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Studies in Neotropical Paleobotany. XIII. An Oligo-Miocene Palynoflora from

Simojovel (Chiapas, Mexico)

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American Journal of Botany 86(1): 17–31. 1999.

STUDIES IN NEOTROPICAL PALEOBOTANY. XIII.

AN OLIGO-MIOCENE PALYNOFLORA FROM SIMOJOVEL
(CHIAPAS, MEXICO)1
ALAN GRAHAM
Department of Biological Sciences, Kent State University, Kent, Ohio 44242

A plant microfossil assemblage of 24 identified and five unknown pollen and spore types is reported from the early
Miocene La Quinta Formation near Simojovel, Chiapas, Mexico. The taxa group into seven paleocommunities representing
versions of the modern mangroves (Pelliciera, Rhizophora), swamp and lowland riparian forest (Ceratopteris, Crudia,
Pachira), tropical rain forest (Selaginella, cf. Antrophyum, Pteris, Sphaeropteris/Trichipteris, cf. Aguiaria, Crudia, Guarea,
Pachira), lower montane rain forest (Alfaroa/Oreomunnea, possibly Eugenia), evergreen cloud forest [Picea, Pinus, Podo-
carpus, Ericaceae (possibly Cavendishia/Vaccinium)], evergreen seasonal forest (Hymenaea, Ilex, possibly Eugenia), and
tropical deciduous forest (Cedrela). Elements of arid and high-elevation habitats were absent or few, and northern temperate
elements (Picea, Pinus?) were few or rare. Paleoelevations are estimated at 1000–1200 m (present average 2000 m, maxi-
mum 3004 m), MAT (mean annual temperature) at least as warm as the present 248C, and annual rainfall near the present
;2500 mm but more evenly distributed. The La Quinta (Simojovel) and other Tertiary floras from the region reflect a trend
toward higher altitudes, more seasonal rainfall, cooling tempertures, increased introduction of cool-temperate elements from
the north after ;15 Ma (million years), and increased introduction of tropical elements from the south after completion of
the isthmian land bridge ;3.5 Ma ago.

Key words: Chiapas; early Miocene; La Quinta Formation; Mexico; palynoflora.

The state of Chiapas in southern Mexico (Fig. 1) en-
compasses a region important to understanding the north-
ern Latin American biota and the events that have shaped
its history. It includes most of the tropical part of Mexico,
but there is sufficient topographic diversity to support an
extensive temperate vegetation. It is positioned just north
of the Central American land bridge and is, therefore,
along the terrestrial migratory pathway for plants and an-
imals between North and South America. The geologic
strata at several sites in Chiapas preserve a record of
climatic change, tectonic evolution, paleovegetation, and
migrations through the region during Cenozoic time.
A number of unpublished theses and dissertations, ab-
stracts, and reports of individual fossil plants are avail-
able for Chiapas. However, there are relatively few de-
tailed studies on extensive paleofloras that focus on es-
tablishing the biological affinities of the specimens and
that provide descriptions and illustrations adequate to
confirm the identifications. Some reports include an array
of fossil plants presently distributed in tropical south-
eastern Asia, temperate eastern Asia, and Africa (e.g., the
early-to-middle Miocene Méndez flora of northern Chia-
pas; Palacios Chávez and Rzedowski, 1993). Although
the southeast Asian mangrove palm Nipa was present in
southwestern Texas during the Eocene (Westgate and
Gee, 1990), and the temperate east Asian Eucommia is
documented for the late Tertiary of Puebla, Mexico (Ma-
gallón-Puebla and Cevallos-Ferriz, 1994), the presence of
other exotics needs to be verified.
Megafossils reported from the region are leaflets and
1 Manuscript received 25 November 1997; revision accepted 20
March 1998.
The author thanks Rodolfo Palacios Chávez for cooperation in the
field work and Shirley A. Graham for reading the manuscript. Research
supported by NSF grant DEB-9206743.
possible sepals of Hymenaea, a fossil flower of Tapirira,
and a leaf of Acacia from Oligo-Miocene amber at Si-
mojovel (Miranda, 1963). The amber has been shown by
infrared spectrophotometry to be derived from Hymenaea
(Langenheim, 1966). Plant microfossils have been de-
scribed from the lower to middle Miocene Méndez For-
mation near Pichucalco/Raudales de Malpaso (Palacios
Chávez and Rzedowski, 1993; Fig. 1) from the middle to
upper Miocene Ixtapa Formation along Highway 195
near Ixtapa (Martı́nez-Hernández, 1992) and from a se-
ries of exposures of Eocene to latest Oligocene age from
the region of Simojovel (Tomasini-Ortiz and Martı́nez-
Hernández, 1984).
Pollen and spore assemblages have also been reported
from other outcrops of Oligo-Miocene age near Simojov-
el by Langenheim, Hackner, and Bartlett (1967). That
study was concerned primarily with establishing the de-
positional environment for amber inclusions within the
Simojovel sediments. Pollen of the mangrove Rhizophora
was studied in detail, while other palynomorphs were
mentioned only briefly. Some additions to the list of pa-
lynomorphs also were made by Graham and Palacios
Chávez (1996). In this study a more complete inventory
is presented (Table 1), along with an estimate of the pa-
leocommunities and the Oligo-Miocene climate and pa-
leophysiography.
MATERIALS AND METHODS
The fossil palynomorphs were isolated according to standard pro-
cessing procedures (Gray, 1965; Traverse, 1988) and recently outlined
for Neogene sediments from Guatemala (Graham, 1998). The samples
were macerated in a mortar and pestle and placed successively in HCl
(1 h), HF (1 h), and HNO3 (overnight), with four rinses of distilled
water between each acid treatment. The residues were actolyzed (nine
parts of acetic anhydride to one part of concentrated H2SO4), with rinses
in glacial acetic acid before and after acetolysis, mounted unstained in

17
18 AMERICAN JOURNAL
Fig. 1. Physiographic diagram of Chiapas (after Frost and Langenheim, 1974, fig.1). Used with permission of Northern Illinois University Press.
glycerine jelly, and sealed with CoverBond. The specimens were ex-
amined and photographed at 4003 magnification using a Leitz Ortho-
plan Photomicroscope and black and white Panatomic X film. Identi-
fications were made through comparisons with a pollen and spore ref-
erence collection of ;24 000 slides, and through published illustrations
and descriptions. Location of the specimens on the slides is by England
Slide Finder (ESF) coordinates. Slides, residues, unprocessed samples,
negatives, and duplicate prints are in the palynological collections at
Kent State University.
LOCALITIES, AGE, AND STRATIGRAPHY
The plant microfossils are from two localities in Chia-
pas, Mexico near Simojovel (178 N; Fig. 1). One site is
at the east end of the village, just east of the cemetery,
and adjacent to a large landslide area. At the time (March/
April, 1993), the steep landslide surface was a cultivated
field. The exposure is an ;8 m vertical section of sand-
stone with lenses of lignite and lignitic sandstone (Figs.
2, 3). The thickest lens, near the base of the outcrop, is
;10 cm, and it yielded the most diverse and well-pre-
served palynomorphs (Locality A, sample 4). In addition,
OF BOTANY [Vol. 86
other collections were made at this site by J. Langenheim
[D-621 in Langenheim, Hackner, and Bartlett (1967), and
in the Harvard University (HU) collections]. Both sets of
slides were used in this study.
The second locality is from the Pabuchil (Rancho Ale-
gre) landslide area 3.2 km northwest of Simojovel. This
site is designated PL-7 in the HU collections, and slides
from these samples also were used in this study. The
locality is similar to that described for A-4(KE)/D-
621(HU), but it is a smaller outcrop. Although these sites
are roadside or near roadside exposures, access to most
outcrops in the area now requires permission from the
local Lacandon/Tzotzil inhabitants who have mining
rights to the amber, which occurs as inclusions in the
sandstone.
The area around Simojovel is in the Front Ranges and
High Plateau physiographic province of Frost and Lan-
genheim (1974; Fig. 1). In the diagram presented by
Breedlove (1981), based on Müllerried (1957), this prov-
ince is divided into the Central Plateau (adjacent to the
Central Depression), Eastern Highlands, and Northern
January 1999] GRAHAM—OLIGO-MIOCENE PALYNOFLORA FROM CHIAPAS, MEXICO 19

TABLE 1. Plant fossils from the early Miocene La Quinta Formation, scribed by Frost and Langenheim (1974; Fig. 4). In these
Simojovel, Chiapas, Mexico. Figures are percentages based on a summaries, the strata at the sites studied here, and others
count of 200 specimens from HU preparation D-621 (Simojovel
landslide area), excluding Rhizophora, which constituted .95% of
mentioned in Langenheim, Hackner, and Bartlett (1967),
the sample. are assigned to the transition between the late Oligocene
and early Miocene. They belong to the La Quinta For-
Megafossils mation, which is divided by Allison (1967) into the Cam-
Anacardiaceae ino Carretero, Florida Limestone, and Finca Carmitto
Tapirira durhamii members. In the stratigraphic charts of Frost and Lan-
Leguminosae genheim (1974, p. 29), lignites are described only from
Caesalpinioideae the Florida Limestone Member. They note (p. 31) that
Hymenaea
Mimosoideae
‘‘the Florida Limestone Member contains a coral fauna
Acacia which is unequivocally Early Miocene’’ and ‘‘thus the
Microfossils Florida Limestone Member and the uppermost part of the
Selaginellaceae Camino Carretero Member are Early Miocene.’’ These
Selaginella subdivisions of the La Quinta Formation, however, are
Monolete fern spore not evident at all sites, and there have been no subsequent
Type 1
Type 2
detailed stratigraphic studies published for the region.
Cyatheaceae The palynomorphs described here are probably part of
Sphaeropteris/Trichipteris the Florida Limestone Member of the La Quinta For-
Pteridaceae mation and are considered early Miocene in age. The
Ceratopteris 4 composition of the Simojovel flora is given in Table 1,
Pteris and the age, location, and references to other palynofloras
Vittariaceae
cf. Antrophyum 8
mentioned in the text are given in Table 2.
Pinaceae
Picea COMPOSITION
Pinus?
Podocarpaceae Selaginella (Selaginellaceae; Fig. 5). Oblate-spheroi-
Podocarpus dal; trilete, laesura 10–12 mm long, extending to spore
Palmae
cf. Crysophila 4
margin, inner margin entire; echinate, spines 4–5 mm in
Type 1 13 length, wall 2–3 mm thick; size 36–38 mm. Loc. Pl-7,
Aquifoliaceae slide 1, ESF H-42,3. Previous records: Cucaracha, Cu-
Ilex lebra, Gatun, Gatuncillo, La Boca, Padre Miguel, Paraje
Asteraceae Solo, San Sebastian, and Uscari formations/group/se-
Bombacaceae quence.
cf. Aguiaria type 0.5
Pachira-type 0.5
Monolete fern spore type 1 (Fig. 6). Reniform; mon-
Ericaceae olete, laesura straight to slightly undulating, situated
Juglandaceae along concave margin of spore, 34–36 mm long, extend-
Alfaroa/Oreomunnea 5 ing ;⅔ spore length, inner margin entire; verrucate, ver-
Leguminosae rucae low, irregular, ;4–6 mm in diameter; size 65–67
Caesalpinioideae 3 50–52 mm. Loc. Pl-7, slide 1, ESF G-43,1–3.
Crudia 2
Meliaceae
Spores of this generalized type occur in virtually all
Cedrela Cenozoic palynofloras in northern Latin America, al-
Guarea though these specimens are distinctive by their relatively
Myrtaceae large size. Similar spores are produced by members of
Eugenia/Myrcia 0.5 the Blechnaceae, Polypodiaceae, Pteridaceae, and the par-
Pelliceriaceae ent plants grow in a wide range of habitats (Tryon and
Pelliciera 52
Rhizophoraceae
Tryon, 1982).
Rhizophora (.95%) Monolete fern spore type 2 (Fig. 7). This spore type
Unknown 1 4 (52–54 3 36–38 mm) is similar to type 1, and it is dis-
Unknown 2 4 tinguished by the more prominent verrucae. It is also pro-
Unknown 3 duced by several members of the Blechnaceae, Polypo-
Unknown 4 2 diaceae, and Pteridaceae, which grow in a variety of hab-
Unknown 5 0.5
itats, and the spore type is widespread in Cenozoic
deposits of northern Latin America. Loc. Pl-7, slide 1,
ESF T-40,4.
Highlands. The locality is ;90 km inland from the pre- Sphaeropteris/Trichipteris (Cyathaceae; Fig. 8). Ob-
sent Gulf Coast, but at the time the sediments were being late, oval-triangular, apices rounded, outer margin lobate
deposited, the coast was farther west as indicated by the due to pits; trilete, laesurae straight, ;18 mm long, ex-
presence of marine invertebrates and Rhizophora-con- tending ;⅔ distance to spore margin; wall conspicuously
taining lignites in the Simojovel sediments. pitted, pits circular, 1.5–4 mm in diameter, inner margin
The general geology of Mexico has been summarized entire, raised border ;1.5 mm wide, irregularly distrib-
by de Cserna (1989) and by Ferrusquı́a-Villafranca uted; size 45 mm. Loc. Pl-7, slide 1, ESF T-30,1. Previous
(1993), and the regional geology around Simojovel is de- record: Paraje Solo Formation.
20 AMERICAN JOURNAL
Figs. 2–3. Collection locality A(KE)/D-621(HU), east side of Simojovel. 2. General view showing exposure of the early Miocene La Quinta
Formation. 3. Detail of exposure showing principal lignite lens from which sample A-4 was collected.
Sphaeropteris is a genus of ;23 species of tree ferns
in the neotropics. It grows from Veracruz, Mexico, to
Andean South America, Peru, Bolivia, and Amazonian
and southeastern Brazil. It is primarily a plant of the low-
land tropics, but it ranges up to 2000 m in elevation
(Tryon and Tryon, 1982). In Chiapas, S. horrida grows
on the edges of pine-oak-Liquidambar forests, and S.
myosuroides grows in the montane rain forest (Smith,
1981). Trichipteris is a genus of ;55 species of tree ferns
distributed from Veracruz, Mexico, to Brazil and Argen-
tina. It grows in lowland rain forest, wet montane and
cloud forests, and can extend into pine-oak-Liquidambar
forests, savannas, grasslands, swamps, and brackish-wa-
ter habitats. It is mostly found up to elevations of ;2000
m, but it can extend to 3500 m (Tryon and Tryon, 1982).
Ceratopteris (Pteridaceae; Fig. 9). Oblate, oval-trian-
gular, apices rounded; trilete, laesura straight, 30–32 mm,
extending ½ to ⅔ distance to spore margin; striate, striae
in numerous, broad, smooth bands 4–5 mm wide, area
between bands 1–2 mm wide; wall 2–3 mm thick; size
;140 mm (folded). Loc. A-4 (5 HU D-621, Simojovel
landslide area), slide 1, ESF S-22 (also D-621, slide 1,
ESF F-39). Prevous records: Cucaracha, Gatun, Gatun-
cillo, Paraje Solo formations.
OF BOTANY [Vol. 86
Ceratopteris consists of two closely related species in
the New World (two others occur in the Old World trop-
ics). They typically grow as floating ferns in freshwater,
but can range into brackish water habitats, and are most
commonly found at elevations up to ;300 m. The genus
is unreported for Chiapas, but probably occurs there
(Smith, 1981).
Pteris (Pteridaceae; Fig. 10). Oblate, triangular, apices
rounded; trilete, laesura obscure, ;26–28 mm long, bor-
dered by lip ;4–5 mm wide; proximal surface laevigate
(obscure), distal surface coarsely verrucate, verrucae low,
irregular, plate-like, ;6–8 mm in diameter; flange 6–8 mm
wide bordering outer spore margin, flange smooth, outer
margin entire; wall ;2 mm thick; size 46–48 mm. Loc.
Pl-7, slide 1, ESF P-34,3. Previous records: Artibonite,
Cucaracha, Culebra, Gatun, Gatuncillo, Ixtapa, La Boca,
Rio Banano, Uscari formation/group/sequence.
Pteris (;280 species) is a tree fern with ;60 species
in the neotropics typically growing at low-to-mid-eleva-
tions in tropical moist and tropical wet forests.
cf. Antrophyum (Vittariaceae; Fig. 11). Oblate, amb tri-
angular, apices rounded; trilete, laesura straight, 25–27
mm long, extending ;¾ distance to spore margin, inner
margin entire; laevigate; wall ;2 mm thick; size 76–78
January 1999] GRAHAM—OLIGO-MIOCENE
Fig. 4. Stratigraphy and correlations of Tertiary strata in the region of Simojovel (after Frost and Langenheim, 1974, fig. 9). Used with
permisssion of Northern Illinois University Press.
mm. Loc. Pl-7, slide 1, ESF M-39. Previous records: Ar-
tibonite, Cucaracha, Culebra, Gatuncillo, Herrerı́a, La
Boca formations/group.
Antrophyum is a genus of ;35 species, with ;10 in
the neotropics. It is a member of rain forest vegetation,
but it extends up to ;1500 m in elevation into cloud
forests. The spore is larger than those in other assem-
blages from northern Latin America (e.g., Herrerı́a For-
mation of Guatemala), and this may be due to swelling
from storage of the HU specimens in glycerine jelly since
;1965.
Picea (Pinaceae; Fig. 12). Monocolpate, colpus ex-
tending between air sacs on lower side of body; vesicu-
late, air sacs 2, approximately spherical, reticulate and
grading into sculpture pattern of body, 28 3 30 mm, body
scabrate, 42–45 3 40–42 mm; no reentrant angle, no
marginal flange; wall ;2 mm thick; size 61–63 3 40–42
mm. Loc. Pl-7, slide 1, ESF Q-41,1–2. Previous records:
Paraje Solo, Padre Miguel formation/group.
These specimens are slightly corroded and possibly en-
larged through long-time storage in glycerine jelly. In
other respects they are similar to pollen of Picea. The
PALYNOFLORA FROM CHIAPAS, MEXICO 21
closest stands of spruce are presently in the mountains of
northern Mexico (P. mexicana, Coahuila, Nuevo Leon;
P. chihuahuana, Chihuahua, Durango).
cf. Crysophila (Palmae; Fig. 13). Prolate-spheroidal;
monocolpate, colpus straight, 20–22 mm long, extending
entire length of grain, inner margin entire; reticulate, re-
ticulum regular, width of muri (;1 mm) approximately
equal to diameter of lumen, surface of muri smooth; tec-
tate-perforate, wall ;2 mm thick, columellae evident
(4003 magnification); size 25–27 3 22–24 mm. Loc. Pl-7,
slide 1, ESF U-30. Previous records: Cucaracha, Culebra
formations.
Palmae type 1 (Fig. 14). Prolate; monocolpate, colpus
straight, 19–20 mm long, extending nearly entire length
of grain, inner margin entire; scabrate; tectate, wall ;2
mm thick, columellae evident (4003 magnification); size
25–27 3 17–19 mm (widest part just off equator of
grain). Loc. Pl-7, slide 1, ESF 29,2.
This generalized type of palm pollen is widespread in
Cenozoic deposits of northern Latin America. The fossil
pollen is not of the Sabal-Scheelea type, which are the
two prominent genera in the modern palm forest of Chia-
22 AMERICAN JOURNAL OF BOTANY [Vol. 86

TABLE 2. Approximate age, location, and references to Tertiary palynofloras of the Gulf/Caribbean region mentioned in the text.

Flora/age Location References

Pliocene
Paraje Solo SE Veracruz, Mexico Graham, 1976, 1993a
Herrerı́a SE Guatemala Graham, 1998
Rio Banano SE Costa Rica Graham and Dilcher, 1998
Mio-Pliocene
Artibonite Haiti Graham, 1990
Gatun (to Miocene) Central Panama Graham 1991a, b, c
Padre Miguel SE Guatemala Graham, 1998
Middle Miocene
Ixtapa Chiapas, Mexico Martı́nez-Hernández, 1992
Méndez (to early Miocene) Chiapas, Mexico Palacios Chávez and Rzedowski, 1993
Early Miocene
Culebra Central Panama Graham, 1988a
Cucaracha Central Panama Graham, 1988b
La Boca Central Panama Graham, 1989
Uscari SE Costa Rica Graham, 1987
La Quinta (Simojovel) Chiapas, Mexico Graham and Palacios Chávez, 1996; Langeheim,
Hackner, and Bartlett, 1967; this report
Oligocene
Punta Alegre Central Cuba Areces-Mallea, 1987
San Sebastian Puerto Rico Graham and Jarzen, 1969
Eocene
Gatuncillo Panama Graham, 1985
Chapelton Jamaica Graham, 1993b

pas (see Discussion: Modern vegetation, below). It prob-
ably represents a palm of the lowland to premontane wet/
moist forests.
Bombacaceae (cf. Aguiaria type; Fig. 15). Peroblate,
amb oval-triangular to nearly circular; tricolpate, colpi
equatorially arranged, meridionally elongated, equidis-
tant, situated in interapex area, short (4–5 mm, apex to
equator), bordered by costae colpi ;3 mm wide; reticu-
late, reticulum regular, width of muri slightly less than
diameter of lumen (;2 mm), surface of muri smooth;
tectate-perforate, wall ;2 mm thick, columellae evident
(4003 magnification); size 25–27 mm. Loc. Pl-7, slide 1,
ESF U-38,4. Previous records: Gatun, La Boca, Uscari
formations/sequence.
These specimens are of the general Aguiaria type, but
the wall is thinner; the Gatun, La Boca, and Uscari grains
are more similar. Aguiaria is presently a monotypic genus
of Brazil.
Ericaceae (Fig. 16). Tetrahedral tetrads, individual
grains roughly spherical (compressed in tetrads); tricol-
pate, colpi equatorially arranged, meridionally elongated,
equidistant, straight, inner margin entire, bordered by nar-
row margo ;2 mm wide, colpi shared (continuous across
contact between grains); psilate; tectate, wall 2–3 mm
thick, homogeneous (4003 magnification); size—indi-
vidual grains ;15 mm, tetrads 25–27 mm. Loc. Pl-7, slide
1, ESF P-39, 3–4. Previous records: Gatun, Uscari for-
mation/sequence.
Pollen of the Ericaceae cannot be identified to genus.
The family is represented in Central America by ;20
genera, which are most common in upland habitats.
Alfaroa/Oreomunnea (Juglandaceae; Fig. 17). Oblate,
amb oval-triangular; triporate, pores equatorially ar-
ranged, meridionally elongated, equidistant, circular, 3–4
mm in diameter, inner margin entire; psilate to finely sca-
brate; tectate, wall 2 mm thick, homogeneous (4003 mag-
nification); size 24–25 mm. Loc. Pl-7, slide 1, ESF M-
40, 1–3. Previous records: Artibonite, Cucaracha, Gatun,
Gatuncillo, Herrerı́a, La Boca, Paraje Solo, San Sebastian
formations/group.
Pollen of the complex that includes Alfaroa, Engel-
hardia, and Oreomunnea cannot be distinguished on the
basis of pollen characters. In the treatment of Stone
(1972; Stone and Broome, 1975), Alfaroa and Oreomun-
nea are New World genera distributed from southern
Mexico to northern South America, while Engelhardia is
an Old World genus. The neotropical genera grow pri-
marily at the transition between wet mountain forests and
premontane cloud forests, and often in association with
Podocarpus.
Crudia (Leguminosae, Caesalpinioideae; Fig. 18). Pro-
late; tricolporoidate, colpi equatorially arranged, meridi-
onally elongated, equidistant, straight, 40–42 mm long,
extending nearly entire length of grain, pore area faint,
circular, situated at midpoint of colpus; distinctly and
coarsely striate, striae oriented parallel to long axis of
grain, surface of striae smooth; tectate, wall ;2 mm thick,
homogeneous (4003 magnification); size 48–50 3 32–
34 mm. Loc D-621, slide 1, ESF K-30. Previous records:
Cucaracha, Gatun, Gatuncillo, La Boca formations.
Crudia is primarily an Amazonian genus of approxi-
mately ten species growing in riverine habitats. In addi-
tion to the microfossil record, pods of Crudia have been
reported from the Eocene of western Tennessee (Heren-
deen and Dilcher, 1990).
Cedrela (Meliaceae; Figs. 19–21). Prolate-spheroidal,
amb circular; tetracolporate, colpi equatorially arranged,
meridionally elongated, equidistant, straight, 25–27 mm
January 1999] GRAHAM—OLIGO-MIOCENE PALYNOFLORA FROM CHIAPAS, MEXICO 23

Figs. 5–16. Fossil pollen and spores from the early Miocene La Quinta Formation, Simojovel, Chiapas, Mexico. 5. Selaginella. 6. Monolete
fern spore type 1. 7. Monolete fern spore type 2. 8. Sphaeropteris/Trichipteris. 9. Ceratopteris. 10. Pteris. 11. cf. Antrophyum. 12. Picea. 13. cf.
Crysophila. 14. Palmae type 1. 15. Bombacaceae (cf. Aguiaria type). 16. Ericaceae.
24 AMERICAN JOURNAL
Fig. 17–31. Fossil pollen and spores from the early Miocene La Quinta Formation, Simojovel, Chiapas, Mexico. 17. Alfaroa/Oreomunnea. 18.
Crudia. 19–21. Cedrela. 22. Guarea. 23. Eugenia/Myrcia. 24. Pelliciera. 25–26. Rhizophora. 27. Unknown. 28. Unknown 2. 29. Unknown 3. 30.
Unknown 4. 31. Unknown 5.
long, pores situated at midpoint of colpus, slightly elon-
gated equatorially, 4–6 3 3–5 mm, costae pori ;4 mm
wide; psilate; tectate, wall ;3 mm thick, homogeneous
(4003 magnification); size 46–48 3 37–39 mm. Loc. Pl-7,
slide 1, ESF X-31, S-45, S-38,3. Previous records: Gatun,
Paraje Solo formations.
Cedrela includes approximately six species of trees in
Mexico and Central America. The two most common
species are C. odorata, distributed from northern Mexico
to South America, and C. angustifolia found along the
Pacific slopes. Both grow at low-to-moderate elevations,
and mostly in tropical moist and premontane wet forests,
although C. angustifolia can extend into the drier habitats
of the tropical deciduous forest.
Guarea (Meliaceae; Fig. 22). Oblate-spheroidal, amb
circular; tetracolporate, colpi equatorially arranged, me-
ridionally elongated, equidistant, straight, relatively short
(8–9 mm), inner margin entire, pores situated at midpoint
OF BOTANY [Vol. 86
of colpus, elongated equatorially, 2 3 5 mm, inner margin
entire; psilate; tectate, wall ;3 mm thick, homogeneous
(4003 magnification); size 22–24 mm. Loc. Pl-7, slide 1,
ESF T-32,2. Previous records: Paraje Solo, San Sebastian,
Gatun formations.
Guarea is a genus of ;150 species, with G. glabra
and G. multiflora being two of the common Central
American species. Both are distributed from Mexico to
South America and grow as understory trees at low-to-
moderate elevations in tropical wet and tropical moist
forests.
Eugenia/Myrcia (Myrtaceae; Fig. 23). Oblate, amb tri-
angular; tricolporate to tricolporoidate, syncolpate, colpi
equatorially arranged, meridionally elongated, equidis-
tant, straight, 7–9 mm long, inner margin entire, pore ob-
scure, situated at mid-point of colpus, ;1 mm in diame-
ter; psilate to scabrate; tectate, wall 2 mm thick, homo-
geneous (4003 magnification); size 19–21 mm. Loc. Pl-7,
January 1999] GRAHAM—OLIGO-MIOCENE PALYNOFLORA FROM
slide 1, ESF Q-30,2–4. Previous records: Cucaracha, Cu-
lebra, Gatun, Gatuncillo, Paraje Solo, San Sebastian, Us-
cari formations/sequence.
The pollen of these genera cannot be distinguished at
the level of light microscopy (Graham, 1980), and the
family is widespread in a wide variety of habitats.
Pelliciera (Pelliceriaceae; Fig. 24). Oblate, amb cir-
cular; tricolporate, colpi equatorially arranged, meridio-
nally elongated, equidistant, 22–26 mm long, inner mar-
gin entire, pore circular, 3–4 mm in diameter, situated at
midpoint of colpus, inner margin entire; sculpture vari-
able from finely to coarsely verrucate; tectate, wall 4 mm
thick, columellae evident (4003 magnification); size var-
iable (45–65 mm). Loc. A-4 (5 HU D-621, Simojovel
landslide area), slide 1, ESF A-49. Previous records: Ga-
tuncillo, La Boca, Rio Banano, San Sebastian, Chapelton
formations.
Pelliciera (5 Pelliceria) is a small mangrove tree pres-
ently distributed from Costa Rica to northwest Colombia
and Ecuador. Its more widespread distribution in the Ter-
tiary of the Caribbean region has been summarized by
Graham (1977, 1995).
Rhizophora (Rhizophoraceae; Figs. 25–26). Prolate to
prolate-spheroidal, amb circular; tricolporate, colpi equa-
torially arranged, meridionally elongated, equidistant,
narrow, straight, 13–16 mm long, inner margin entire,
costae colpi ;2 mm wide, pores elongated equatorially
(colpi transversalis), ;1 3 4 mm, situated at midpoint of
colpus, inner margin entire; finely reticulate; tectate-per-
forate, wall 2–3 mm thick, columellae evident (4003
magnification); size 18–22 3 23–25 mm. Loc. Pl-7, slide
1, ESF G-39,2, U-37,1–3. Previous records: Cucaracha,
Culebra, Gatun, Gatuncillo, Herrerı́a, La Boca, Paraje
Solo, Rio Banano, San Sebastian, Uscari formations/se-
quence.
Rhizophora is a small tree or shrub growing in brack-
ish water along the coasts of warm-temperate to tropical
regions. Its geologic history in the Caribbean region,
along with that of other mangroves, has been summarized
by Graham (1977, 1995).
Unknown 1 (Fig. 27). Prolate, amb circular; tricolpor-
ate, colpi equatorially arranged, meridionally elongated,
equidistant, 32–34 mm long, inner margin entire, bor-
dered by costae colpi 3–4 mm wide, pore circular, ;4 mm
in diameter, situated at midpoint of colpus, inner margin
entire; reticulate, reticulum regular, becoming finer to-
ward colpus to form narrow margo, width of muri ap-
proximately equal to diameter of lumen, surface of muri
smooth; tectate-perforate, wall 3 mm thick, columellae ev-
ident (4003 magnification); size 40 3 30 mm. Loc. Pl-7,
slide 1, ESF O-31.
Unknown 2 (Fig. 28). Prolate, amb circular; tricolpor-
ate, colpi equatorially arranged, meridionally elongated,
equidistant, 20 mm long, inner margin entire, bordered by
costae colpi 3–4 mm wide, pore circular, ;4 mm in di-
ameter, situated at midpoint of colpus, inner margin en-
tire; scabrate; tectate, wall 3 mm thick, homogeneous to
columellae just evident (4003 magnification); size 28 3
18 mm. Loc. Pl-7, slide 1, ESF Q-35, 1–3.
Unknown 3 (Fig. 29). Prolate to prolate-spheroidal,
amb circular, tricolporate, colpi equatorially arranged,
meridionally elongated, equidistant, 12 mm long, inner
margin entire to minutely dentate, bordered by costae col-
CHIAPAS, MEXICO 25
pi ;3 mm wide, pores slit-like, ;0.5 3 3 mm, situated
at midpoint of colpus; finely reticulate, reticulum regular,
width of muri approximately equal to diameter of lumen
(;1 mm), surface of muri smooth; tectate-perforate, wall
;3 mm thick, columellae evident (4003 magnification);
size 21 3 16 mm. Loc. Pl-7, slide 1, ESF M-32.
Unknown 4 (Fig. 30). Oblate to oblate-spheroidal, amb
circular; tricolpate (pores, if present, obscure), colpi
equatorially arranged, meridionally elongated, equidis-
tant, 10–12 mm long, inner margin entire to minutely
dentate; finely reticulate, reticulum regular, width of muri
approximately equal to diameter of lumen (;1 mm), sur-
face of muri smooth; tectate-perforate, wall ;2 mm thick,
columellae evident (4003 magnification), size 30 mm.
Loc. Pl-7, slide 1, ESF N-39,1.
Unknown 5 (Fig. 31). Oblate, amb circular; tricolpor-
ate, colpi equatorially arranged, meridionally elongated,
equidistant, 6–8 mm long, inner margin entire; pores cir-
cular (obscure), ;4–5 mm in diameter, situated at mid-
point of colpus; finely reticulate, reticulum regular, width
of muri approximately equal to diameter of lumen (;1
mm), becoming finer toward poles and colpi, surface of
muri smooth; tectate-perforate, wall ;2 mm thick, colu-
mellae evident (4003 magnification); size 27 mm. Loc.
Pl-7, slide 1, ESF U-39,2–4.
In addition to the 19 identified and five unknown pa-
lynomorphs described here from the Simojovel assem-
blage, Palacios Chávez (in Graham and Palacios Chávez,
1996) provided photographs of Ilex and Asteraceae pol-
len from samples processed at the Instituto Politecnico
Nacional, Mexico City. Also, Langenheim, Hackner, and
Bartlett (1967, p. 319) found ‘‘a few poorly preserved
grains probably of Pinus,’’ Podocarpus, and Pachira,
which were not seen during scans from two (D-621, Pl-7)
of their 13 localities in the HU collections. The glycerine
jelly in many of the HU slides has partly or completely
deteriorated and some specimens are no longer recogniz-
able. Ilex is widespread in northern Latin America and
grows in a variety of habitats, Pinus and Podocarpus are
members of the upland evergreen cloud forests, and Pa-
chira is a lowland tropical, riverine plant occupying hab-
itats similar to those of Crudia in South America.
DISCUSSION
Modern vegetation—The extant plant communities of
Mexico are described by Rzedowski (1978) and those of
Chiapas by Breedlove (1973, 1981). The latter uses the
classification scheme of Beard (1944) and, where possi-
ble, incorporates or correlates Beard’s units with those of
other authors (Miranda, 1952/1953; Miranda and Her-
nández X., 1963; Wagner, 1964; Gómez Pompa, 1965;
Holdridge, 1967; Pennington and Sarukhán K., 1968;
Flores et al., 1971; Holdridge et al., 1971; Rzedowski,
1978). Systems using climatic data for classifying vege-
tation (e.g., Holdridge, 1967; Holdridge et al., 1971) are
difficult to apply uniformly to Chiapas because these data
are lacking for many areas and especially for sites above
2700 m.
Breedlove (1981) classifies the vegetation of Chiapas
into 19 formations [18 in Breedlove, (1973)]:
Tropical rain forest (5 selva alta siempre, in part; selva
alta perennifolia, in part; bosque tropical perennifolio, in
26 AMERICAN JOURNAL
part). This forest is mostly restricted to valleys bordering
the Rio Usumacinta on the low eastern slopes of Front
Ranges and High Plateaus and on Gulf Coastal Plain. It
is surrounded by lower montane rain forest, and includes
as the prominent genera Aspidosperma, Brosimum, Di-
alium, Erblichia, Ficus, Guatteria, Manilkara, Poulsenia,
Protium, Swietenia, Talauma, Terminalia, and Vatairea,
with an understory of Alchornea, Alibertia, Belotia, Ble-
pharidium, Blumea, Bursera, Cassia, Cephaelis, Dracae-
na, Forchhammeria, Guarea, Hasseltia, Hirtella, Lacis-
temma, Licaria, Orthion, Ouratea, Piper, Pithecellobium,
Quararibea, Sickingia, Wimmeria, and Zuelania.
Lower montane rain forest (selva alta siempre verde,
in part; selva alta perennifolia, in part; bosque tropical
perennifolio, in part; tropical premontane moist, wet, or
rain forest). Remnants of this forest suggest it potentially
would cover much of the Front Ranges and High Pla-
teaus, and it also occurs as a narrow band along the west-
ern lowlands of the Sierra Madre de Chiapas. It is similar
to the tropical rain forest, but it lacks the uppermost can-
opy layer of tallest trees, and lianas and epiphytes are
more common. Most of the tropical rain forest genera are
present, in addition to Alseis, Belotia, Calophyllum,
Chaetoptelea (Ulmus), Licania, Nectandra, Ocotea,
Quercus, Sebastiana, Talauma, and Vochysia in the can-
opy, and Chrysophyllum, Cleidion, Cymbopetalum, Far-
amea, Piper, Rinorea, Sloanea, Stemmadenia, and Tro-
phis in the understory.
Montane rain forest (5 selva mediana y baja siempre
verde, in part; selva mediana o baja perennifolia, in part;
bosque tropical perennifolio, in part; tropical montane
moist, wet, rain forest). This forest is found at elevations
of 900–2200 m in the Front Ranges and High Plateaus,
and on the eastern and western slopes of the Sierra Madre
de Chiapas. It is characterized by numerous epiphytic
ferns, aroids, bromeliads, and orchids. The prominent
genera include Alfaroa, Ardista, Brunellia, Calatola,
Hedyosmum, Hyperbaena, Lunania, Matudea, Melisoma,
Mirandaceltis, Oecopetalum, Oreopanax, Quercus, Pla-
tanus, Synardisia, Trichilia, and Turpinia. Understory
trees and shrubs are Acalypha, Billia, Centropogon,
Cephaelis, Chamaedorea, Eugenia, Hampea, Miconia,
Mollinedia, Oreopanax, Palicourea, Parathesis, Psycho-
tria, Siparuna, Trophis, and Urera.
Evergreen cloud forest (5 cloud forest, in part; selva
mediana y baja siempre verde, in part; selva mediana o
baja perennifolia, in part; subtropical montane wet or rain
forest). This forest is found near the crests of mountains
at elevations of 1900–3200 m in the Front Ranges and
High Plateaus and in the Sierra Madre de Chiapas. Hang-
ing mosses and epiphytic ferns are common, fog and
clouds are usually present, and minimum January tem-
peratures may approach 08C, although frost is rare. The
common trees are Abies, Pinus, Podocarpus, Acer, Chir-
anthodendron, Clethra, Drimys, Magnolia, Meliosma,
Microtropis, Olmediella, Oreopanax, Persea, Photinia,
Quercus, Weinmannia, Wimmeria, and Zinowiewia. Un-
derstory plants include Cavendishia, Cleyera, Deppea,
Eugenia, Fuchsia, Gentlea, Greigia, Mahonia, Miconia,
Rapanea, Saurauia, Symplocos, Vaccinium, and Vibur-
num.
Evergreen seasonal forest (5 selva alta subdecidua;
selva alta or mediana subperennifolia; selva alta or me-
OF BOTANY [Vol. 86
diana subcauducifolia; bosque tropical perennifolio, in
part; tropical or subtropical premontane, moist forest).
There are several transitional communities between the
tropical rain forest and other vegetation types as rainfall
decreases and becomes more seasonal. In the evergreen
seasonal forest there is a marked dry season, and this
vegetation covers much of the slopes of the Sierra Madre
de Chiapas up to an elevation of ;1200 m, the southern
and western part of the northern Front Ranges and High
Plateaus, and the Gulf Coast Plain. Important trees in-
clude Astronium, Bernoullia, Brosimum, Brumelia, Ca-
lycophyllum, Ceiba, Coccoloba, Cordia, Cupania, Elaeo-
dendron, Enterolobium, Ficus, Guettarda, Hymenaea,
Lafoensia, Licania, Platymiscium, Rheedia, Sapium, Ster-
culia, Styrax, Tabebuia, and Vatairea. Associated shrubs
and small trees are Amyris, Ardisia, Bourreria, Calyp-
tranthes, Clusia, Eugenia, Gentlea, Karwinskia, and Psy-
chotria.
Tropical deciduous forest (5 deciduous seasonal forest,
selva baja decidua, selva baja cauducifolia, bosque trop-
ical deciduo, tropical or subtropical dry and very dry
forest). In regions covered by this forest there is a pro-
nounced dry season of 4–6 mo. It occurs along the Gulf
Coastal Plain, on the lower hills of the Sierra Madre de
Chiapas, and in the Central Depression. The common
trees are Annona, Bucida, Bursera, Calycophyllum, Cec-
ropia, Cedrela, Ceiba, Cochlospermum, Cordia, Eysen-
hardtia, Gliricidia, Godmania, Hauya, Heliocarpus,
Hura, Ipomoea, Leucaena, Luehea, Lysiloma, Plumeria,
Pseudobombax, Spondias, Stemmadenia, Swietenia, Ta-
bebuia, and Triplaris.
Short-tree savanna (5 sabana, selva baja subperenni-
folia, vegetacion sabanoide, savanna). This community
grows on gradual slopes or flat bottomlands on shallow,
poorly drained soils where it is usually associated with
various other vegetation types. The common trees are
Byrsonima, Crescentia, and Curatella, with Acacia, Al-
varadoa, Cordia, Piscidia, Quercus, and Tetracera.
Thorn woodland (5 selva baja espinosa cauducifolia,
bosque espinoso, tropical and subtropical thorn wood-
land). This community is prominent in the Isthmus of
Tehuantepec and it extends into Chiapas only in the
northern part of the Central Depression and along the
Pacific Coastal Plain. Important members include Acacia,
Bauhinia, Buettneria, Bursera, Casearia, Cordia, Croton,
Diphysa, Erythroxylon, Jacquinia, Karwinskia, Piptad-
enia, Pisonia, Pithecellobium, Randia, Sageretia, Steno-
cereus, and Zizyphus.
Pine-oak-Liquidambar forest (5 cloud forest, in part;
deciduous forest; bosque cauducifolio; bosque deciduo;
bosque mesofilo de montãna; subtropical montane moist
forest). This forest occurs on moist north and east-facing
upland slopes in the Front Ranges and High Plateaus and
in the Sierra Madre de Chiapas. Many of the genera also
grow in the deciduous forest of eastern United States and
include Pinus, Carpinus, Cassia, Citharexylum, Clethra,
Cornus, Cupania, Erythrina, Fraxinus, Liquidambar,
Meliosma, Montanoa, Nyssa, Ostrya, Perrottetia, Quer-
cus, Rhus, Saurauia, Styrax, and Turpinia. Associates in-
clude Aphelandra, Baccharis, Bocconia, Cestrum, Fuch-
sia, Gaultheria, Guamatela, Hibiscus, Liabum, Parathes-
is, Phenax, Rapanea, Solanum, Symplocos, Triumfetta,
Verbesina, and Viburnum.
January 1999] GRAHAM—OLIGO-MIOCENE PALYNOFLORA FROM CHIAPAS, MEXICO 27

TABLE 3. Representation of extant community elements of Chiapas in TABLE 3. Continued.

the early Miocene La Quinta assemblage. Vegetation classification
follows Breedlove (1973, 1981), and placement of ferns follows Bunch grassland
Tryon and Tryon (1982) and Smith (1981). Fossil pollen of the Lycopodium
Cyperaceae, Gramineae, Ericaceae, and many Palmae and Com- Possibly some Gramineae
positae cannot be identified to genus; pollen of the Lauraceae usu- Possibly Vaccinium (as Ericaceae)
ally does not preserve in the fossil record; and some taxa with Evergreen cloud scrub
species occurring in different communities (e.g., Pinus, Quercus) Lycopodium (widespread)
cannot be distinguished on the basis of pollen characters. Taxa in Gramineae
parentheses are names reported in the fossil record. Possibly Vaccinium (as Ericaceae)
Herbaceous marsh
Tropical rain forest Possibly some Gramineae
Canopy/dominants—none Coastal strand vegetation
Understory/associates Possibly some Gramineae
Selaginella (widespread) Second-growth and successional forest
cf. Antrophyum (to cloud forests) Acacia
Pteris (ecologically diverse, cloud forests, deciduous forest, pine Compositae (widespread)
woods, secondary forests
Sphaeropteris (to pine-oak-Liquidambar, montane forests)/Trichip-
teris (ecologically diverse, to pine-oak-Liquidambar, wet mon-
tane-cloud forests, savanna, grasslands, swamp forests) Pine-oak forest (5 pinares and encinares, bosque de
Guarea pino y encino). This forest is best developed on dry south
cf. Aguiaria type (not present in the modern Chiapas flora, South and west-facing slopes of the central Front Ranges and
American; identification only as example of morphology) High Plateaus and on the east side of the Sierra Madre
Lower montane rain forest de Chiapas, mostly at elevations between 1300 and 2500
Alfaroa (to transition with montane rain forest)
Montane rain forest
m. It includes Pinus, Arbutus, Buddleia, Crataegus, and
Canopy/dominants Quercus. Common shrubs are Ceanothus, Chiococca,
Alfaroa (as Alfaroa/Oreomunnea; lower limits) Garrya, Holodiscus, Lippia, Litsea, Mahonia, Monnina,
Eugenia (as Eugenia/Myrcia) Myrica, Rhus, Senecio, Solanum, and Viburnum.
Understory/associates—none Swamp and lowland riparian forest (5 manglar, in part;
Evergreen cloud forest canacoital). This forest is well developed along lowland
Canopy/dominants
Picea (not present in the modern Chiapas flora)
rivers and in swamps directly behind the mangroves. The
Pinus? prominent trees are Taxodium, Andira, Bravaisia, Bucida,
Podocarpus Calophyllum, Haematoxylum, Pachira, and Salix.
Understory/associates Mangrove swamp (5 manglar). Mangrove vegetation
Possibly Cavendishia/Vaccinium (as Ericaceae) extends along the entire Pacific Coast of Chiapas. The
Eugenia (as Eugenia/Myrcia) dominants are Avicennia, Conocarpus, Laguncularia, and
Evergreen seasonal forest
Canopy/dominants
Rhizophora. Batis and Philoxerus are common associated
Hymenaea herbs.
Ilex (widespread) Palm forest (5 palmar). Vegetation dominated by
Understory/associates palms is found along rivers in the Central Depression,
Eugenia (as Eugenia/Myrcia) along the Rio Usumacinta and tributaries in the eastern
Tropical deciduous forest portion of the Front Ranges and High Plateaus, and in
Canopy/dominants
Cedrela
the southern portion of the Pacific Coastal Plain. The
Short-tree savanna common palms are Sabal and Scheelea. Palm forests ben-
Acacia efit and may result from human actitivies
Thorn woodland Temperate riparian forest. This forest occurs along
Acacia streams at elevations above 1500 m in the eastern Front
Pine-oak-Liquidambar forest Ranges and High Plateaus. Members include Acer, Alnus,
Pinus?
Tapirira (moist habitats)
Baccharis, Berchemia, Cornus, Crataegus, Cuphea, Pla-
Pine-oak forest tanus, and Salix.
Canopy/dominants Bunch grassland (5 zacatonal). Communities charac-
Pinus? terized by tall bunch grasses are found in the eastern
Possibly Arbutus (as Ericaceae) Front Ranges and High Plateaus, and include Briza, Bro-
Understory/associates—none mus, Festuca, Muhlenbergia, Stipa, and Trisetum.
Swamp and lowland riparian forest
Ceratopteris
Evergreen cloud scrub (5 paramo; paramos de altura;
Crudia tropical or subtropical montane or subalpine moist, wet,
Pachira or rain paramo). This vegetation occurs on the highest
Mangrove swamp peaks of the Front Ranges and High Plateaus and the
Rhizophora (dominant in the fossil assemblage) Sierra Madre de Chiapas. The common members are Ly-
Pelliciera (common in the fossil assemblage, not present in the copodium, Gleichenia, Alchemilla, Arenaria, Beshorner-
modern Chiapas flora)
Understory/associates—none
ia, Calamagrostis, Comarostaphylis, Draba, Festuca,
Palm forest Gaultheria, Gnaphalium, Haplopappus, Holodiscus, Lu-
as Palmae, cf. Crysophila pinus, Luzula, Muhlenbeckia, Myrrhidendron, Pernettya,
Temperate riparian forest—none Potentilla, Ternstroemia, Trisetum, Vaccinium, Weldenia,
and Werneria.
28 AMERICAN JOURNAL OF BOTANY [Vol. 86

TABLE 4. Distribution of dominant and associated taxa in the modern TABLE 4. Continued.
vegetation of Chiapas (see Discussion: Modern vegetation) in other
Tertiary palynofloras of the Gulf/Caribbean region. 1 5 Chapelton Vegetation/taxa Eocene Oligocene Miocene Pliocene
Formation, 2 5 Gatuncillo, 3 5 San Sebastian, 4 5 Punta Alegre,
Understory/associates
5 5 Cucaracha, 6 5 Culebra, 7 5 La Boca, 8 5 Uscari Sequence,
Eugenia—as above
9 5 Ixtapa, 10 5 Artibonite Group, 11 5 Gatun, 12 5 Padre Mi-
guel Group, 13 5 Paraje Solo, 14 5 Herrerı́a 15 5 Rio Banano. Tropical deciduous forest
cf. Bucida 11
Vegetation/taxa Eocene Oligocene Miocene Pliocene Bursera—as above
Tropical rain forest Cedrela 11 13
Ceiba—as above
Canopy/dominants Hauya 3 11
cf. Ficus 2 Pseudobombax 7, 11
Protium 13
cf. Protium 2 Short-tree savanna
Terminalia 2 6 13 Canopy/dominants—none
(Combretum/Terminalia) Understory/associates
Understory/associates Acacia 3 6, 11
Alchornea 3 5, 6, 7, 8, 13 cf. Acacia 13
10, 11, 15 Quercus—as above
cf. Alibertia 13
Thorn woodland
cf. Bumelia 11
Bursera 3 11 13 Canopy/dominants
Guarea 3 11 13 Acacia—as above
Buettneria 13
Lower montane rain forest Bursera—as above
Canopy/dominants Casearia 2 3 6 13
Ulmus/Chaetopetala (Ul- 13
Pine-oak-Liquidambar
mus)
Quercus 11, 12 13 Canopy/dominants
Pinus—as above
Understory/associates
Cupania—as above
Chrysophyllum 3
Erythrina 11
cf. Chrysophyllum 2
Liquidambar 3 13
Cymbopetalum 11
Meliosma—as above
Faramea 2 3 11 13
Nyssa 3
Montane rain forest Quercus—as above
Canopy/dominants Understory/associates
Alfaroa (Alfaroa/Oreo- 2 3 5, 7, 9, 13, 14 Hampea/Hibiscus—as
munnea, Engelhardia) 11 above
Brunellia 3 Symplocos—as above
Hedyosmum 13
Pine-oak forest
Meliosma 13
Quercus—as above Canopy/dominants
Pinus—as above
Understory/associates
Quercus—as above
cf. Chamaedorea 13
Eugenia/Myrcia 2 3 5, 6, 8, 11 13 Understory/associates
Hampea/Hibiscus 6, 11 13 Ericaceae (possibly Arbu-
tus)—as above
Evergreen cloud forest Myrica 13
Canopy/dominants
Swamp and lowland riparian forest
Abies 13
Pinus 4 9, 10, 12 13, 14 Canopy/dominants
Podocarpus 3 8, 11 13 Bravaisia 13
Meliosma—as above cf. Bucida 11
Quercus—as above Salix 3
Understory/associates Mangrove swamp
Cleyera 13 Laguncularia
Ericaceae (possibly Caven- 8, 11 Rhizophora 2 3 5, 6, 7, 8 13
dishia) 9, 11 13, 14
Eugenia—as above
Symplocos 11 Palm forest
cf. Astrocaryum 13
Evergreen seasonal forest cf. Attalea 13
Canopy/dominants cf. Brahea 13
Bernoulia 3 11 cf. Chamaedorea 13
Ceiba 11 Chrysophila type 5, 6
cf. Ceiba 7 Desmoncus type 6
Coccoloba 2 13 Manicaria type 5, 6, 7
Cupania 6, 11 13 cf. Maximiliana 13
Ficus—as above Synechanthus type 6, 7
Sapium 6 Palmae 2 10, 11, 12 14, 15
cf. Sapium 13
January 1999] GRAHAM—OLIGO-MIOCENE
TABLE 4. Continued.
Vegetation/taxa Eocene Oligocene Miocene
Temperate riparian forest
Alnus
Cuphea
Salix—as above
Bunch grassland
Gramineae 6, 7, 11, 12
Evergreen cloud scrub
Lycopodium 3 6, 7, 8, 9
11, 12
Gramineae—as above
Ericaceae (possibly Vaccin-
ium)—as above
Herbaceous marsh
Cyperaceae 12
Ludwigia 12
Gramineae—as above
Coastal strand vegetation
cf. Alibertia
Coccoloba 2 deciduous forest (Cedrela).
Gramineae—as above
Second-growth
Acacia—as above
Herbaceous marsh (5 tulare, popal). Marshes occur
scattered through Chiapas in areas of low elevation. The
composition is variable, but frequently includes Calathea,
Caperonia, Carex, Caladium, Cyperus, Heliconia, Hy-
menachne, Juncus, Leersia, Ludwigia, Lythrum, Neptu-
nia, Nymphaea, Paspalum, Phragmites, Rhynchospora,
Scirpus, Sesbania, Thalia, and Typha.
Coastal strand vegetation (5 cordon littoral). This
community is found on sandy beaches of the Pacific
Coast and is characterized by scandent shrubs and herbs.
The common members are Acanthocereus, Alibertia,
Canavalia, Capparis, Coccoloba, Distichlis, Ipomoea,
Monanthocloë, Okenia, Pectis, Prosopis, Salpianthus, Se-
suvium, Stegnosperma, Uniola, and Ximenia.
Second-growth and successional forest and shrub as-
sociations. This is a disturbance and recovery vegetation
of variable composition, but frequently includes Abutilon,
Acacia, Arctostaphylos, Baccharis, Calliandra, Ceano-
thus, Clibadium, Coaxana, Lantana, Lippia, Malvaviscus,
Muhlenbeckia, Pavonia, Smallanthus, Rhamnus, Rubus,
Salvia, Tithonia, Vernonia, and Viburnum.
Paleocommunities and physical setting—The distri-
bution of taxa identified from the La Quinta flora is ar-
ranged according to the extant community types in Chia-
pas in Table 3. The occurrence of fossil representatives
of the modern vegetation elements in other Tertiary floras
of the Gulf/Caribbean region is given in Table 4. One of
the principal paleocommunities present during the early
Miocene in eastern Chiapas was mangrove. This com-
munity now grows on the Pacific Coast Plain, and on the
Gulf Coast Plain of Veracruz and Tabasco beyond the
political boundaries of Chiapas. In the Tertiary, however,
it was part of the shallow, brackish-water vegetation in
the vicinity of Simojovel. It was different from the mod-
ern mangrove vegetation of Mexico and northern Central
PALYNOFLORA FROM CHIAPAS, MEXICO 29
America because in addition to Rhizophora it included
Pelliciera, which is now distributed from Costa Rica to
Pliocene
northwestern South America.
Inland from the mangroves there was a swamp and
13 lowland riparian forest represented by fossil spores and
13 pollen of Ceratopteris, Crudia, and Pachira. Also in the
lowlands was a version of the tropical rain forest. None
of the dominants of the modern rain forest of Chiapas
13, 14 are represented in the La Quinta assemblage, but several
others are typical of that community. In addition to the
13 riparian Crudia and Pachira, there were the tree ferns
Pteris and Sphaeropteris/Trichipteris, Guarea, possibly
Antrophyum and Aguiaria, and the widespread Selagi-
nella.
At higher elevations there was a montane rain forest
as suggested by Alfaroa and possibly Eugenia, which are
13, 14 mostly lower montane/montane transition plants, and an
evergreen cloud forest of Picea, Pinus, Podocarpus, and
possibly Cavendishia/Vaccinium. In somewhat drier hab-
itats there was a version of the evergreen seasonal forest
13 (Hymenaea, Ilex, and possibly Eugenia), and the tropical
Absent or poorly represented in the La Quinta flora are
defining members of distinctly dry to arid commuities
and high-elevation vegetation. The absence of pollen
from the latter community may be a result of distance
from the lowland basin of deposition, but no dominant
plants of high-montane or paramo habitats were being
blown or washed into the coastal sediments. Pollen of
one of the highest-elevation trees in Mexico (Abies) does
occur in other Neogene floras (e.g., Paraje Solo). There
is also meager representation of pollen of northern tem-
perate elements (Picea, possibly Pinus) in the La Quinta
flora. This is consistent with the pattern based on other
Tertiary assemblages in northern Latin America (Graham,
1973, 1997). These elements are mostly absent to poorly
represented in early Miocene and older floras, prior to the
middle Miocene temperature decline (Miller, Fairbanks,
and Mountain, 1987), and appear in greater numbers in
late Miocene and younger floras. There is also a trend
from the greatest number of northern temperate taxa in
northern Latin American Tertiary palynofloras (e.g., 12
in the Paraje Solo flora of Veracruz, Mexico), to fewer
numbers to the south (e.g., only Quercus in the Gatun
flora of central Panama). This places the principal time
of introduction for these elements in the late Miocene and
later, with the early and middle Miocene being the tran-
sitional time during which they first begin to appear.
Thus, the paleovegetation in the vicinity of Simojovel
in the early Miocene included versions of mangrove
swamp, swamp and lowland riparian forest, tropical rain
forest, lower montane/montane rain forest transition veg-
etation, evergreen cloud forest, evergreen seasonal forest,
and tropical deciduous forest.
Average elevation in the Sierra de Chiapas is presently
;1000 m, and the highest peak is in the south near the
Guatemalan border (;4000 m). In the Front Ranges and
High Plateaus province, which includes the Simojovel lo-
cality, the average elevation is presently ;2000 m and
the highest peak is San Cristobal de Las Casas (3004 m).
The geologic history of the region provides some con-
straints on estimates of the paleoelevation. The highlands
arose by compressive deformation from the northeast to
30 AMERICAN JOURNAL
the southwest primarily in the earliest Miocene. Because
Plio-Pleistocene strata show little deformation, this brack-
ets the principal time of folding and uplift between the
early Miocene and the Plio-Pleistocene. The La Quinta
flora was being deposited near the beginning of highland
formation, while the younger Pichucalco and Ixtapa floras
were deposited after somewhat greater elevations had de-
veloped. The paleoelevation in the vicinity of Simojovel
in the early Miocene is estimated at 1000–1200 m, which
would allow for the presence of a midelevation forest of
Picea, Pinus, Podocarpus, and Alfaroa, but not a high-
elevation forest of Abies or paramo.
Paleoclimate—The present annual rainfall in the vi-
cinity of Simojovel is ;2500 mm and summer wet. With
the more inland position of the sea, and slightly greater
and more evenly distributed precipitation in the late Mio-
cene, the region would likely support a rain forest in the
lowlands without converting the midelevation evergreen
cloud forest into a more tropical selva, or the drier ev-
ergreen seasonal/tropical deciduous forests into more me-
sic formations. The present MAT at midelevations in the
region is a relatively uniform 248C, and it was at least at
that value during the early Miocene. At 248C, tempera-
ture is not a limiting factor for the growth of most low-
to midelevation tropical forests if there is no great sea-
sonal variation. Thus, it is difficult to detect warmer tem-
peratures on the basis of plant microfossils from lowland
basins of deposition. The absence of upper montane cool-
temperate, and possibly paramo vegetation, is more likely
a reflection of the absence of highlands in the immediate
vicinity of Simojovel than of substantially higher MAT.
By this scenario, Tertiary floras in the region younger
than the Simojovel should have better representation of
cool-temperate and higher elevation communities, and
the Ixtapa (Martı́nez-Hernández, 1992) and Méndez flo-
ras (Palacios Chávez and Rzedowski, 1993) reflect this
trend. Further modernization of the flora occurred with
additional immigrations from the north concomitant with
continued late Cenozoic cooling (Miller, Fairbanks, and
Mountain, 1987), and tropical elements from the south
with closure of the isthmian land bridge between ;3.5
and 2.5 Ma (Coates et al., 1992; Graham, 1992; Jackson,
Budd, and Coates, 1996; Webb and Rancy, 1996).
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