Axelrod 1970

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MESOZOICPALEOGEOGRAPHYAND EARLY
ANGIOSPERMHISTORY
DANIEL I. AXELROD
Departments of Geology and Botany
University of California
Davis, California95616
Abstract -----277
Introduction -----277
Some Early Angiosperms -----279
Paleogeography -- ---279
Pre-Cretaceous History - - - ---280
Environment --- --280
Early Evolution -----282
CretaceousHistory -----284
Invasion of Lowlands -----284
Evolutionary Significance of Megafossil-MicrofossilRecords --- 287
The Cradle of Flowering Plants -----290
Cretaceous Plant Geography - - ---- 292
Tropical Links - - - ---292
Desert Links -----305
Conclusions -----312
Literature Cited -----314
ABSTRACT
During the Early Cretaceous, ocean-floor spreading gradually opened up
the tropicalAtlantic and the Indian Ocean basin widened as the eastern segments
of Gondwanalandwere conveyed farther apart. At the same time, epeiric seas
were advancing on all continents, reaching maximum extent during the
Cenomanian. The resultanttrend to widespread, more equable climate favored
the invasion into the lowlands of angiosperms whose postulated origin was in
mild uplands at low latitudes during pre-Cretaceous times. As tropical and
subtropical lands were rafted farther apart by ocean-floor spreading following
Albian-Cenomaniantimes, new taxa (species, genera, tribes, families) evolved
in isolation. This accounts in part for the increasing richness of the three major
tropical floras following the Cretaceous. Changes in Mesozoic paleogeography
also appear to clarify several other puzzling aspects of early angiospermhistory
and distribution,including their early appearanceat middle latitudes.
INTRODUCTION*
For a number of years I have been concened with the time and place
of origin of angiosperms (Axelrod, 1952, 1959, 1960, 1961, 1966b). The
paucity of their record in rocks older than Cretaceous has generally been
* Some of the general principles and ideas presented in this article are an out-
growth of my studies of Tertiary floras in the western United States, a project
supported by the National Science Foundation (Grants G 9441, G 24339, G3 2625,
GB 4926, GB 7480).
277
278 THE BOTANICAL REVIEW
interpreted as resulting from rapid evolution from an unknown ancestor

just prior to their assumption of dominance over the lowlands. However,
I have favored the idea that their sparse occurrence in the fossil record
at earlier times resulted from their origin in, and occupation of, upland
habitats from which leaves and other recognizable organs were only rarely
preserved in the sedimentary record. I have also challenged the
hypothesis of angiosperm origin in high northern latitudes by the alterna-
tive of their radiation from equable uplands at low tropical latitudes.
From this standpoint it follows that the tropical floras of Cretaceous and
later times were derived by adaptation to hotter lowland climates; that
the mesophytic hardwood forests of the temperate Northern Hemisphere
developed from basically tropical alliances which spread into less equable
northern latitudes; that the broad-leaved evergreen forests of the temper-
ate Southern Hemisphere evolved from tropical taxa and have occupied
regions of highly equable climate throughout their history; and that the
floras of the dry regions have largely evolved in isolation since the middle
Cretaceous.
The present review brings into consideration new evidence that bears
on their antiquity, center of origin, and later evolution. This reassessment
has become necessary for several reasons. During the past few years new
principles regarding the nature of ocean basins and continents have been
discovered by geologists and geophysicists. There is now a reliable basis
for accepting the fragmentation of continental and oceanic plates and
their large lateral movement by the accretion of new ocean floor which
has spread the buoyant lands apart. This has a critical bearing on in-
ferences regarding the area and environment of early angiosperm evolu-
tion, and on their subsequent history. There also are new records of
angiosperms in pre-Cretaceous rocks that provide further evidence of
their antiquity, and recent reviews of Early Cretaceous floras of the USSR
make available critical evidence regarding their history in that region.
In addition, it is dlesirable to discuss the problem raised by palynologists
regarding the apparent absence of angiosperm pollen in rocks older than
the latest Aptian or earliest Albian, in rocks that yield numerous un-
doubted angiosperm organs. Finally, clarification of certain aspects of
early angiosperm history comes from recognition of the importance of
equability of climate, a factor that provides a measure of thermal ex-
tremes. Developed by Bailey (1960, 1964, 1966), the concept has been
applied to problems concerning the distribution and composition of dif-
ferent kinds of Tertiary and living forests (Axelrod, 1964, 1965, 1966a,
1966b, 1967a, 1967c), the altitude of Tertiary floras and topography
(Axelrod, 1965, 1968), and the nature of Cretaceous and Tertiary paleo-
climate in the western United States (Axelrod and Bailey, 1968, 1969).
Especially important to early angiosperm evolution is the crucial role that
equability appears to have played in the early adaptive radiation of
the alliance (Axelrod, 1966b, 1970), as well as in its rapid assumption
of dominance over the lowlands.
AXELROD: EARLY ANGIOSPERM HISTORY 279
SOME EARLY ANGIOSPERMS
At the present time, leaves that appear to be angiosperms are in the
Triassic (Sanmigueliain Colorado: Brown, 1956),' and there are leaves
and undoubted angiosperm (palm) wood in Jurassic rocks (Pro-
palmophyllumin France, Lignier, 1895, 1908; Sassendorfitesin Germany,
Kuhn, 1957; Phyllites in England, Seward, 1904; Palmoxylonin Utah,
Tidwell & others, 1970).2 Angiosperm fruits are in very early Cretaceous
rocks of France (Carpolithus in Valanginian: Chandler, 1958) and
California (Onoana in Hauterivian: Chandler & Axelrod, 1961). Leaves
occur locally and rarely in rocks of Hauterivian and Barremian age at
middle latitudes. Angiosperms are recorded at more numerous localities
across middle latitudes during the Aptian, but are still low in diversity
and subordinate to gymnosperms and ferns. They become more frequent
in most regions during the Albian, but it is only near the end of that
stage and in the Cenomanian that angiosperms surge to dominance, and
in great variety. In the light of this information it seems appropriate
to ask: Why did angiosperms take so long to assume dominance over
lowland regions?
PALEOGEOGRAPHY
That their rapid invasion of the lowlands may have resulted from
changing paleogeographic relations has been suggested (e.g. Seward,
1941, p. 379, 383; Krystofovich, 1946; Takhtajan, 1954, p. 12; Umbgrove,
1947, p. 291). However, the principal conditions and circumstances that
enabled them quickly to assume dominance in the later part of the Early
Cretaceous have not been identified. That climate was primarily re-
sponsible seems likely because it is the all-pervasive factor that controls
plant distribution. Furthermore, global land-sea relations-which greatly
affect regional climates-were being materially altered as angiosperms
commenced to enter the record in increasing number and diversity during
the Early Cretaceous.
1-Professor G. Ledyard Stebbins has pointed out (verbal communication, Dec.

1969) that the leaf blade of Sanmiguelia is closely matched by that of the Cyclantha-
ceae. Although Sanmiguelia does not have a midrib, the resemblances with Cyclan-
thaceae are otherwise remarkable considering they are separated in time by
approximately190 million years.
2 Some investigators (e.g. Krausel, 1956; Scott, Barghoorn and Leopold, 1960)
reject all records of pre-Cretaceousplants previously reported as angiosperms. While
some of them may not representangiospermsthese investigatorshave not demonstrated
that those which resemble palm (Propalmophyllum,Palmoxylon), poplar (Phyllites),
and magnolia (Sassendrofites) are either pteridophytes or gymnosperms. The recent
discovery of palm wood in the middle Jurassic of Utah (Tidwell and others, 1970)
was immediately discredited by many paleobotanists at the International Botanical
Congress (Seattle, 1969) on "stratigraphic"grounds. Through the cooperation of
Dr. Tidwell, I have examined the sites (April, 1970). There is no doubt that the
fossils occur in situ, in rocks certainly dated as Jurassic by both stratigraphic and
paleontologicevidence.
It was at this time that Gondwanaland ceased to exist. The continued

accretion of new ocean floor by upwelling lavas along the traces of the
deep rifts at the site of the present Atlantic and Indian mid-ocean ridges
gradually conveyed segments of Gondwanaland farther apart following
later Triassic time (e.g. Hess, 1960, 1962; Heirtzler & others, 1968;
Hurley, 1968; Le Pichon, 1968; Isacks, Oliver & Sykes, 1968; Vine, 1966,
1968). Current evidence indicates that whereas the North Atlantic basin
began to form as an ocean about 200 million years ago (middle Triassic),
and the South Atlantic about 150 million years ago (late Jurassic), the
middle Atlantic opened up only near the close of the Early Cretaceous,
about 110 million years ago (early Albian). Fragmentation, as shown
by the penetration of the oceans along the opening coasts, is now well
documented by stratigraphic, structural, and paleontologic evidence (e.g.
Allard & Hurst, 1969; DuToit, 1937; Freake, 1966; Hawkes & Smith, 1965;
Hoeken-Klinkenberg, 1964; Hurley & others, 1967; King, 1962; Melville,
1966; Martin, 1968; Reyment, 1967, 1969; Smith & Hallum, 1970). The
earliest marginal sediments on the Atlantic shores of tropical South
America-Africa are lagoonal Aptian, followed by marine Albian deposits.
Disruption along the east side of Africa had separated an eastern realm
of Madagascar-India-Australia by the early Cretaceous. As the Atlantic
and Indian Ocean basins widened by ocean-floor spreading, epeiric seas
encroached on all the continents and attained maximum extent in the
Cenomanian. Thermal extremes were progressively lowered as the ratio
of sea- to land-surface gradually increased, producing a general trend
toward increased equability.3 When taken in conjunction with the con-
ditions which preceded it, this trend appears to explain the rapid spread
of angiosperms over the lowlands, and also certain puzzling aspects of
their early history and later distribution. To develop these events
sequentially, evidence concerning the probable environment of pre-
Cretaceous angiosperm evolution is considered first.
PRE-CRETACEOUS HISTORY
Environment. It seems evident that angiosperms were not living
and evolving in lowland areas during pre-Cretaceous times. Numerous
Jurassic and Triassic floras are known from lowland regions throughout
the world, yet they contain few angiosperms, and plants that may
certainly be considered their progenitors are unrecorded. Since angio-
sperms made their entry into the lowlands in the Early Cretaceous, they
must earlier have inhabited sites remote from the major basins of ter-
restrial deposition (Axelrod, 1952, p. 33-35). Such a postulated occur-
3As used throughout this paper, equability refers to the degree of thermal
extremes. As described by Bailey (1960, 1964, 1966), it may be rated on a scale
from 100 (maximum) to 0 (minimum equability), defined by the departurein hours
of the year from an ideal isothermal condition of T (mean annual temperature)
57.20F (or 14?C) and A (mean annual range of temperature) 00. It may be deter-
mined by equation, or by nomogram (also see Axelrod & Bailey, 1969).
rence is not unique for angiosperms: it is inferred for the evolution of
all higher categories of land vertebrates and plants during their rapid
phase of divergence from an ancestral group (Simpson, 1944, p. 105-124;
1953, Chap. 11). Their general absence (or extreme rarity) in pre-
Cretaceous rocks seems explicable on the basis that the alliance that gave
rise to them was spreading out into a new adaptive zone, presumably in
equable, warm upland areas, to judge from their adaptive relations, and
from stratigraphic evidence as well.
There can be but little doubt that the angiosperm alliance is basically
adapted to warm, frostless climate. More than half of the 300-odd
families are found primarily in tropical to subtropical regions, and scarcely
extend beyond their margins. Another fourth find optimum development
there, and are represented outside the tropics by relatively few genera
(Axelrod, 1952). Furthermore, Bews (1927) has shown that there is a
pattern of evolution displayed by order after order, by family after
family, throughout the entire angiosperm phylum: in general, the more
primitive living taxa of each alliance are adapted to the more equable
warm temperate and montane tropical climates, whereas the derived
families and genera range into drier and colder regions, and into those
with hotter, drier, and less equable climates. This agrees with Camp's
(1952, p. 210-211) observation that the rainforest of the Amazonian low-
lands includes numerous taxa whose nearest ancestors occur in the
bordering hilly regions above the torrid lowlands. Takhtajan (1969, p.
166) also notes that in Southeast Asia: "despite the fact that the tropical
lowlands are characterized by the greatest diversity of species, genera,
families, and life forms amongst the angiosperms, they contain signifi-
cantly fewer primitive and phylogenetically intermediate fo-rmsthan the
mountain forest of tropical Asia and the subtropical forests of (that
region)." Takhtajan (1969, p. 170-171) emphasizes that in the eastern
Himalayas, Assam and southeast China: "Numerous existing phylogenetic
series can be observed in the flora... .there is an abundance not of isolated
'missing links', but whole chains of such links, and this makes it possible
to establish the pattern and direction of morphological and ecological
evolution of several exceedingly important lines of angiosperm develop-
ment. The phylogenetic series of forms transitional between subtropical
and temperate elements is clearly shown in many genera, families, and
even orders...."
This basic pattern of radiation from equable upland regions is con-
sistent with stratigraphic evidence, for Cretaceous angiosperms appear
in progressively younger rocks at higher latitudes (Axelrod, 1959;
Teslenko & others, 1966), implying radiation from warm latitudes to
cooler ones. Since stratigraphic evidence shows Cretaceous angiosperms
appearing in lowlands at equatorial latitudes after they entered the record
in middle latitudes, an earlier history under climates not so torrid as those
in tropical lowlands, and also more equable, is clearly implied. The
Cretaceous record thus suggests that angiosperms radiated from equable
climates in generally warm latitudes, to the less equable colder and hotter
and drier regions.
Some of their early development no doubt occurred on the united
Gondwana shield of South America-Africa-Madagascar-India-Australia.
Geologic maps reveal the absence of marine Jurassic and Triassic rocks
from large sectors of its crystalline basement, showing that at those times
much of it was an upland region. A survey of this broad terrain indicates
that there were scattered hills and mountains, as seen in the Guiana High-
lands (Venezuela to British Guiana) and the Massif de Ladamqua
(Cameroun). The region was well drained and composed of broad areas
of Precambrian metamorphic and igneous rocks, as seen now in the shield
areas of South America, Africa, Madagascar, India, and Australia-New
Zealand. Thus, geologic evidence shows that the nonmarine basins were
limited in number and frequently hundreds of miles removed from the
equable uplands where, it is inferred, scattered early angiosperm popula-
tions lived. From such sites there would have been little or no opportunity
for angiosperms to contribute to the pre-Cretaceous nonmarine record
accumulating locally in the lowlands. Furthermore, if the unknown
phylum that gave rise to angiosperms also inhabited this region during
the early Mesozoic, the absence of an ancestral alliance in the lowland
record, as well as taxa intermediate to angiosperms, becomes under-
standable.
Early Evolution. In searching for an upland tropical environment that
would be propitious for pre-Cretaceous angiosperm evolution, we can rule
out montane regions of ever-wet climate (e.g. "montane rainforest," or
"cloud forest") early in their history. Available evidence suggests these
areas would have supported dank tree fern glades, towering dense stands
of coniferophytes and ginkgophytes, and cycadophyte thickets. The
ground presumably was covered with a wet carpet of mosses, liverworts,
selaginellas, lycopodiums, and small ferns. Inasmuch as the rise of a new
alliance such as angiosperms involves the evolution of a new grade of
organization and the development of a wholly new way of living, wet
densely forested terrain obviously would not provide as favorable a
setting for their early evolution as open areas. Open sites were present
in the uplands at generally lower latitudes, in regions where there were
regular periods of seasonal drought. In such an environment angiospermy
would have high selective value in terms of the adaptive advantages of
enclosed (protected) fertilization and seeds. Whether angiospermy
originated as a response to seasonal drought is not certainly determinable,
but seems highly probable.
To judge from modern evolutionary studies, open areas in a region
of regular drought would provide an ideal setting for early angiosperm
evolution. These sites would be the more exposed, better-drained slopes,
dry intermontane valleys, and rocky slopes, areas where a dry season
would provide a powerful stimulus for rapid, continuing evolution. As
formulated by Stebbins (1952), and discussed further by Axelrod
(1967b), several factors would meet here in optimal interplay to favor
rapid, divergent evolution. It is recalled that many different specialized
structures can evolve concurrently which would adapt plants to live in
drier regions. Since the array of life forms possible in warm regions
subject to some drought is great, the potential for considerable adaptive
diversity would favor rapid evolution. Varied topography, soil, and ex-
posure have a greater control over plant distribution in areas where
moisture is limited than in humid regions. Diversity of local habitats and
geographic isolation combine to provide numerous opportunities for the
origin of isolating mechanisms, and for speciation. As populations become
subdivided into smaller units, new gene combinations become established,
and interdeme migration enables them to draw on a large supply of
genes. This interchange, which is enhanced by normal climatic fluctua-
tion, would be especially effective in disrupted, spatially isolated popula-
tions near the margins of their ranges. The genetic reorganization of
peripherally isolated, potential founder populations would favor rapid,
major change (Mayr, 1963; Lewis, 1966). In this connection, it is
emphasized that there is continuous flowering in areas of high equability
and adequate moisture. Under highly equable climates, therefore, small
founder populations in a diverse climatic-edaphic-topographic setting
might become established more readily. This would especially be true
at times of normal climatic fluctuation (e.g. "dry cycles," "wet cycles") in
regions of seasonal drought, for new adaptive combinations might have
high selective value for conditions somewhat warmer or cooler, drier or
moister, more equable or less, or for some new biotic relation (e.g.
pollination) closely attuned to one or more of the physical factors.
Finally, since all thermal zones (tropical to alpine) are compressed in
montane climates of high equability, they are spatially and thermally
closer. Therefore, rapid radiation into diverse thermal-moisture fields
is favored as compared with less equable regions; here tolerance for
greater extremes of moisture and/or temperature would be required for
entering the same zones (Axelrod, 1970). All these conditions must have
been met in tropical montane regions on the Gondwana shield. Terrain
was diverse; rock and soil type was varied; high relief favored regional
climatic gradients ranging from torrid to cool (or cold); precipitation
over the lowlands was strongly seasonal (see below) but at middle levels
rainfall increased and seasonality was less marked; equability decreased
from a maximum at middle altitudes to lower values at higher and lower
levels.
On this basis, position of the diverse terrain of Gondwanaland in
warm latitudes would favor the early evolution of taxa adapted to equable
montane tropical, temperate, and possibly alpine (depending on altitude)
climates at an early date. In terms of moisture-temperature gradients,
taxa would be radiating into diverse environments. Some would exploit
drier regions, and their derivatives might well return to more humid areas
(Stebbins, 1952; 1967). This early exploitation of a variety of equable
upland sites by divergent growth forms with different adaptive features

of floral and fruiting structures, would not only result in the emergence
of numerous categories, but would also stimulate evolution of unique
adaptive and morphologic alliances during early stages of their radiation
(Axelrod, 1952, p. 47).
CRETACEOUS HISTORY
Invasion of Lowlands. The question has frequently been asked: If
angiosperms were evolving in warm, upland areas during the pre-
Cretaceous, why did they not enter the lowlands during Jurassic or
earlier times? The paleogeography of the Gondwana shield provides a
basis for answering this question. In view of the large land area, low-
lands would be characterized by widespread, strongly monsoonal climate
to judge from climatic principles (Willett & Saunders, 1959, p. 199-210).
That such climate characterized the area from Triassic to Early Cretaceous
(Aptian) is demonstrated by the red bed, evaporite and aeolian deposits
in basins throughout the region (see maps in Narin, 1964). Furthermore,
broad belts of dry climate were also present across middle latitudes
of Eurasia and North America (see Takhtajan, 1954; 1969, p. 176-7;
Vakhrameev, 1964, 1966; in Narin, 1964). These relations suggest that
broad areas of torrid climate, with seasonally dry climate over the low-
lands, probably confined early angiosperms to upland areas where
extremes of temperature and drought were not severe. A similar climatic
relation occurs across lower latitudes today, for warn equable climates
are in the uplands, well removed from the torrid lowlands covered with
tropical rainforest, deciduous forest, savanna, grassland, or thorn scrub.
If angiosperms evolved initially in tropical uplands under climates
of high equability, they would appear first in the record at middle, not
at low latitudes. This follows because the warm temperate montane
zone of the tropics descends to sea level at middle latitudes, as may be
seen today in China, and also in eastern Australia and Brazil. Signifi-
cantly, dicot leaves in the Early Cretaceous floras from middle latitudes
(California, Portugal, Maryland-Virginia) are of moderate size (nanophyll,
microphyll, notophyll) and include taxa with entire margins as well as
those that are serrated or lobed. Such an assemblage is uniquely char-
acteristic of upland climates of high equability, the conditions postulated
for areas in which pre-Cretaceous angiosperms appear to have evolved.
Furthermore, a site in the uplands at generally low, warm latitudes is
also implied by the lack of angiosperms or their forerunners in Triassic
or Jurassic rocks at generally higher latitudes: their absence there in-
dicates that during those periods they must have been confined to areas
of warmer climate.
Pre-Cretaceous fold-belts (Umbgrove, 1947, see plates) were available
as upland dispersal routes to middle latitudes (Axelrod, 1952, fig. 11).
With some existing gaps they still connect the Gondwana shield areas
with Kazakhstan, Virginia-Maryland, California and Portugal, the regions
that yield the earliest Cretaceous angiosperms. Furthermore, old orogenic
belts are suggestively close to areas where fossils that represent angio-
sperns are recorded from Jurassic (Sassendorfites in Germany, Phyllites
in England, Propalmophyllum in France, Palmoxylon in Utah), and
Triassic rocks (Sarnmiguelia in Colorado). Since palms regularly inhabit
streambanks, it is not surprising that their remains have been discovered
in the pre-Cretaceous lowland records; presumably they were transported
down drainage-ways from the bordering, more equable and moister
uplands. Significantly, these fossils either occur with saline deposits
(Palmoxylon), or are closely associated stratigraphically with them
(Sanmiguelia), or are at least geographically near them (Sassendorfitesv,
Phyllites, Propalmophyllum). This is consistent with the inference that
pre-Cretaceous angiospenns were largely restricted to upland regions
by drier and hotter, less equable climates over the lowlands, an analysis
also consistent with the distribution of Early Cretaceous angiosperms
now known from. the Soviet Union. As Samylina (1968, p. 212)
emphasizes, they occur at middle latitudes (400 to 500 N) in both the
Siberian and Indo-European paleo-floristic regions described by
Vakhrameev (1964), but "within the latter they occur only in areas
outside the arid belt."
The gradual entry of angiosperms into the lowland record may thus
be correlated with the spread of moister, more equable climates that re-
sulted from the gradual widening of the Atlantic and Indian Oceans by
sea-floor spreading, and from the progressive inundation of all the con-
tinents during the Early Cretaceous which culminated in the Cenomanian
transgression. Evidence for a general trend toward climatic moderation
is well documented by late Jurassic to late Albian floras from the Kolyma
River area in northeastern Siberia (Samylina, 1964), by the early Jurassic
to Albian floras from most of the Asian platform (Vakhrameev, 1964,
1966), and by the Early Cretaceous floral sequences on the arctic coast
of Alaska (Smiley, 1966, 1967, 1969a, 1969b) .4 In each region evidence
for the northward shift of milder climate from the early into the medial
Cretaceous, is revealed by the shifting percentages of ferns, conifers,
cycadophytes, ginkgophytes, and angiosperms. Clearly, angiosperms
were entering the record during the Early Cretaceous trend toward
increased equability, and they rapidly increased in numbers and diversity
as it reached its peak.
Fig. 1 shows a hypothetical setting for Jura-Triassic to medial
Cretaceous angiosperm evolution. The sinuous boundaries imply that
in their movement to higher and lower altitudes and latitudes, the rate
of advance depended chiefly on local climate. The progressively younger
areas represent approximately the expanding regions occupied by angio-
4 All these investigators refer to a trend toward more tropical conditions,

or to a
general warming during the sequential change. It seems more probable that there
was not so much a major trend to greater warmth, but rather a significant increase in
equability (see Axelrod, 1968; Axelrod & Bailey, 1968, 1969).
C)~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~C
:3
90 60 30 0 30
N L a t i t u d e
FIG. 1. Generalized age-space relations of early angiosperm flora. The alliance radiated from
hotter lowlands and into cooler climates at higher altitudes and latitudes. Becaus
austral lands are still uncertain.
AXELROD: EARLY ANGIOSPERM HSTORY 287
sperms as climatic moderation increased. They are very rare in the
Neocomian, they are still rare though at more numerous localities in the
Aptian, they are uncommon and still subordinate in the early Albian,
and they surge to dominance at the end of that epoch and in the
Cenomanian as the equatorial Atlantic was fully established, and as
marine transgressions reached their greatest extent on all continents.
Evolutionary significance of Cretaceous megafossil-microfossil records.
The first angiosperms in the Cretaceous lowland record are megafossils
(leaves, wood, fruits) and they preceded definitely recognizable angio-
sperm pollen. The pre-Albian angiosperm megafossils do not display any
primitive features from a morphological standpoint; even the palm woods
(2 species) from the Jurassic are anything but primitive. As Samylina
(1968, p. 213) remarks in her review of the Early Cretaceous angiosperms
of the USSR: "As soon as we find fossil remains of obvious angiosperms
we immediately come across a great systematic diversity." By contrast,
Kemp (1968) and Doyle (1969) point out that the oldest angiosperm
pollen (probably latest Aptian or earliest Albian) is very simple and
presumably represents a very primitive or ancestral form. Doyle states
(1969, p. 28):
"The expansion and diversification of angiosperm pollen in the
Cretaceous is believed to reflect the basic adaptive radiation of the
group, within which morphological series documenting evolutionary
trends and the origin of major types may be recognized. Though
the angiosperms may have originated well before the observed radia-
tion, the idea that they are highly differentiated at their first ap-
pearance in the fossil record conflicts with the low diversity of Albian
angiosperm pollen and the regular sequential appearance of morpho-
logical types."
The factors that account for these divergent interpretations may be
considered in terms of the fossil sample and its interpretation. As for the
sample, medial Cretaceous megafossil and microfossil floras from the
same strata are often markedly different. A notable example is provided
by the low (5 per cent) representation of angiosperm pollen in the
Dakota formation (Stanley, 1967) of Kansas, for it has yielded a pre-
dominantly angiosperm leaf flora of no less than 200 taxa. Muller (1969)
points out that angiospermous pollen does not assume dominance until
post-Cenomanian time (Turonian-Senonian), yet Cenomanian and latest
Albian floras are wholly dominated by leaves of angiosperms that
represent numerous, diverse taxa. In the case of Kaolak River flora from
the arctic slopes of Alaska, of probable late Cenomanian age, Langenheim,
Smiley and Gray (1960) emphasize that the fossil pollen grains and
spores could not have been derived from plants in the megafossil record,
and that many of the latter are not represented by microfossils. Of the
factors that may account for the differences, failure of preservation and
selective preservation are significant. Some angiosperrn pollen (e.g.
laurels, Cercidiphyllum, Salix, Populus) is thin-walled and is not readily

preserved. If most angiosperms in the Early Cretaceous had thin-walled
grains, they would scarcely be expected to contribute to the record.
Furthermore, thin-walled grains frequently are destroyed during labora-
tory preparation so that a part of the record is lost in this manner. Selec-
tive sorting during transportation also produces an incomplete record.
Whereas the leaves of a tree carried by a current come to rest early as
velocity decreases on a floodplain, pollen and spores settle out of suspen-
sion at a more distant site-often in the sea in the case of Cretaceous
sequences-and are therefore lost so far as the terrestrial record is con-
cerned. Plants pollinated by wind may contribute to a record many
miles distant, but those pollinated by insects contribute relatively little,
even to one near at hand. If early angiosperms were largely insect
pollinated, very little evidence of them would be preserved even though
the alliance was already diversified.
Pollen of Early Cretaceous (or older) angiosperms may have differed
from that of most living angiosperms, so that its identification is not
possible or at least uncertain. The pollen may have resembled that of
ferns or gymnosperms, as does that of some few flowering plants today,
notably those at the middle level (distal aperture) of evolution discussed
by Kuprianova (1969); thus their absence may be more apparent than
real (Axelrod, 1961, p. 450). Furthermore, since the diversity of pollen
morphology was stimulated by entomophily, pollen evolution in some
measure has been independent of other characters on which taxa are
traditionally based (Kuprianova, 1969). It is tempting to suggest that
the increase in diversity of pollen taxa following the Cenomanian may
reflect primarily the rapid development of the crucial interrelation be-
tween the principal groups of pollinators and flowers-not necessarily
the adaptive radiation of the basic groups of angiosperms.
To judge from the record, diverse flowering plants were already
present when the first undoubted pollen grains of generally simple
morphology entered the latest Aptian or early Albian record: the basic
adaptive radiation of the group had already occurred.5 The Aptian woods
from England (Stopes, 1913) are not annonalian, or magnolian, or
hamamelidean, but are complex and structurally advanced in phyletic
position; Woburnia has been compared with Dipterocarpaceae (see
Boureau, 1957). Since there is no doubt about their stratigraphic position
(Casey, 1961, p. 566, 572), the distant hills from which -they were trans-
5"As far as adaptive radiation can be distinguished from progressive occupation

of numerous zones, a phenomenon with which it intergrades, the distinction is that
adaptive radiation strictly speaking refers to a more or less simultaneous divergence
of numerous lines all from much the same ancestral adaptive type into different, also
diverging adaptive zones. Progressive occupation of such zones is not simultaneous
and usually involves in any one period of time the change of only one or a few lines
from one zone to another, with each transition involving a distinctly different an-
cestral type" (Simpson, 1953, p. 223).
ported obviously supported diverse taxa of advanced morphology.
Furthermore,Chandler (in Chandler & Axelrod, 1961, p. 466) has
emphasized that the fruit of Onroanafrom rocks of Hauterivianage in
Californiais not primitive,but representsa plant that may provisionally
be referred to Icacinaceae, or a precursorof that family. This implies
that evolutionhad reachedthe level of Celastrales,an intermediateorder,
by the earliest Cretaceousat least. A mere glance at the leaves in the
Early CretaceousCheyenneflora of Kansas (Berry, 1922), the Blairmore
flora (Berry, 1929; Bell, 1956) of Canada, the Potomac of Virginia-
Delaware (Fontaine, 1889;Berry,1911), the Cercalof Portugal(Teixeira,
1948), the Shastaflora of California(Ward, 1905), or the floras of USSR
(Samylina, 1968), demonstratesthat diverse taxa had already evolved:
these angiospermleaves do not have any obviously primitive features.
Not only are they diverse in morphology,but distinct adaptive types had
already evolved. As Samylina (1968) emphasizes,angiospermsnot only
included small trees and shrubsin divergenttaxonomicgroups,they also
had adapted to aquatic habitats, as shown by the thin-textured,peltate
leaves rathersimilarto those of water-lily (Nelumbites) and Hydrocotyle
(Hydrocotylophyllum). Furthermore,leaf blades reminiscent of those
of a pandanus-likeplant (Pandanophyllum)are found in the Aptian,and
grass-likeplants (Carecopsis) are in the Albian. The latter taxa indicate
a relatively high degree of differentiationfor monocots, which is con-
sistent with the records of palms (Propalmophyllum,Palmoxylon) in
Jurassic rocks that are basically modern. The megafossil record thus
shows floweringplants were already diversifiedto a considerabledegree
in the Early Cretaceous,supportingthe view that their adaptiveradiation
took place much earlier (Axelrod, 1952,p. 44 48).
This is consistentwith evidence which indicates that regional floras
were already established during the Early Cretaceous. For example, in
the Siberian paleofloristic region Samylina (1968) notes that entire,
elongate dicot leaves account for more than one-half of the taxa as com-
pared with about one third in the Indo-Europeanpaleofloristic region
where impressions of trilobate and pinnate leaves, recalling those of
Aralia, Sassafrasand Crataegus,are more frequent. Furthermore,dicot
leaves in the Indo-Europeanpaleofloristic region are predominantly
entire-margined(12 of 22 species) whereas in the Siberianpaleofloristic
province serrate-marginedleaves are more numerous (24 of 42 species),
implying evolution in response to cooler climate at higher latitudes.
Comparisonof the leaves from the Early Cretaceousfloras of the Soviet
Union with those from the Potomac group in Virginia-Marylandpoints
up a significantdifference. Leaves from the latter areas are larger,which
is consistentwith their position at lower latitudes nearer the east coast
of a continentwhere warmerclimate would be expected. Furthermore,
some of the leaves in the Early Cretaceous Shasta flora of Califomia
(Ward, 1905) are microphyllousand thick, suggesting that angiosperms
were alreadyadaptedto dry climatein that region.
Another point that suggests we are not witnessing the adaptive radia-
tion of angiosperms during Albian-Turonian time is that this premise
calls for rates of evolution that are inconsistent with the observable
record. Scores of families and many tribes and genera that represent
ancient as well as derived groups are confined today to the inner tropics
and provide links between tropical America-Africa-Madagascar-India-
Australasia. They are at least as old as Albian-Cenomanian, their present
distribution having resulted chiefly from ocean-floor spreading which
conveyed these tropical lands farther apart. In addition, late Albian
and early Cenomanian floras have numerous taxa that represent advanced,
highly divergent adaptive types, including various Amentiferae, and
pollen compared with that of Sapindaceae or Proteaceae appears near
the end of the upper Albian in tropical Africa and Brazil (Doyle, 1969, p.
16-17). These records are separated by not more than 10 million years
from the earliest preserved undoubted pollen of the late Aptian-Albian.
Clearly, angiosperms could not have evolved to such a level in so brief
a time. The above-mentioned records of complex Aptian woods
(Woburnia, Aptiana), the Hauterivian fruit of Onoana, and the Jurassic
palm wood are also inconsistent with the view that the Albian and
younger pollen record provides us with a glimpse of the adaptive radiation
of angiosperms.
In this regard, an analysis of the climatic requirements of ancient
taxa that are still living (e.g. cycads, araucarians, tree ferns, podocarps,
redwood) demonstrates that they have survived chiefly in areas of very
high equability (Axelrod, 1964, p. 54-59; 1967a, fig. 4; Axelrod & Bailey,
1968, fig. 3). This is also true of the more "primitive" angiosperm al-
liances-magnoiioids, annonids, hamamelids-as well as the surviving
primitive members of the major angiosperm orders. Inasmuch as basic
adaptive radiation of angiosperms most probably occurred in relatively
open sites in equable tropical uplands characterized by some seasonal
drought (see above), they must have spread into mesic regions at a later
date. On this basis, the early angiosperm pollen may represent that of
ancient taxa which were shifting into moister, more equable areas-away
from the principal theatre of evolution. That the early Albian-Cenomanian
pollen is relatively simple seems clear, but the sequential relations suggest
not an ancestral evolutionary sequence resulting from adaptive radiation,
but a clisere that displays increasing complexity-a phemonenon common
in the history of land plants.
The Cradle of the Flowering Plants. Under this chapter heading,
Takhtajan (1969) has summarized evidence based on his own observa-
tions in Southeast Asia, as well as data presented by Smith (1967) and
others. He proposes that the extensive region between Assam and Fiji
may represent "a fragment of the ancient area which was first colonized
by the angiosperms" (p. 159). This view is based chiefly on the present
distributions of the more primitive living angiosperms. Takhtajan properly
asks the question: "Are we not taking a mere refugium, or center of
survival, of primitive angiosperns in the 'paradise garden' of the islands
and peninsulas of Southeast Asia for the cradle of the angiosperms and
their initial distribution center?" (p. 157). In attempting to answer the
problem, he notes that the geographic distribution of living angiosperms
is very different from that of the Tertiary, and even more so from that of
Cretaceous. However, living angiosperms do in some cases exhibit suf-
ficiently complete phylogenetic series, and their patterns of distribution
can be studied so much more fully, that they are of considerable signifi-
cance for a solution of the problem. He notes that the rich flora of tropical
America is significantly poorer in primitive angiosperms than the flora
of the southwestern Pacific basin, and that the flora of tropical Africa
is almost devoid of them. "It is difficult to explain why no members of
the Magnoliaceae, Winteraceae, Himantandraceae, Degeneriaceae, or any
other archaic family have survived in the African flora, if they had
existed there previously. Of the families of the Magnolia.les, only the
Canellaceae and the Annonaceae are in Africa. In Madagascar there
are rather more representatives of primitive families than in Africa,
as besides the Canellaceae and Annonaceae, we also find Bubbia
(Winteraceae) and Ascarinopsis (Chloranthaceae), as well as Didy-
melaceae. . . Although primitive angiosperms are appreciably more
numerous in America than in Africa and Madagascar, even in America
they are still far fewer and, as a rule, more specialized than in east and
Southeast Asia. The American members of the Magnoliaceae and
Winteraceae, for example, are markedly inferior in number and diversity
to those of the western Pacific and are representatives of the less primitive
forms. It is exceedingly unlikely that the concentration of so large a
number of primitive angiosperms in east and Southeast Asia, Australasia
and Melanesia can be explained solely by the presence in this region of
optimum conditions for their survival. The variation in paleogeographical
conditions in this region has not been less, but significantly greater, than
in tropical America and in tropical Africa. In Assam, upper Burma,
Yunnan, in the mountain regions of Thailand, Laos, and Vietnam, and in
Malesia and Melanesia grow several very primitive Magnoliales and also
many other 'living fossils'. When one considers this striking concentration
of primitive angiosperms, amongst which are not a few that are 'missing
links,' . . . one is forced to the conclusion that this region of the world is a
fragment of the ancient area which was first colonized by the angio-
sperms. Admittedly, this conclusion is still far from proven, but it is
very likely that the initial center of angiosperm expansion (and perhaps
also their center of origin) was situated somewhere in or near this region"
(p. 159).
Viewed from the standpoint of ocean-floor spreading and plate
tectonics, the Indo-Australasian region rich in ancient angiosperms is
composed of the eastern fragments of Gondwanaland which were rafted
apart following the Early Cretaceous. (See Paleogeography.) India
was separated from Madagascar in the Cretaceous and conveyed north
by sea-floor spreading to reach its present position during the late

Tertiary. Australia, and its continuation into New Caledonia-Fiji-New
Zealand, were rafted to their present positions during the Cretaceous and
Cenozoic. Since these eastern micro-continents were surrounded by seas
and were under the influence of "marine"climates, they were largely re-
moved from the more extreme climates which developed over the larger
land masses of South America and Africa. Survival in the equable uplands
from Southeast Asia into Australia-New Caledonia of more numerous
primitive living angiosperms may thus be correlated with a more favor-
able, equable climate during most of their history following the mid-
Cretaceous.
When the fragments of Gondwanaland are reassembled for the
Mesozoic, we find it was centered in the tropics. It is this region that
harbors the numerous taxa that form the continuous "phyletic trend line"
championed by Camp (1947; 1952): it is also the area that contains the
greatest concentration of angiosperm families, including those relatively
low, intermediate, and high in phyletic position (see Axelrod, 1952, figs.
3-9). All evidence presently available is consistent with the suggestion
that the "cradle" of early angiosperm evolution was in the uplands of
Gondwanaland, situated in tropical latitudes during the Mesozoic. From
there, angiosperms radiated into hotter and cooler climates, first with
respect to altitude and then with latitude-spreading poleward during
the Cretaceous as temperate Austral (evergreen) and Boreal (mixed
deciduous and evergreen) floras. Although evolution of the upland
tropical flora commenced in the pre-Cretaceous, the Neotropical and
Paleotropical floras were not isolated until the Cenomanian, when the
Atlantic was opened by ocean-floor spreading. There were more
numerous links between tropical and subtropical regions into Paleogene
time, but most had disappeared by the end of that epoch, leaving the
tropical floras of each region nearly as distinct as they are today.
CRETACEOUS PLANT GEOGRAPHY

The preceding discussion provides a basis for reevaluating certain
patterns of angiosperm distribution at generally low to middle latitudes,
two of which are reviewed here briefly.
Tropical Links. Numerous alliances provide links across the inner
tropics today (Vester, 1940; Hutchinson, 1946; Good, 1964; Camp, 1947;
1948; Croizat, 1952; Axelrod, 1952). There are 54 large pantropical
families that attain optimum development and diversity in tropical areas
and scarcely extend out of the region; 36 additional families are dis-
continuous in the inner tropics, linking America with Africa-Madagascar
(12), and Africa-Madagascar with the Indo-Australasian region (16), or
the latter area and America (8 families); and 125 other families attain
optimum development and diversity in the tropics, are largely pantropic,
and have generally derived and comparatively few taxa in extratropic
regions. Furthermore, some of the 80-odd families that are now endemic
to tropicalregions (25 American,23 African,40+ Australasian)occurred
earlier on other continents (e.g. Cyclanthaceaein India). Of the inner
tropical links, it is noteworthy that 287 genera are pantropic, 91 are
common to America and Africa-Madagascar,289 link the latter area
and the Indo-Australasianregion, and 37 are common to the Indo-
Australasianarea and the Americas.
When these groups are looked into a little more closely, we find that
related species of the same genera, or genera of the same tribes, or tribes
of the same family, are now separatedby broad tracts of tropical ocean.
These relations suggest the splitting apart of a more homogenous flora
by fragmentationof Gondwanaland,and subsequentevolutionin isolation
leading to divergenttaxa of the larger alliances. Some examplesof these
relationsare noted briefly.
1. A well known example is that of Gossypium (Hutchinson, Silow
and Stevens, 1947), where taxonomic sections are associated with dry
climates in the four tropical and subtropicalcontinents. Thus, the sec-
tions Erioxyla (Southern California-N. Mexico), Klotzschiana (Peru,
GalapagosI., BajaCalif.), and Thurberana(Ariz.-Mexico)are American,
Sturtianais Australian,and Anomalaand Stocksianaare centered in the
dry parts of Africaand Arabia. As discussedby Hutchinson,and Stevens
(in Hutchinson,Silow and Stevens (1947, p. 56) and also by Hawkes and
Smith (1965), the relations seem consistent with the breakup of
Gondwanaland.
2. Hawkes and Smith (1965) have reviewed the distribution of
Solanumwhich is pantropical.The subgeneraLycopersiconand Bassoyia
are in South America;Archaeosolanumis in Australiaand New Zealand;
and Lyciosolanumis African. Two other subgenera,Solanumand Step-
patipoliumare widespread and have been divided into 42 sections and
subsections,and 32 of these (about 75 per cent) are confineidto a single
continent. Thus, the taxonomicgroupings agree with the geographical
separationof South America-Africa-India-Australia-New Zealand, all of
which are componentsof Gondwanaland.
3. In his discussion of inner tropical links, Camp (1947) presented
numerousmaps to show that there are close ties acrossthe inner tropics.
(a) In the tribe Agaveae, the genera Agave, Furcraea,and Beschorneria
are in America,but the related Doryanthes is Australian(Camp, 1947,
fig. 11). (b) In the family Strelitziaceae,Heliconia is Americantropical,
Strelitziais in tropicalAmericaand Africa-Madagascar, Phenakospermum
inhabits eastern tropical South America,and Ravenalais in Madagascar
(Camp, 1947, fig. 20). (c) In the tribe Dianelleae (Liliaceae), Walleria
is central east-African, Dianella is widespread through the Indo-
Australasianregion, reaching North Island, New Zealand, Stypandrais
Australian,and Excremisis in Chile (Camp, 1947, fig. 9).
Others (e.g. Engler, 1905, Hutchinson, 1946, p. 12-14; Camp, 1947;
Good, 1964; Aubreville, 1959) have pointed out that trans-Atlanticcon-
nections are suggested by a number of families that are commonchiefly
FIG. 2. Distribution of 3 families (1-Mayacaceae, 2-Rapateaceae, 3-Bromeliaceae) that
and subtropics, and have only isolated occurrences of a single genus in tropical Afr
aceae (15 genera, 1400 species) into the United States represents the genus Tillands
to tropical America and Africa. For instance (see Fig. 2), the large
family Bromeliaceae (60 genera, 1400 species) is entirely American
except for a single species of Pitcairnia which is endemic to Guinea. The
Vochysiaceae (9 genera, 187 spp.) have all of their species in tropical
America except for the genus Erismadelphus (2 spp., trees) in the rain-
forests of Cameroon and lower Guinea. The Rapateaceae (16 genera, 80
spp.) are centered in the dense tropical forests of eastern South America
but are also represented by a monotype (Maschalopcephalus) which is
endemic to the rainforest of Liberia, west Africa. The family Velloziaceae
has two genera, with Vellozia in south Arabia, tropical and South Africa,
Madagascar, and tropical South America, and Barbacenia endemic to
Brazil. The Mayacaceae form a monogeneric monocotyledonous family
of aquatic herbs, with one species (Mayaca baumii) in Angola, west
Africa, and the remainder in tropical America. This finds a parallel in
the distribution of the family Loasaceae, which is chiefly in the warmer
parts of the Americas, but has one genus (Kissenia) in Africa, with two
species-one in South Africa and the other in Somalia and adjacent Aden.
In a broader, more significant way, the distribution of genera of
tribes of large typically tropical families provides indirect, though reliable
evidence for ancient connections across the Atlantic. Aubreville (1959)
has summarized the distribution of the tribes of the families (or sub-
families) of Leguminosae in tropical Amazonia and in equatorial Africa.
Distribution of the genera of the respective tribes of these families (8
tribes in Caesalpiniaceae, 5 in Papilionaceae, and 4 in Mimosaceae)
shows that they have an unequal representation in tropical America
(Amazonia) and tropical Africa. The data for the Caesalpiniaceae are
as follows (Aubreville, 1959, p. 52):
Genera in forests Genera in forests
of equatorial Africa of Amazonia
Bauhinieae 1 1
Amphimanteae 1 0
Cassieae 4 6
Cynometreae-Amherstieae 50 13
Swartzieae 3 4
Eucaesalpineae 1 2
Dimorphandreae 4 2
Sclerolobieae 0 11
Inasmuch as these are inner-tropical alliances, they not only indicate
ancient connections across the Atlantic, but suggest that some of the taxa
have evolved independently in each region.
The evolutionary results of the separation of inner tropical lands by
ocean-floor spreading during the later Cretaceous are suggested by the
distribution of genera that represent natural tribes of large pantropical
families. The tribes of these alliances have distributions that may include
only one, or two, or all three major tropical regions-America, Africa-
Madagascar, and the Indo-Australasian region. Recalling that today
there are only about 280 pantropical genera and that more numerous
17- ~ ~ ~ ~ ~ ~ ~ ~ ~ ~
/ 3. ,-.- /,
~~~~~~~~~~~~~2
FIG. 3. Distribution of 5 tribes (34 genera) of the subfamily Moroideae (Moraceae): 1-Fato
Strebleae, 5-Dorstenieae. Of the tropical genera, 2 are American,4 are confined to A
2 are on Madagascar, 18 are in the Australasianregion, and 2 link the latter area w
pantropical and it also reaches into temperate regions. Only one genus (Maclura) is
genera were common both to the Old and New World well into Paleogene
time, it is evident that at a minimum, the genera of the respective tribes
have largely developed since ocean-floor spreading separated lands
formerly connected. In this study, the distribution of genera of tribes
representing the following 18 families was plotted, using the data in
Hutchinson (1964), as a source:
Annonaceae Dilleniaceae Monimiaceae
Araliaceae Flacourtiaceae Moraceae
Bombacaceae Lauraceae Myristicaceae
Connaraceae Magnoliaceae Passifloraceae
Canellaceae Malpighiaceae Tiliaceae
Erythroxylaceae Mimosaceae Winteraceae
Inasmuch as the same general pattems of distribution are repeated, the
relations of only several of them are summarized here; several figures are
included to illustrate the general areal relationships of the tribes.
Moraceae. Fig. 3. This large family of 76 genera and over 1000
species is composed of 3 subfamilies, Moroideae with 5 tribes, Arto-
carpoideae with 4 tribes, and Concephaloideae which forms a natural
alliance of 8 genera. Only 3 of the genera in the entire family are
pantropical, notably Ficus, Artocarpus and Morus. Only two genera
(Maclura, Morus) reach into temperate climate, and most live in regions
well removed from frost. The distribution of the 5 tribes of the subfamily
Moroideae reveals that two occupy the Old World and find optimum
development in the southeast Asian-Melanesian region. The other three
tribes have a pantropical distribution, though most of the genera occur
only in one tropical region. Distribution of the genera of the Moroideae
is as follows.
Tribe 1. FATOUEAE. This small tribe of 4 genera is confined to the
Old World tropics, and two of the genera (Fatoua, Bleekrodea) link
both of the African-Madagascar region with that of Southeast Asia and
Melanesia. Pseudostreblus: 1 sp., India and Hainan; Fatoua: 3 spp.,
Old World tropics; Ctenocladus: 1 sp., West tropical Africa, Cameroons;
Bleekrodea: 3 spp., Madagascar, Indo-China, Borneo.
Tribe 2. MOREAE. This tribe of 11 genera has only one genus
(Morus) that is pantropical. The other genera are confined to single
regions, with most occurring in the area from Southeast Asia to Melanesia
and Polynesia. Paratrophis: Malaya to Polynesia and New Zealand;
Dimerocarpus: 1 sp., Indo-China; Pseudoirophis: 2 spp., New Guinea;
Pseudomorus: 3 spp., New Guinea to Polynesia and Norfolk I.; Morus:
60 spp., tropical and temperate regions; Ampalis: 1-3 spp., Madagascar;
Pachytrophe: 2 spp., Madagascar; Trophis: 4 spp., tropical America;
Metatrophis: 1 sp., Polynesia; Calpidochlamys: 2 spp., New Guinea;
Smithiodendron: 1 sp., SW China.
Tribe 3. BROUSSONETIEAE. The genera of this tribe have a
pantropical distribution, but only one genus (Chlorophora in America-
Africa) occurs in more than one tropical region. Maclura: 6 spp., North
16~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~
FiC;.4. Distribution of 6 tribes (23 genera) of the family Bombacaceae: 1-Durioneae, 2

mateae, 5-Adansonieae, 6-Ceibeae. Four tribes are American tropical, one (Durio
the other (Adansonieae) is pantropical, with one genus (Bombacopsis) linking Am
tropical Africa-Madagascar-NWAustralia, and with the others more localized.
America; Chlorophora: 11 spp., tropical Africa, tropical to temperate
South America; Bagassa: 3 spp., tropical South America; Broussonetia:
6 spp., eastern Asia, Indo-China, Malaya, Polynesia; Malaisia: 3 spp.,
eastern Asia, Malaya, tropical Australia, Polynesia; Allaeanthus: 5 spp.,
Indomalaya, China, Philippine I.; Cardiogyne: 1 sp., tropical and sub-
tropical Africa; Plecospermum: 2 spp., tropical Asia.
Tribe 4. STREBLEAE. Of the 6 genera in this alliance, only one
(Maillardia in the Mascerene I.) is found outside the southeast Asian-
Melanesian region. Taxotrophis: 7 spp., China, Indomalaya; Diplothorax:
1 sp., Tonkin; Streblus: 3 or 4 spp., Indom.alaya, Philippine I., New
Guinea; Teonongia: 1 sp., Indo-China; Phyllochlamys: 4 spp., Hainan,
Indo-China and Indo-Malaya; Maillardia: 5 spp., Mascerene I.
Tribe 5. DORSTENIEAE. This tribe of 5 genera is pantropical, and
one of the genera (Dorstenia) occurs in each tropical region though it
is rare in the Asian tropics. Sloetia: 5 spp., Malaya; Sloetiopsis: 1 sp.,
east tropical Africa; Noesloetiopsis: 1 sp., tropical Africa; Dorstenia: 200
spp., tropical, but rare in Asian region; Craterogyne: 4 spp., tropical
Africa.
Bombacaceae. Fig. 4. This tropical family of 31 genera and about
225 species represents highly specialized members of the Tiliales which
have retained their woody character. Of the 6 tribes in the alliance 4
are restricted to the American tropics, one is Australasian, and the other
is pantropical. The taxa are distributed as follows:
Tribe 1. DURIONEAE. All 8 genera are in the Asian-Australasian
region. Neesia: 8 spp., Burma, Thailand, Malaya; Papuodendron: 2 or 3
spp., New Guinea; Durio: 25 spp., Burma, Malaya, Philippines;
Camptostemon:2-3 spp., N. Australia, N. Borneo, New Guinea, Philip-
pines; Coelostegia: 5 spp., Malaysia; Boschia: 6 spp., tropical Asia;
Kostermansia: 1 sp., Malay peninsula; Cullenia, 2 spp., Ceylon, S. India.
Tribe 2. HAMPEAE. 3 genera, restricted to American tropics.
Hampea: 112 spp., Mexico and Honduras; Scleronema: 4 spp., American
tropical; Cavanillesia: 5 spp., tropical South America.
Tribe 3. MATISIEAE. 8 genera, confined to the American tropics.
Quararibea: 12 spp., tropical America; Ochroma: 11 spp., American
tropical; Mati*ia: 25 spp., Central and tropical South America; Patinoa:
2 spp., Colombia, Brazil; Bernoullia: 2 spp., tropical American;
Septotheca: 1 sp., Brazil, Peru, Colombia; Phragmotheca, 1 sp., Colombia;
Huberodendron: 2 spp., Brazil.
Tribe 4. CATOSTEMMATEAE. Two genera, both in Brazil.
Aguiaria: 1 sp., in Brazil; Catostemma: 4 spp., tropical South America.
Tribe 5. ADANSONIEAE. This tribe of 7 genera is pantropical in
occurrence. Four of the genera are American, one links the American-
African tropics, one links Africa-Madagascar with Australia, and the other
ranges from New Guinea to India. Bombax: 8 spp., Old World tropics,
India and Ceylon to New Guinea; Adansonia: 12 spp., Africa-Madagascar,
FIG. 5. Distribution of 3 tribes (18 genera) of tropical family Dilleniaceae: 1-Delimeae,
limeae (6 genera) is American tropical except for Tetracera (130 spp.), the only pa
lenieae (7 genera) has 3 genera of local distribution in the Australasianregion. Of
region with Madagascar, and Neowormia is endemic to the Seychelles. Tribe Hibb
Caledonia-New Guinea region, and one genus (Hibbertia) provides a link with Mad
northwest Australia; Pseudobombax: 20 spp,., tropical American;
Eriotheca: 20 spp., tropical American; Bombacopsis: 20 spp., all of which
are in tropical America and 1 or more in tropical Africa; Pachira: 2 spp,.,
tropical American; Gyranthera: 2 spp., tropical American.
Tribe 6. CEIBEAE. Three genera, all confined to the American
tropics. Ceiba: 10 spp., tropical and subtropical America; Chorsia: 9
spp., tropical South America; Neobuchia: 1 sp., Haiti.
Dilleniaceae. Fig. 5. This tropical family is composed of trees, shrubs
and vines distributed among 18 genera and about 530 species, which
are grouped into 3 tribes. Delimeae (6 genera) is American tropical
except for Tetracera (130 spp.) which is the only pantropical genus in the
family. Tribe Dillenieae (7 genera) is chiefly southeast Asian, but has
links with African region. Tribe Hibbertieae (5 genera) is primarily
Australian but one genus (Hibbertia) also occurs in Madagascar, New
Guinea and New Caledonia. As listed below, some of the genera in each
tribe have very restricted occurrences.
Tribe 1. DELIMEAE. This alliance of 6 genera is entirely American
except for the pantropic genus Tetracera. Empedoclea: 1 sp., Brazil;
Tetracera: 130 spp., tropics and subtropics, including Australia;
Curatella: 6 spp., tropical America, including West Indies; Pinzon: 2 spp.,
Brazil; Doliocarpus: 40 spp., Central and tropical South America;
Davilla: 40 spp., Central and tropical South America, West Indies.
Tribe 2. DILLENIEAE. The tribe of 7 genera is entirely Old World,
centered in southeast Asia, and with links to Madagascar and Seychelles.
Reifferscheidia: 1 sp., Philippine Is.; Wormia: 50 spp., tropical Asia to
Australia, Fiji and Madagascar; Neouwormia: 1 sp., Seychelles; Dillenia:
70 spp., Indo-Malaya, Philippine I.; Schumacheria: 3 spp. Ceylon;
Didesmandra: 1 sp., Borneo; Acrotrema: 10 spp., Southern India, Ceylon,
Malay peninsula.
Tribe 3. HIBBERTIEAE. Five genera, in New Guinea, Australia,
New Caledonia, and with one linking the region to Madagascar. Trisema:
7 spp., New Caledonia; Hibbertia: 150 spp., New Guinea, New Caledonia,
Australia, Madagascar; Candollea: 20 spp,., W. Australia; Adrastaea:
1 sp., E. Australia; Pachynema: 5 spp., Australia.
Tiliaceae. Fig. 6. This large family of 67 genera and about 1500
species is found chiefly in the tropics, more rarely in the subtropies,
and is composed principally of trees and shrubs. Tilia is the only genus
that reaches into temperate regions. The family includes 12 tribes
(Hutchinson, 1967), and their distribution suggests that some have
probably developed in isolation. Their general distribution can be sum-
marized as follows. A. Tribes with genera in all three tropical regions:
Enteleeae, Tilieae, Grewieae, Triumfetteae, and Brownlowieae. Only 3
of the 24 genera among these 5 tribes have a pantropical occurrence,
notably Corchoris (100 spp.), Grewia (400 spp.), and Triumfetta (160
spp.). B. Tribes with genera in two or more tropical regions: America-
8~~~~~~~~~~~~~~
F. Dtuo f tb ( ge of iTa e Po e pi a
7~~~~~~~~~~~~~~~~~~
Fic. 6. Distribution of 12 tribes (67 genera) of family Tiliaceae. 1-Prockieae, 2-Akpeibeae,

Corchoropsideae, 6-Sparrmanieae, 7-Lueheeae, 8-Tilieae, 9-Desplatzieae, 10-G
lowieae. Five tribes have genera in all tropical regions, but only 3 genera-Corchoru
iinfetta (160 spp.)-are pantropical. Tribe Apeibeae links the American-Africantro
eae link the Arnerican-Australasian tropics. Tribes that are confined to one tropical a
Pseudocorchoreaein Madagascar, and Sparrmanieaein Africa.
Africa: Apeigese (3 genera); America-Asia: Prockieae (5 genera),
Lueheeae (3 genera), Desplatzieae (4 genera). No genera link only
Africa and Asia. Among the genera with very restricteddistributionsare
Hainania (Hainan), Erinocarpus(Bombay), Tetralix (Cuba), Tahaitia
(Tahiti), Brachypodandra(Tonkin), and Eleutheranthus(New Guinea).
Tribe 1. PROCKIEAE. This tribe of 5 genera is entirely American
except for the genus (Plagiopteron) in Burma. Prockia: 18 spp., tropical
America,West India, Argentina;Hasseltia: 10 spp., Centraland tropical
America; Hasseltiopsis, 3 or 4 spp., Mexico, Central America, Peru,
Colombia; Plagiopteron: 1 sp., Burma; Neosprucea: 2 spp., tropical
South America.
Tribe 2. APEIBEAE. Three genera are confined to the American
or African tropics. Glypheae: 4 spp., tropical Africa; Apeiba: 15 spp.,
tropicalAmerica;Ancistrocarpws:2 or 3 spp., tropicalAfrica.
Tribe 3. ENTELEEAE. Pantropicalin occurrence,but 5 of the 6
generahave restrictedoccurrences.Entelea: 1 sp., New Zealand;Nettoa:
1 sp., island of Australia;Mortoniodendron:5 spp., Mexico and Central
America;Burretiodendron:3 or 4 spp., SW China, Burma,Indo-China;
Sicrea: 1 sp., Indo-China;Corchorus: 100 spp., pantropic.
Tribe 4. PSEUDOCORCHOREAE.Pseudocorchorus:6 spp., Mada-
gascar.
Tribe 5. CORCHOROPSIDEAE.Corchoropsis:4 spp., easternAsia.
Tribe6. SPARRMANIEAE.3 genera,all African-Madagascar.Sparr-
mania: 4 spp., tropical and South Africa, Madagascar;Clappertonia: 2
spp., tropicalAfrica;Cephalonmema: 1 sp., tropicalAfrica.
Tribe 7. LUEHEEAE. 3 genera in the Americantropics, and one of
them occurs also in SoutheastAsia. Luehea: 40 spp., Mexico to tropical
South America, Colombia, Brazil; Trichospermum:25 spp., tropics of
Americaand Asia; Mollia: 18 spp., tropical South America.
Tribe 8. TILIEAE. Pantropical,with 4 of the 5 genera centered in
the tropics,and the other (Tilia) ranging into temperateregions. Tilia:
80 spp., temperate Northern Hemisphere, reaching south into central
Mexico; Duboscia: 4 spp., West Tropical Africa; Muntingia: 3 spp.,
tropical America,Mexico, CentralAmerica,West Indies, tropical South
America;Brachypodandra:1 sp., Tonkin;Schcutenia: 11 spp., Malaysia,
Borneo, Java.
Tribe 9. DESPLATZIEAE.4 genera, in the American and African
tropics. Desplatzia: 6-8 spp., chiefly tropicalwest Africa;Cotylonychia:
1 sp., tropical Africa;Vasivaea: 2 spp., tropical South America;Hydro-
gaster: 1 spp., Brazil.
Tribe 10. GREWIEAE. Pantropicalin distribution,but best repre-
sented in the region from SoutheastAsia into New Guinea. Grewia: 400
spp., tropics and subtropics chiefly, rarely into mild temperate areas;
Eleutherostylis:1 sp., New Guinea; Tetralix: 2 spp., Cuba; Althoffia:
5 spp., Philippine Is., Malaya archipelago,New Guinea; Luehopsis: 6
spp., easterntropicalSouth America;Colona: 30 spp., S. Yunnanthrough
Malaya, Philippines Is., to New Guinea and east Pacific Is.; Diplo-
phractum: 3 spp., Indo-China, Malaya.
Tribe 11. TRIUMFETTEAE. Pantropical in distribution. Erino-
carpus: 1 sp., Bombay; Triumfetta: 160 spp., tropics and subtropics;
Heliocarpus: 11 spp., Central America and tropical South America.
Tribe 12. BROWNLOWIEAE. Composed of 3 genera, each of which
is in a different tropical region. Christiania: 2 spp., tropical Africa;
Brownloowia: 30 spp., SE India through Malaya, Philippine Is. to, New
Guinea and east Pacific Islands; Pentaplaris: 1 sp., Costa Rica.
Similar relations have been documented for other alliances, notably
the leafy hepatics (Fulford, 1951), rainworms and scorpions (in Dutoit,
1937, see figs. 43, 44), copepods (Sewall, 1956), and non-marine molluscs
(Pilsbury, 1911). The distribution of angiosperm families, or sections,
or genera, as well as other alliances (mosses, scorpions, etc.), in two or
more tropical areas today is not to be explained by migration around
higher latitudes of Holarctica, from one tropical area to another, as has
been suggested (e.g. Sharp, 1966; MacGinitie, 1969). In view of the
high sensitivity of tropical taxa to chill or frost, it seems probable that
these alliances have always been confined to tropical or subtropical
regions because the climatic (physiologic) requirements of larger taxa
are genetically controlled and circumscribe natural alliances just as do
their morphologic features. Furthermore, the so-called "tropical" and
"subtropical" fossil plants in early Tertiary floras at middle to higher
latitudes lived under climates of high equability and were members of
mixed temperate, not tropical, rainforests. This is in accord with fossil
evidence which shows that tropical to subtropical forests have never
lived at higher latitudes. Equally convincing is the fact that there are
no records of tropical forests in eastern Asia at the latitude of Japan, for
they would be expected there in Eocene-Paleocene times if there was
interchange of taxa between low tropical latitudes via the Beringian
region. The forests recorded from Japan are-at the most-no more than
warm temperate (ET 60(F at a maximum), and most probably could
have lived under warmth ET 58?F to 590F if equability was high (+ M
65), as it was.
As first documented and discussed by Engler (1905), and reiterated
by others (e.g. Hutchinson, 1946, p. 12; 133-135; Camp, 1947; 1948; 1952;
Good, 1964; Boughey, 1957; Croizat, 1952; Axelrod, 1960; Aubreville,
1969), these distributions indicate former east-west connections across the
tropics. Foundered continents, drifting continents, isthmian links, and
island steppingstones have all been called upon to explain the relations.
It is now evident that the taxa occupied the region prior to the breakup
of Gondwanaland (Boughey, 1957; Hawkes & Smith, 1965; Melville,
1966; Aubreville, 1969), and the rafting of its segments apart by ocean-
floor spreading. If this was the case, lowland floras close to the suture
line, represented now by the opposing shores of the tropical Atlantic,
must have been essentially homogeneous into medial Cretaceous time.
This is confirmed by the composition of the Cretaceous spore-pollen
floras recovered from sedimentary rocks in Brazil (Sergipe) and Congo/
Gabon, which reveal 34 of 39 taxa common to each area (Freake, 1966).
There are also marked similarities between the Cretaceous pollen floras
of Nigeria and Colombia (Hoeken-Klinkenberg, 1964), and comparable
relations are revealed by fresh-water ostracods and fish (in Reyment,
1967, 1969).
The preceding data lead directly to the general conclusion that the
angiosperm, families and genera (excluding widely distributed littoral
plants) that now link the tropical regions probably are at least as old
as the Albian-Cenomanian transition (100 million years), and they may
be considerably older (cf. Hawkes & Smith, 1965, p. 49). This means that
bradytely pervades the phylum to a far greater degree than has been
previously supposed. Since the transtropic links represent "primitive,"
"intermediate," and "advanced" alliances, angiosperms must have been
in existence long before the Cretaceous. Rare finds of plants which evi-
dently are angiosperms in Jurassic and Triassic rocks support an early
dating of angiosperm origin; it may well have commenced in the late
Permian.
Desert Links. The desert floras of the Northern and Southern Hemi-
spheres are now separated by broad tracts of tropical forests and savannas.
Those in the Southem Hemisphere (South America-Africa-Australia) are
isolated by wide oceans, those in the Northern (southwestern North
America-southern Eurasia) by ocean and frozen lands. The floras of
each of these desert regions are largely distinct today. They have long
been isolated and have largely had separate origins and histories. Never-
theless, in spite of their isolation there has been interchange between
them. Since links between the herbaceous floras of North and South
America have recently been reviewed (Raven, 1963), attention here is
focused chiefly on problems raised by the woody plants.
It is essential to keep in mind that the floras of each of the 6 major
warm-desert regions are characterized by two contrasting alliances in
terms of their general affinities. One group, which includes by far most
of the taxa in each desert, either ranges outside of the desert into
bordering communities, or if the species are confined to the desert they
are related to alliances in moister regions; the second group is relict,
with no close allies today. As for the first group, judging from the degrees
of difference the taxa display-from plants that are scarcely distinct
varietally, to subspecies, to distinct species, to unique species, to paired
genera, to related genera-plants in each of the dry regions have been
adapting to increasing aridity during the Cenozoic. Most of this evolution
did not take place in regional deserts, but in local dry areas. In these
sites, where evolution was speeded up (Axelrod, 1967b), taxa became
adapted to increasing aridity. Thus they were able to expand with
progressively drier climates commencing in the early Eocene as forests
and savannas withdrew from areas presently desert.
Although ready interchange of taxa between the deserts in the

Northern and Southern Hemisphere is limited today by tropical forests
and savannas, this barrier was broader and far more effective in the
Tertiary and Late Cretaceous. The fossil record shows that forests and
savannas inhabited all the present desert areas in the Cenozoic and
also in the Late Cretaceous (see Axelrod, 1950). They have been re-
corded from the deserts of Egypt, the Red Sea area, western India-
Afghanistan-Kazakhstan; from southwest Africa; from south Australia;
from the Patagonian-Argentine and Chilean deserts, and the desert of
Bahia in South America; and from the North American desert in south-
eastern California, southern Nevada, southern Arizona, northern Baja
Califomia and the northern Chihuahuan region. It is apparent that im-
portant interchange was scarcely possible between floras of dry areas
during most of the early Tertiary and Late Cretaceous, but gradually
increased following early Eocene time as dry climate commenced to
expand.6
To judge from the available record, it was during the later Cretaceous
and earliest Tertiary that a number of taxa gradually differentiated from
widely distributed species of savanna-border areas, giving rise to related
(paired) species in distant areas of dry climate. This appears to explain
the development of similar species in the desert and desert-border areas
and in the bordering thorn scrub and deciduous tropical forests of the
Americas, and presumably in the Old World as well. The plants involved
are in genera such as
Acacia Pithecolobium Randia
Condalia Caesalpinia Zizyphus
Bursera Cercidium Ficus
Dodonaea Passiflora Lycium
Erythrina
That dry-adapted taxa had already evolved by early Paleogene time is
apparent from the record, for the Florissant (MacGinitie, 1953) and
Green River (MacCinitie, 1969) floras contain numerous taxa of this
alliance. Furthermore, a pollen flora near Del Mar, California, contains
a number of plants of similar relations (W. S. Ting, oral communication),
indicating they were already developed by early Eocene time.
The second group typical of desert and semidesert areas is represented
by taxa (mostly genera, some families) that have no close living relatives;
many of them are unique in adaptive type. Representative of this group
in southwestern North America are Fouquieria, Pachycormus, Chilopsis,
Pachycereus (or other cacti), Koeberlinia, Holocantha, Yucca, Sim-
mondsia; in the desert stretching from northwest Africa across Asia Minor
and eastward, are Adenia, Alhagi, Balanites, Genitsia, Haloxylon,
6
There was interchange between the floras of the drier and cooler parts of
western North America and South America in the late Tertiary and in the Quater-
nary-a topic not probed here (see Raven, 1963, for an excellent review).
Pyrenacantha,Randonia,Tamarix,and Zilla;in SouthAfricaare Aeonium,

Euphorbia (cactoid forms), Aloe, Ceraria, Conophytum, Eucela,
IHaworthia,Hoodia, Lithops, Moringa, Pachypodium,and Sarcocaulon;
in the drier parts of the Argentine-Chile deserts are Donatia, Browningia,
Puya, Grabouskya, Monttea, Bredemeyera, Bulnesia, and Zuccagnia;
while the Australian desert is marked by Amperea, Beyeria, Bertya, Bous-
siaea, Kingia. Since they include isolated monotypes in their respective
families and are unique in adaptive type, their pre-Tertiary origin is
strongly indicated. Thus a part of the problem is clarified: desert floras
contain taxa of different ages (Johnston, 1940; Axelrod, 1952; Rzedowski,
1962). One group has been developing gradually in response to a trend
to aridity during the Cenozoic, but the other is much older and its
history is obscure.
With respect to their evolution, the unique adaptive types are found
preponderantly in the southern deserts. They exhibit conspicuous modi-
fications in vegetative habit which give rise to bizarre life-forms that
make them seem out-of-place in their respective floras: they "look
ancient." Some show a tendency to leaf reduction, the development of
short shoots, and aphylly. Others are knobby forms with dense leaf
clusters; some have stems covered with scale-like leaves; others have
spine-tipped leaves; shrubs are commonly divaricately branched and
very spinescent. These modifications are not confined to plants of the
austral deserts. They occur regularly in other plant formations in austral
lands. They typify the thorn scrub of South America-Africa-Australia
which is marginal to and grades into the desert, and many are in the
drier savannas which border the austral thorn scrub. Plants of bizarre
habit also contribute to the monte and chaco of Argentina that live under
mild equable climate, and they make up similar communities in South
Africa and Australia as well. They are on the Patagonian steppes, and
in the dry alpine (puna) zone of the Andes and in the Cape Highlands
of South Africa; furthermore they contribute to the moist paramo of the
high tropical Andes and the central African highlands (Ruwenzori, Mt.
Kenya).
The austral floras of each of these vegetation zones have been living
under climates of high equability more continuously than their counter-
parts in the north. Thus, they have been removed from the vicissitudes
of major climatic change, and from the competition of taxa better-adapted
to the harsher conditions in the north where equability was progressively
lowered during the Cenozoic. As a result, the austral regions have not
only preserved many "primitive" and "ancient" taxa, but also numerous
bizarre adaptive types in divergent environments-including the desert-
that are highly successful today even though they are very ancient.
Clearly, to interpret the history of the unique taxa that now characterize
the austral deserts, the history of similar taxa in bordering vegetation
zones must also be considered, for all of them have diverged from
basically tropical alliances. Although they have developed in response
to different (tropical to alpine) dry climates of high equability, they are

generally similar in terms of their antiquity as unique adaptive types.
Attention is directed to the fact that they regularly occur in open
environments, that they inhabit areas which have a period of drought
or are physiologically dry (e.g. paramo), and that they find optimum
development in climates of moderate to high equability. These are
essentially the conditions postulated above for early angiosperm evolution.
The savannas and tropical deciduous forests of austral areas (e.g. Mada-
gascar, Brazil, Mozambique) contain so many unique and bizarre adap-
tive types that they impart to those regions the aspect of another, more
ancient plant world. The impression is that these plants may be only
slightly modified survivors from an Early Cretaceous-or earlier-world,
a suggestion consistent with the presence of palms (another bizarre
adaptive type) in the Jurassic. In the modern world of "normal' angio-
sperms, they find an analogy with Welwistchia and Ephedra among the
gymnosperms, or with the persistence of armadillos, sea-cows, and bats
(all bizarre in type) in the world of "normal"mammals.
That an ancient flora adapted to seasonally dry climate had opportuni-
ties for migration across the tropics during Early Cretaceous and earlier
times now seems highly probable from geologic and paleontologic evidence
(see above), and may also be inferred from the distribution of taxa that
link the tropical areas today. It is recalled that the tribe Agaveae is com-
posed of the American genera Agave, Furcraea, and Beschorneria and
the Australian genus Doryanthes-all found in dry areas marginal to or
within the tropics. The baobab (Adansonia) links the dry areas of
Madagascar and east Africa with northwestern Australia, and it has an
ecologic counterpart in Charisla of the Argentine chaco. The cactoid
euphorbias of south and central Africa provide a link with Madagascar-
India, and palms are found commonly in all savanna and dry deciduous
tropical scrub regions, ranging also into desert border sites. The
Byblidaceae, composed of only 2 genera, link western and northeastern
Australia (Byblis) with the Cape region of South Africa (Roridula).
In the small family Tecophilaeaceae, which appears to provide a link
between Liliaceae and Iridaceae (Hutchinson, 1959, p. 613), there
are related genera linking California (Odentostomum) with Chile
(Conanthera), and Chile (Zephyra) and South Africa (Cyanella) with
the more primitive genera (Cyanastrum, Walleria) in tropical Africa.
There also are taxa common to the arid climates of the Americas,
South Africa, the Mediterranean-Near East region, and-occasionally-
Australia (see Engler, 1914). For instance, similar species of Menodora
are in South Africa, southwestern North America, and South America;
related species of Hoffmanseggia are in southwestern North America,
South America and South Africa; paired species of Fagonia occur in the
Mediterranean region, southwestern North America, and Chile; related
species of Thamnosma are in southwestern North America, Socotra
I.-Somaliland; and related species of Gossypium and Malvastrum link
the dry parts of all these areas, as well as India. In the Northern Hemi-
sphere, the links between the arid regions of southwestern North America
and the extended Mediterranean region include species of Fagonia,
Hoffmanseggia, Helianthemum, Lavatera, and Lycium. Furthermnore,
plants in the bordering woodland and chaparral vegetation adjacent to
the desert provide additional ties between these areas. Among these is
Arbutus, which is found in the Mediterranean, the Canary Islands, Mexico
and the western United States. Related species of Cupressus and Pinus
(remorata of California, oocarpa of Mexico, halepensis of the Mediter-
ranean) are represented; and Simmundsia of Arizona-southern California-
Baja California is related (distantly) to Buxus (Buxaceae) of the Mediter-
ranean region which also occurs in dry tropical areas (for others, see
Stebbins & Day, 1967).
To explain the links between the dry areas of southwestern North
America and the extended Mediterranean region, Engler (1914, p. 619)
favored a North Atlantic connection. More recently, Stebbins (in Steb-
bins & Major, 1965; Stebbins & Day, 1967) has suggested the links may
have resulted from long-distance dispersal via Beringia during Paleogene
time. He proposes that dry tracts in the lee of ranges in northwestern
North America and adjacent Siberia provided sites suitable for a sweep-
stakes route (sensu Simpson, 1940). This possibility seems to be ruled out
for several reasons. 1) Although Stebbins assumed that high Paleogene
mountains in the far north provided dry rainshadows in their lee, geologic
evidence shows that the cordillera was not elevated there prior to the
Miocene. 2) With sea-surface temperatures in arctic basin and north
Pacific basin considerably higher in the Paleogene than today, ample
moisture would have fallen over the interiors of Alaska and Siberia, and
dry tracts would not have been present even if mountains were there.
3) There is no paleobotanic evidence to show that climate at high latitudes
was sufficiently warm to support ancestors of the disjuncts which pre-
sumably required warm, sunny semiarid climates at least. 4) Taxa moving
via a sweepstakes route across northeast Siberia would encounter the
coldest climate in the Northern Hemisphere, a climate sufficiently cool for
summer-green deciduous hardwood-conifer forests, and therefore un-
suited for plants of warm, dry climates. 5) The migrants would have
to adapt to a very different photoperiod in the far north immediately
upon germination-which seems unlikely judging from present disjunct
patterns that regularly occur between generally similar latitudes (or
photoperiods), as Chile-California, Argentina-Texas, or British C;olumbia-
Fuegia.
An alternate explanation is that the unique taxa common to the dry
tropical and subtropical regions of the Americas and Eurafrica, as well
as the present links between the desert and desert-border areas, may be
the remnants of a dry flora that inhabited Gondwanaland prior to its
breakup by ocean-floor spreading near the close of medial Cretaceous
time. On this basis, the distributions shown by certain woody plants
commonto three or four desert and desert-borderareasnoted above may,

be relict and ancient rather than comparativelyrecent. This possibility
is consistentwith several lines of evidence providedby the fossil record.
In the first place, angiospermswere already adapted to dry climate
by the Early Cretaceous(Neocomian to Aptian), as shown by the small,
thick, occasionallyrevolute leaves in the early Cretaceousfloras of Cali-
forniaand Kazakhstan.Furthermore,as noted above, palmswere already
living in areasof seasonaldroughtduring the middle Jurassic. Inasmuch
as there are fewer ties today between floras of dry than of humid regions
on opposite sides of the Atlantic,as well as north-southin the Americas,
one could argue that the connections were less effective for plants of
dry than of the more humid regions. Patches of dry climate would thus
serve as a filter bridge for some and as a sweepstakesroute for others
(sensu Simpson,1940), depending on the degrees of drought, extremes
of temperature,and modes of dispersal, physiology, self-compatibility,
and other factors that may have determinedtheir establishment. While
such an analysismay be essentiallycorrect,it is nonethelessevident that
genera now in semiarid austral areas lived formerly to the north-
demonstratingthat transtropicconnections were more frequent in the
past and that interchangehad already taken place prior to Eocene time.
The early Tertiaryflorasof Colorado-Utah-Wyoming have taxa whose
nearestrelativesnow inhabit the warm, subhumidchaco of northwestern
Argentina and occur elsewhere in subhumid parts of South America.
The transitionalEo-Oligocene Florissant flora (MacCinitie, 1953) has
Athyana, Lomatia, and Prosopis, and the middle Eocene Green River
flora (MacGinitie, 1969) has Allophyllus,Athyana,Astronium,Lomatia
and possiblySchinopsis. Fossil woods from the Green Riverflora (Kruse,
1954) resemble Schinws, which lives in South America today, and
Aspidios'perma, which is centered in Brazil. Among several other genera
represented,the wood of Forchammeria(Capparidaceae)is noteworthy,
for this unique monotypeinhabitsthe Sonorandesert as well as semiarid
subtropicalto tropical areas (thorn scrub, savanna) at lower latitudes,
including South America. Furthermore,both the Florissant and Green
River floras have numeroussmall-leavedplants similar to those now in
the dry tropics and subtropics;they occur also in the Eocene of the
southeasternUnited States (Berry, 1916, 1930) and they are in the South
American Tertiary (Berry, 1938). Inasmuch as the Eocene species
resemble plants now of warm, seasonallydry regions in both Americas,
earlier (pre-Eocene) connectionsby patches of dry savannaclimate seem
indicated.
Available evidence lends support to the inference that the unique
taxa of the dry regions may be relicts of ancient dry climates (Engler,
1914; Johnston, 1940; Axelrod, 1952; Rzedowski, 1962), and it seems
probablethat they are Early Cretaceousor of greaterage (Axelrod,1960,
p. 236). Althoughsome of the unique alliances are confined now to one
continental area (e.g. cactoid euphorbs to Africa-India), this need not
AXELROD: EARLY ANGIOSPERM HSTORY 311
mean that their absence from another region (e.g. South America) in-
dicates that they evolved after the Cretaceous link was severed. They
may only have had a restricted distribution, one that was not favorably
situated for migration during the Cretaceous. Alternatively, they may
have occurred more widely in both hemispheres and then became extinct
in one region, as shown by the early Tertiary occurrence of numerous
plants in the New World that are now only in the Old (e.g. Nipa,
Cinnamomum, Aponogeton) or vice versa (Sabal, Regnellidium,
Cyclanthaceae). In this connection, the distribution of Rhipsalis is sug-
gestive (Camp, 1948). It is a common epiphytic cactus of the American
tropics, but also occurs in the montane forests of West Africa, Mada-
gascar, the Mascerene Islands, and Ceylon as the sole representative in
the Old World of the large family Cactaceae (84 genera, nearly 2,000
species) that is otherwise solely American. Rhipsalis may represent a
surviving member in the Old World of a Cretaceous flora that formerly
linked tropical America-Africa-Madagascar-India, a link shown by genera
of other families.
As the Atlantic and Indian Ocean basins widened during the later
Cretaceous, and as seaways lapped far onto (or across) the continents,
precipitation increased over regions previously dry. Semiarid to desert
taxa were then confined to local dry intenmontane valleys and scattered
rocky dry slopes in the southwestern parts of the continents chiefly. At
the same time, diverse taxa that had been adapted to dry climate probably
became extinct as dry areas contracted and locally disappeared. This
would leave the survivors (e.g. Colletia, Forchammeria, Larrea, Adan-
sonia, Fagcmia, Koeberlinia, Simmondsia, etc.) as isolates in their re-
spective families, with no close relatives. Then, as dry climates expanded
following the early Eocene, the survivors, having undergone no significant
change from their Cretaceous relatives, could then spread in southwestern
North America, in the extended Mediterranean region, and also in the
expanding warm, dry areas of South America, South Africa, and
Australia. These ancient relicts now became associated with autochtho-
nous subhumid to semiarid floras that were evolving in response to in-
creasing aridity.
When we realize that the semiarid regions of Laurasia probably were
at least discontinuously connected with Gondwanaland during the Early
Cretaceous, and that the floras of tropical rainforest, savanna, and thorn
scrub vegetation had greater continuity in the past, the problem of the
desert interrelations becomes clearer. Relatively free and isolated from
competition by northern shrubs, and surviving under a more equable
climate, some bizarre ancient taxa have persisted in larger numbers in
austral lands, not only in the semidesert to desert areas, but also in the
dry tropical forest, savanna and thorn scrub vegetation adjacent to them.
But in North America and the Eurasian region, they were subjected to
more extreme climates and were unable to compete with the new
xerophytes that originated there and have largely supplanted them. In
this connection, it is recalled that a number of genera in the Late

Cretaceous and Paleogene floras of the western United States occur now
in South America. Among these are Araucaria, Athyana, Astronium,
Allophyllus, Aspidiosperma,Lamanonia, Lomatia, and Schinus, all of
which inhabit regions with seasonally dry climates. Most of them had
disappeared from the western United States by the close of Paleogene
time, though some plants, notably Stipa, sect. Piptochaetium, persisted
into the mid-Pliocene on the High Plains (Elias, 1942; Stebbins, 1947).
On this basis, it seems probable that more numerous taxa of dry regions,
and notably those of bizarre adaptive type, were represented earlier in
the north and were eliminated there following Paleogene time as
equability was progressively lowered.
Thus, many of the unique living shrubs in the north that now have
austral affinities may be remnants of an old semiarid to arid Gondwana
flora. They appear to have migrated north during the Early Cretaceous
or earlier, and were no doubt accompanied by others of similar adaptive
type which have since been eliminated. During the milder, moister
regional climates of the late Cretaceous and early Tertiary, semiarid
floras probably survived in local dry sites provided by deep, inter-range
valleys, probably not unlike the cactus savanna in the Rio Balsas valley
of Guerrero, Mexico, where it is surrounded by deciduous tropical forest.
In such sites they were associated with ancient, autochthonous taxa of
southwestern North America, notably Idria, Yucca, Pachycereus, Pachy-
cormus, Washingtonia and others that may have evolved in the region
during pre-Cenomanian time. During the post-early Eocene trend toward
increasing aridity and decreasing equability, the bizarre adaptive types
of austral alliance gradually became subordinate to the new, hardier
taxa that developed locally from plants on the drier borders of woodland,
chaparral, deciduous tropical forest, thorn forest and savanna vegetation.
By the close of Paleogene time, when equability was progressively low-
ered, some taxa of unique adaptive type probably became extinct in North
America. Their descendants now survive in the drier equable parts of
South America, a pattern shown by other plants in the fossil record.
CONCLUSIONS
Angiosperms were definitely contributing to the fossil record as far
back as the middle Jurassic, if not in the late Triassic, during a span of at
least 50 to 80 million years prior to their assumption of dominance. To
judge from their adaptive and evolutionary relations, and from paleo-
climatic and stratigraphic evidence, they probably inhabited seasonally
dry, open country in equable tropical uplands during pre-Cretaceous
time. Angiosperms were confined to upland areas early in their history
because the lowlands at lower latitudes were characterized by hotter,
drier and less equable climate-conditions to which they adapted later.
They first entered the record at middle latitudes, where the warm
temperate zone of the montane tropics descends to sea level. Their
gradual appearance during the Early Cretaceous coincides with a climatic
trend toward increased equability which resulted from the increasing
ratio of sea- to land-surface. This was produced by ocean-floor spreading
which opened up the tropical Atlantic Ocean and widened the Indian
Ocean, and by advancing seas on all continents which reached maximum
extent at the close of the Albian and in the Cenomanian-as angiosperms
surged to dominance, and in great diversity.
The idea that taxa now common to the Old and the New World
tropical regions migrated from one area to the other via the North Atlantic
or Beringia during Paleogene times finds no support from paleobotanic or
paleoclimatic evidence, and is also inconsistent with their adaptive and
evolutionary relations. The numerous taxa (families, sections, genera)
that now link tropical regions probably had attained their distribution
by the middle Cretaceous, prior to the final breakup of Gondwanaland
or shortly thereafter, and before the tropical Atlantic had widened ap-
preciably. On this basis, related taxa now in the New and Old World
tropical regions largely evolved in isolation following the opening of
the tropical Atlantic by sea-floor spreading. Ties between Africa and
the Indo-Australasian region probably were not wholly severed by ex-
panding dry climate until Eocene time. This analysis requires the ap-
pearance of numerous angiosperm families before Cenomanian time, and
suggests that bradytely pervades the phylum to a far greater degree than
heretofore suspected.
Floras of warm to tropical deserts are composed of plants that
represent two discordant groups. In the first group are morphologic
isolates that often are bizarre adaptive types with no close living relatives.
Since numerous bizarre adaptive types also contribute to other types of
vegetation marginal to the desert and well out of the desert (e.g. savanna,
puna, dry deciduous forest) their evolution was not shaped by desert
climate. They appear to be derivatives of taxa that developed under
seasonally dry tropical to alpine climates in upland montane regions on
Gondwanaland and Laurasia in Early Cretaceous and still earlier times.
As moister climates spread widely during the Late Cretaceous and
Paleocene, close allies of many of the unique and bizarre taxa probably
became extinct, leaving isolated monotypes persisting in scarcely modi-
fied form in local dry areas. They probably were more numerous in the
drier, warmer parts of Laurasia into Paleogene time, but decreased as
equability was lowered and persist now in greatest diversity in equable
climates with seasonal drought. Alliances now common to the warm dry
regions of the Old World and the New probably had attained this dis-
tribution across Gondwanaland by medial Cretaceous time.
The second group, which accounts for by far the majority of plants
in desert and desert-border regions, was derived from taxa in moister,
bordering regions by gradual adaptation to increasing aridity during the
Late Cretaceous and Cenozoic. Evolution in each of the six major isolated
dry areas largely accounts for the distinctness of the desert floras today.
The evidence suggests there have been three major periods of angio-
sperm evolution in response to dry climate. First, it is hypothesized that
angiospermy evolved in response to seasonal drought, and that the alliance
underwent its basic adaptive radiation in open, equable monsoonal areas
in tropical uplands on Gondwanaland, probably in Permo-Triassic time.
The second period was Trias-Jurassic, and accounts for most of the bizarre
taxa found in warm deserts, and also in vegetation zones of the drier parts
of the continents in lower latitudes, ranging from tropical savannas into
the.alpine puna. The second period has been essentially continuous since
the Cenomanian, first in local dry valleys and on exposed slopes in the
southwestern parts of the continents, then spreading as the trend to aridity
commenced following Paleocene time. Most of these taxa find relation-
ship to those in bordering, moister regions.
Changes in continental position, configuration, size and altitude, as
governed by ocean-floor spreading, plate fragmentation or joining, and
continental rafting ("drift"), have been operating throughout angiosperm
history. With plate fragmentation and separation, lands formerly desert
have become moist tropical (e.g. Brazil-Congo), lands formerly connected
have become isolated (South America, Africa, India, Australia), moun-
tains have been elevated as plates collided (e.g. Alpine belt, American
cordillera, etc.), and climates have been profoundly affected. The im-
portance of the "new tectonics" to angiosperm history is not so much
as a basis for explaining the geographic disjunction of taxa, which has
been the chief concern of most investigators, but rather as a means for
comprehending angiosperm evolution more clearly. New taxa evolve
by developing effective barriers to gene flow, with resulting changes in
frequency of alleles. The degree of variability in terms of novel recom-
binants which may have adaptive or preadaptive significance increases
greatly under isolation, as shown by insular floras. Thus, climatic change
coupled with isolation that accompanied plate fragmentation, ocean-floor
spreading, and continental rafting, has provided a major stimulus for
evolution of floras. The great diversity of taxa in numerous families
in the tropics and subtropics, as well as the evolution of unique floras
of arid to semiarid regions and those of the temperate climates as well,
seems directly related to the breakup of Gondwanaland following the
medial Cretaceous and subsequent evolution in isolation.
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interpreted as resulting from rapid evolution from an unknown ancestor

1-Professor G. Ledyard Stebbins has pointed out (verbal communication, Dec.

It was at this time that Gondwanaland ceased to exist. The continued

upland sites by divergent growth forms with different adaptive features

4 All these investigators refer to a trend toward more tropical conditions,

laurels, Cercidiphyllum, Salix, Populus) is thin-walled and is not readily

5"As far as adaptive radiation can be distinguished from progressive occupation

by sea-floor spreading to reach its present position during the late

CRETACEOUS PLANT GEOGRAPHY

FiC;.4. Distribution of 6 tribes (23 genera) of the family Bombacaceae: 1-Durioneae, 2

Fic. 6. Distribution of 12 tribes (67 genera) of family Tiliaceae. 1-Prockieae, 2-Akpeibeae,

Although ready interchange of taxa between the deserts in the

Pyrenacantha,Randonia,Tamarix,and Zilla;in SouthAfricaare Aeonium,

to different (tropical to alpine) dry climates of high equability, they are

commonto three or four desert and desert-borderareasnoted above may,

this connection, it is recalled that a number of genera in the Late

CAMp,W. H. 1947. Distribution patterns in modem plants and the problems of

REYMENT, R. A. 1969. Ammonite stratigraphy, continental drift and oscillatory

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