.
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Recibido en febrero de 2013.
Aceptado en abril de 2013.
