Biogeographical Relationships of North American Tertiary Floras

51 Pages·1999·44.4 MB·

by  Steven R Manchester

BIOGEOGRAPHICAL
NORTH
RELATIONSHIPS OF
AMERICAN
TERTIARY
FLORAS'
Europe document
lllo-r
.Hill \sl.l lull
(.1 .1
1
selected conifer and angiospcnii geueia
ll
I
The
ami The
origin of the exianl Mora vegetation of purpose review
this highlight Tertian-
is
<>l l<>
North America has heen the subject of much in- records of selected conifer and angiosperm genera
terest and debate ever since the similarities ma\ be considered soundly and
flori-lie that identified
between North America and Asia were docu- which are significant understanding die hiogeo-
first in
mented. Many of the woody genera comprising graphic affinities of North American Tertiary floras.
North America's present-day forests have excellent present a review of about 90 genera with reliable
I
fossil records that can be traced through the Ter- Tertiary records in North America and other con-
tiary in North America and other continents of ihe tinents. Examination the straligraphic ranges
of of
Northern Hemisphere (Wolfe, 1975; Graham, 1993; different taxa in North America. Europe, and Asia
many
Tiffney, 1985a, b; Mai, 1995). In addition, provides the basis for assessing ph\ logeograpluo
genera that are no longer native to North \uierua patterns and pathways biota dispersal through
of
have well-documented Tertian records. Patterns of the Tertiary. South American and African records
among
geographic disjunction extinct and extant are mentioned when known, but the emphasis on
is
genera provide important clues to the history of affinities within the Northern Hemisphere. For a
North American and former
flora continuity of review of relationships between North and South
lh.
Tertiary forests in the Northern Hemisphere. By America, see Burnham and Graham (1999,
this is-
eomparing
the stratigraphic records of genera
sue).
among
shared two or more continents possible Because of our familiar vantage point of the pres-
is
if
to consider the pathways and timing of plant inter- ent day. neobi botanists alike tend
change through the Tertiary. to regard the modern flora as an endpoint showing
Many
ideas have been published on the paleo- the "true" floristic patterns of extant genera and
botanical origins and development of extant flora species. Thus a genus such as Ginkgo, with a wide
and vegetation of the Northern Hemisphere (e.g.. paleogeographic distribution (Tralau, 1968), mav be
Engler, 1879; Chaney, 1940, 1947; Wolfe, 1975; classified as an "East Asian element." Clearly, such
& &
Raven Latham
Axelrod, 1974; Ricklefs, 1993; "elements" have more to do with extinctions else-
Mai, 1995; Akhmetiev, 1996). In the attempt to be where than they do to the natural geographic affin-
comprehensive
as as possible, investigators have ity or origin of the genus (Wolfe. 1975). Fossils can
sometimes relied uncritically upon genera reported document former
geographic' distribution patterns
many
in the literature. Closer scrutiny reveals er- of both extinct and extant genera and thus provide
roneous generic determinations (Dilcher. 1974). a means of tracking the changing
floristic relation-
ollii
I
W. Wehr, Wen, and V
V.
J.
',',"
.Nln^e,,. Ilongshan War h translation of Chinese Mali
ill.
lil
1 1
m!,
>v R. Stockey, B. ' Iiiln Kvaoek, and editing by
^ I.
erendeen and Can.-
Peter
,,ln. II for
,
1
EAR
This rese ai.li was supported in part h\ grants 9506727 and l\l'')722lll
1
499 Mu
.resents no. in the Contribiations to Paleobiology fron the Florida
i
History. Gainesville, Florida 3261 U.S.A.
iral
1,
Gaud. 472-522.
86: 1999.
Volume Number
2
86,
Biogeographical Relationships
landmasses through Tertian genera Kirchheiincr
diffei is thai of (l'>->7).
mm
Re. siudies have emphasized pholngu ml ml iphasizes the European fossil record but
.-nl
;
molecular phylogenetics as a framework to con- also repol'ls selected reliable occurrences nf genera
Many
log. graphic In-lorv iii^jian la\a ir.- in North America and Asia. of the age as-
- ' |' ; r.
i 1 1 j • . « I I
icuo.l hv Wen. in pi. ss _i signments in Kirchheimers catalog have since been
\ 1! .g
i i II i I [i
ses .lev eloped from phylogenetie and molecular revised, but Kirchheimer was careful to give local-
studies can be tested by observations from the fossil
current age assignments by attention more recent
to
It might be argued that the fossil record is too literature. Helpful, but less comprehensively doc-
ihonm;
f-agmeiiiarv in piov ((!.' a i.m lor-tauditi- ol umented, reviews ta\a icpoiicl the paleobo-
li of in
however,
..graph the alternative of
ic v tanical literature are provided by Taylor (1990),
i I . i ;
i
using only model, -
i il uti >n I * i > I Collinson et al. (1993), and Mai (1995). Useful
potheses
del
to li i - I l.i-i guides to Asian literature on Cenozoic Paleobotany
i | i ' '
leads In an even more incomplete picture. If forced include Tanai (1992a, 1994) and Liu et al. (1996).
we
to rely onlv on extant generic distributions,
Published reviews of the record of
critical fossil
would be Carya had
unlikely to predict that its
particular phylogenetie groups, for example, that of
Europe
greatest diversification Tertiary of
in tin- the magnoliid angiosperms (Friis et ah, 1997), and
(Mai, 1981; Manchester, 1987a) despite absence
its
individual families Retulaceae, Crane. 1989)
(e.g.,
from the extatil European flora, or that Ensete and
or genera Nyssa, Eyde, 1997) are also very
(e.g.,
(both native Africa and Asia
itha to to-
day) would be present in the early Tertiary of North
The decision whether accept generic records
to
America (Manchester, 1994b).
summary was based on whether the organ(s)
for this
American
In this article, review the North Ter-
I
and morphological/anatomical features preserved
and described can be considered diagnostic
truly
tmm
comparison
gmi. w -Iraligm:
-j.,'-!ii ia in ill: la<
genus mod-
the indicated. \- the taxomuiiv of
ol in
Northern Hemisphere.
other parts of the
l make
go
ern plants, often difficult to reliable
is
it
based upon
Rare paleobotanical specimens
showing
and/or
fruits
floi
foliage have revealed "mosaic" plants, in which the
modern
leaves closely resemble those of a genus,
while the or llowers icveal novel characters
limits
indicate belong an extinct genus (Man-
that lliev to
well-known problem paleobotanical
a that
It is
chester, 1989a; Manchester et al., 1998). In the
number
includes a large of generic as-
literature
Vrl
dubious
Mgrunorils \uici
ol '.all lilv. I:i ic; : 'r
unavailable
productive parts arc studies of fertile
much
problen applies of the literature prior to
to
and
material, particularly flowers fruits, prov ide lh.
the 1970s and actually persists to the present day.
best systematic resolution.
ppK
This hecau.se iiivcsiigatnrs <iinVi
is diif.'-tvnl
:i
Reports based on fossil foliage are also impor-
what necessary idenliS
ni -1 h ilards of is to
more commonly
tant, particularly since leaves are
may
nus. also -n
It l>. _
I. I I I I i
many
preserved than depositional environ-
fruits in
names
ohtam
geologists quick, tentative for fos-
to
ments, hut the) require careful seruthn to deter-
may may
that not be well preserved or that not
sils
more
provide characters a secure de-
sufficient for
Leaves some families arc so distinctive as
tions. in
main assigmin
.> par
In nisi in. n! to a
to be reliable in diagnosing a genus, as in Rlalan-
us iiiscnssiiig feu
gci lied
is |iis! l>\ n
!
Herb arc
>etween the fossil species and the mod- a. < i. in. I < -1 Iv lh. i<
convergence and
instances of parallelism in foliage
from
!isliagi.is!i
ei iteii.i r|,.it
ii- lli, i!
me
that cause to be less confident of generic as-
ss hie candidates are not considered,
and
signments. In the Betulaceae, Juglandaceae,
inadvisable accept reports
ion, to
is
it
family using leaf morphology relatively easy, but
is
secure generic determinations icquire In
fruits.
may
good
these families the leaves provide a "best
estimalc" genera are present, but identifi-
ol
lilt
1 1
i '
that the south.
-i
emphasize megafossil reports of fruits, flowers, stratigraphic charts. The Tertiary record for South
I
and leaves, because .mi more confident the Vmeriea and \lina rb known relative
\silli is still
I
|
aUn
and
e\aliiiiln>ii ..I lli.se ..rga ib. \\ |><>ll«'n
I
pro\ide biogeogiaphicallv uiipni laul records, direct exchange of plant species between North and
lull
South America (luring the early to Lite Tertian re-
pollen the de< leased systematic resolution mains weak. For more detailed accounts of rela-
<>l is
i I
aneed
h\ the relatively -I at igraphic resolution tionships inferred between the fossil record of
itz.ii
1 1
(wind-dispersed (Milieu ma\ he lound throughout a North \meriea and the living genera of Africa and
sedimentary sequence, whereas megafossils are South America, see Raven and Axelrod
(1974).
Bumham
eonliiied speeihe laeies). lor eritieal systematic Taylor and and Graham
|,. (1990), (1999).
The
million-year chronology epochs
of Tertiary
refer to Midler (1981). follows Berggren et al. (1995). The Brandon Lignite
In addition, before accepting records for inclu- flora of Vermont significant as one of the few
is
sion summary, was necessary
in this it thai I li«\ I" in ii in I i i il I is ii the north-
documented
fully in the literature with convincing eastern niled Slates. nlortunately. there are no
I
I
photographs.)] had
that the opportunity to observe associated datable rocks or annual fossils to pro-
I
means
ide an independent determining the age.
v ,,l
am
to stu.K eritieal specimens. indebted en though manv voars considered be
to \ lor likely to
I
iatois ol the following collections: niversitv of Oligoeene, the current conscn-u- is that mav be
I il
California, Berkeley (specimen numbers cited have Miocene lW4a;
early (Tiffnev. Traverse, 1994).
I
UCMP);
Moscow;
the prefix University of Idaho, adopt this age the ussioiis involving Bran-
foi
.lis.
Museum Mn
D.nxei of Natural History; IVaho.h don fossils. There are also problems with the pre-
seum
of Natural History, Yale University (YPM); cise age of many floras in eastern \sia. have usu-
I
Museum,
Hinted States National Washington, D.C. ally accepted the assignments given by the most
SNM): Museum
Burke of Natural History and eases there continue
(I >< !>i - -i ui« to
< i 'i' 1 I i i
(UWBM);
Culture, Seattle Department Biolog,. be disagreements among and
al different investigators
..I
Sciences, University of Alberta; Natural llis|or\ mav be expected that the ages of manv -lies will
il
Museum, London (BM-V); Museum.
National continue to be revised in the future.
Senekenhcg Museum
Plague;
of Natural History. have attempted to be conservative in the po-
I
Frankfurt |SM): Botanical Department. Hungarian
sitioning of stratigraphic i.uigcs. hu- the ranges
1
Museum,
Natural History Budapest; Geological presented here
In- are often shorter (begin than
later!
stitute and Geological Museum, Moscow: kniuaiov those indicated by other authors, Mai
e.g., (1995).
Botanical Institute, St. Petersburg; National Sci- lot example, is possible to find reports of En
it
ence Museum,
Tokyo; Nanjing Institute of Geology gelhardia based on pollen from the Paleocene.
and Academia
Paleontology, Siniea, Nanjing;
Insti-
tute of Botany, \cadeinia Siniea. Beijing. Speci- occur until the en. .and
<>,
I
mens
Ih
cited with the preli\ are from the Florida len (usually calf house) known
is
1
.
Museum
of Natural History, Cainesville. Extant have been produced genera
to bv othei of Juglan-
eom|)arative material was studied herbaria daceae
at in- in the Tertiary (Manchester, 1989b). Like-
MO,
cluding A, FLAS, and PE.
wise, there are reports nf ,1/hh.v from the Crela
ceous, but without convincing
infruclesceiices.
may
the paper be found Mabberley The Because scope
in (1997). the ol this treatment limited to
i-
known
genera
the Tertian and/or Becenl
('torn flora
hav
are specified at their first mention in the text. of North America. omitted some interesting
I
Stratigraphic charts presented in tin- paper aic examples of taxa shared only between Europe and
simplistic in thai the\ condense diverse records Asia, many of which are reviewed bv Mai
'><>.">).
( I
from each
different part- of northern continent
into
column. However,
single enables quick
a this a
vi-
summary some
sual that provides constraints on the
liming ol intercontinental exchange events for each The following section highlights records
that
I
taxon. More detail- foi ea. h genu- are provided in consider useful in evaluating the biogeographic
af-
the family narratives of the .Mowing section, lead- finities of North American Tertiary These
floras.
I.
more
ing to the informative literature for the genera paragraphs cite the references upon which the
Pliocene found
ranges I'urope after the Krlclrrriu. to
-tra .die in
:
21-25
(Fig. 19) and in Figures are based. This sec- day only in the broad-leaved evergreen forests of
known
arrange China and Taiwan, based on distinctive
tion is
is
I
Eocene
the (leadings iiih and \ngiosperms, with seeds from the of the CUiilchena flora of
(,-, •.in.
.-.(>«
Columbia Mathewes, unpublished
angiosperms mi h treated un- British (R. data),
ol last,
.1 -
i i
der die (leading lneertae Sedis. Ciiramon patterns and Oligocene of Oregon (Me\ei Mancheslei.
e>
are reviewed the subsequent section. 1997).
in
Taxaceae. Amentotaxus, with four extant spe-
cies China and southeast \sia. has an excellent
in
record North Vineiica and Europe.
fossil in It is
U|M-ess;Hi-ae. Dislillclive leaves of the ex- iva.lil\ recognized b\ its broad needle-like leaves
<
» Hi < and .ismici- with a pair of prominent stomatal bands and dis-
in ,. \1< .\
i 1 ,' I i i i
ated cones occur in the Paleocene of Wyoming, (Ferguson et al., 1978).
f
&
Saskatchewan, Alberta (Mclver Basinger. extends from the ppet Cretaceous
«)<•();
I
I
Mclver, 1992) and in the Paleocene of Altai, Xinj- (Santonian) to Miocene of North America and from
ing. China (Quo et al., 1984, as Ditaxocladus Guo), the Paleocene to ppcr Miocene ol Kurope (Per
I
indicating trausberingial distribution. guson 1978; Jahniehen. 1990). Other genera
a et al.,
and and
Tetraclinis, with one species living in the w c-t.au of the Taxaceae, including 7h.vn.s- Torreya,
Mediterranean today, has a g Tertian record in at least one extinct genus, arc well represented in
I
Europe (Kvaeek, 1989; Mai, 1995). In addition, one the Tertiary of western North America (Manchester,
&
species known tmir goc-:i |o \1io 1994b; Meyer Manchester, 1997) and central
is I'M <\ii l\ < >l
cue western North America based upon cones, Europe (Kvaeek, 1982), but their history best
is
of
.
&
As Mesozoic
seeds, and foliage (Meyer Manchester, 1997). explored through attention to floras.
has not been observed in the east Asian Tertian. Taxodiaceae. This family includes several gen-
it
appears likely that this genu- traversed the North eta with relietual extant distribution. Metasequoia
it
Eocene. has become famous as an example genus once
Atlantic in the late ol a
Hemisphere now
*.iiik»oac«'ae. The record of Ginkgo in the widespread in the Northern thai is
Northern Hemisphere extends back to the Jurassic. native only to China. do not see the need to review
I
biogeographie
but record the Tertiary of
its in is
&
Meyer
interest, documenting the relatively late confine- (most recently in Manchester, 1997).
ment Asia. The Tertian records in North Amer- Cunninghamia grows today in mixed mesophytic
to
ica extend from the Paleocene of the Rocky Moun- and broad-leaved evergreen forests of China and
tains (Brown, 1962) to the Miocene of Oregon Taiwan. Its fossil record includes cones, seeds, and
(Chancy. 1920). A worldwide review of the distri- foliage from the Eocene to Miocene of western
bution genus through space and time was North America (reviewed in Meyer Manchester.
ol this t\
Europe and
presented by Tralau (1968). 1997), the Tertiary of (Mai, 1995), the
m
Pinaceae. Various geneia ol iho I'm ...a. Eocene to Miocene of Japan (Matsuo, 1967; Hori-
well represented in the Cretaceous and Tertiary of uchi, 1996).
the Northern Hemisphere, including, for example, Glyptostrobus is native today only in southeastern
&
&
id'
Abies (Schorn Wehr, 1986), Keteleeria (Meyer Tina. but. h '
L
(
1
&
Manchester, Larix (LePage Basinger, was widespread the Tertian. extends from the
1997), in
It
1991; Schom. 1994). Picea (Crabtree, 1983), Pinus Paleocene (Boulter ex Kvaeek. 1989) to the Plio-
(Miller K Mal.nkv. 1986). and Pseudolarix (LePage cene in Europe (Mai, 1995: Martinetto, 1998).
& cones
Basinger, 1993). Most of these genera are wide- </n europaeus twigs with attached
spread in the Northern Hemisphere today, but Ke- occur in the Eocene to Pliocene of Japan Taiuu.
(
',,.'. re limited lo eastern \sia 1961; Malsumotoet al.. 1997b). In North \„,erica.
/, ., iid ! ',
.
in their ru tiis « m! I!asmg< Glyptostrobus is well represented in the Paleocene
i I. I I '
provided a comprehensive analysis of the Eocene of the books Mountain region (brown.
(1991) to
phytogeographic history of Pseudolarix in the 1962; Hoffman, 1996), and in the Miocene of Ida-
Chaney
Northern Hemisphere based on distinctive ho, Oregon, and Washington (Brown, 1936;
its
&
seeds, ((in.s. and foliage, with earliest records in Axelrod, 1959; Fields, 1996).
Cretaceous Asia and North America followed Sequoia and Taxodium, although restricted in
the of
by Oligocene establishment Europe. They con- then modern distribution to Pacific Coastal North
in
cluded that the genus became extinct in North America and eastern North \nieiK a. respectively.
Hemisphere
America by the middle to late early Miocene, and were widespread in the Northern in the
.
1995: pi. 9, 2l to the Middle Miocene of Swit-
fig.
zerland.
Araliaeeae. western and
Torirellia of eastern
\sia. sometimes placed own laimb. was
in its eoii-
some
sidered by taxonomists to be allied to the Cor-
Aetinidiaeeae. Artimdia
is distributed in In- riaceae. but rec.nl molecular work indicates po-
.,
and cistern was
(lnin,llr-i.i \si;l lodav. lull present sition within the Araliaeeae (Plunkett et 1996).
al.,
North America and Europe
in during the IV-i tiar Torirellia has distinctive fruits (Fig. 1A, B) with
\
known
In North \nicrir.i. mi seeds from three single-seeded
i- lia><-il locules: a central locule with a
i!
mm
Eocene
formation Otegoti (Manclms-
tile (II. o| terminal germination valve and two enlarged, blad-
ter. 1994b). In Europe, fossil seeds occur from the der-like lateral chambers that are infertile. The en-
Eocene
to the Pliocene (Tralau, 1963; Friis. 1985; docaip tissue composed isodiametric
is of sclei'eids.
Martinetto, 1998). These same characters occur Eocene
in fruits (Fig.
Anaeardiaceae.
Pentoperculum
Maneliester
ribed as the
fossil
dm
an extinct genus of belonging Spon-
is fruits to (1994b). There-
dieae shared between die middle Eocene of Oregon fore, now offer the new combination. Tortrrllia
I
(Manchester, 1994b) and the lower Eocene of Eng- honrsii (Manchester) Manchester comb.
nov. (see
&
land (Reid Chandler, 1933, as Draronton,rlon\. \ppendix Tmiii'llm honrsii occurs the middle
in
).
I
The
calyces called Astronium truncatum by Mac- Eocene
of the Clarno Formation, Oregon 1C-
(Fig.
Ginitie (1953) from the Eocene and Oligocene of Roslyn Formation, Washington and
F), (Fig. 1G),
the western lilted Static are \ei\ different ve- Messel, Germain
in (Collinson. 1988:
I pi. 1, fig. 11).
and
nation fruit iimrphologv from the extant genus Fruits are also present the lower Miocene Ob-
in
and no
are longer believed represent \nacardi-
to cidorl localih of Austria (Meller, 1996, and
pers.
&
aceae (Manchester Wang, The comm.
1998). leaves that 1998). These occurrences indicate that 7b-
MacOinilie placed in the -anie species ma\ in fact iitrilio was shared between Europe and North
represent Anacardiaceae,
but they are at least as America during the Eo< cue. although un-
is still
it
Rhus
similar to as they are to Astronium. Rhus is known from the Asian Tertian. The timing of
its
known
from anatomically preserved from
fruits the
Eocene Oregon
of (Manchester, 1994b).
Apocynaeeae.
Elongate seeds with a terminal Catalpa
lives today eastern
in
Asia, eastern North America, and the Canbli.'an
length of the seed body are found in many extant legion. Small biwinged seeds Calalpa have
u beeti
f
genera Apocvnaceae,
of e.g.. Anoilcndron, Clegh- recognized from EaiU Oligocene Oregon
the of
&
ornia, Echites, Forsteronia. Holarrhena, Kibatalia, (Meyer Manchester, 1997). Similarly small Ca-
Odontadenia. Pottsia. Prestonia, Strophantus, and talpa seeds (Catalpa mirrosperma Saportai cur
o, in
\poc\naceous
Wrighlia. seeds can be recognized the Oligocene France and Germain
late of (Saporla.
with ease, but die distinction of genera based onl\ 1889; Weyland, 1937), suggesting a possible North
on seed characters is a difficult challenge. Thus, antii inkage during oi pri.n to the Migocenc
\
i (
I
(=
the fossil generic nam.' Krhitoniuin I'nger 1850 Berberidaceae. Mahonia
distributed
is in
=
Apocynospcrmum X Chan
C\psrlitrs leer lb-id \sia, Malcsia. and North and Central America
to-
I
dler. ( >2<)) is applied to los^l remains. In the North da\. Mihough soiu. iimes subsumed within Herberts
I
m
\rii.iie.in Tertian, such seeds occur the middle iWhittcumre. Mahonia
1997), is easily distin-
Eocene
tireen Ki\ei formation (Apocynospermum compound
guished by genus
foliage. This
its
is
Brown
coloradensis in MacGinitie, 1969: pi. 18, fig. readib recognized b\ the distinctive an '.lure of
Int.
the Clarno shales Oregon (Manchester, un- imparipinnately compound and
1). >f its leaves spiny leaf-
(
published), and the Eocene
late of Florissant, Col- lets, without the need for associated flowers or
orado (Manchester, unpublished). They are not and thus
fruits, is a g candidal.' for recognition
I
known
in North America after the Eocene, but in in the fossil record. Mahonia well represented m
is
Europe they extend from the Middle Eocene (M.s- western North an Tertian extending
\m.-rii floras,
sel, Germany: Senckenberg Collection-ME 7624), from Eocene
the to Pleistocene (Schorn, 1966). Al-
&
the Late Eocene of England (Reid Chandler. though not native in Europe today, occurs the
in
it
1926), Early Oligocene of Budapcst-Obuda, Hun- Oligocene
France and
of (Saporla. North Bo-
Iii<>5)
gai\ (Botanical tepartment, Hungarian Natural hemia (Buzek Neogene Hun-
et al., 1990), in the of
I
Museum BP
lliston Collection, and
63.1039), the gary (Andreanszky, and Miocene
1959),
in late
tin-
Ccske
slredohoff Mountains (Kvacek Walthcr. Abkhasia Mahonia
of (see Takhtajan, 1974).
Ox hil-
—
MO
same showing
Yunnan, China, 52556: A Henri 11907. B. Transverse section of tin- fruit.
tilinclolia
I
(Manchester! comb.
chambers; luo large and cmptv. one smaller ami containing a seed. C-F. Toricellia bonesii
—
Clnmo \ul Beds. Oregon Iransvers. section showing sinall central chamber and larg< late
In
I'i F..—..-.-I1 ( —
UF 9578. D. Delail ihc p specimen in enlarged In show isodiametric sclereids. K. Iransvei
>
i>l -^ <>l ( ..
1 1 1 1 1 1
-
seelioned s—pecimen Intnl.. e .iniiiig a seed, and two larger lateral chambers filled with sedin
In. v. -,; -.i.!,.ll
i ,
UF stripped awav showing two symmetri,
9577. Dorsal surface of a silicified fruit with the pericar]. partially
F.
placed chambers, and the median chamber with a small facet corresponding to the germination valve, holot
lateral
Sedin
F')2;!!i. -(;.
I
A -alb
ng.-M kva<". k k Mi..,-. n,«
,',,/,-,/ i! I'.!./., 1'r.nii
M.
of Bilina, western Czech Republic, is similar to
impl nold from the Oligocene of western North
\i
&
Middle locale (The coal southwestern
America (Kvacek Bfizek, 1994). field
ill
&
Mahonia Honshu: Huzioka Takahashi. and from
Although most diverse in Asia today, is 1970),
Kamchatka Budanfsev m.sid-
poorly represented in the Asian fossil record. Tauai strata in that 1<)<>7|
|
Mahonia
and Suzuki (1963) described lanveojolia lale first
I i
Eocene Bournemouth lug
from the Miocene of southwestern Hokkaido, corded from the lied- of
bill
land (Chandler, 1963).
Mahonia may he questioned. Basel on Betula, which widespread in the Northern
is
affinity to
Mahonia Hemisphere has commonly been reported
appears todav. in
the available fossil record, that
it
because
spread from North America Europe in the early the fossil record on the basis of leaves, but
to
between Betula and
Tertiary and to Asia in the late Tertiary. Its arrival of overlap in venation pall, ins
such remain
Asia may have been either from Europe or North other genera of the Betulaceae, reports
in
speculative unless accompanied by the diagnostic
America.
Betulaceae. The birch family has an excel- trilobed infructesoeiiee brails. North \nicno.i
Iii
Hemisphere Betula documented by such bracts
lent fossil record in the Northern the earliest is
&
Wehr Eocene
(Crane, 1989). Alnus, which distributed today B. leopoldae Wolfe from the middle
is
&
around the Northern b mispl ere d -oub lb< of One Mile Creek, British Columbia (Crane
ai
'.<
I
Subsequently, another species with
Andes, oufii ied on of inirael Stockey, 1987).
is is
'I
( 1 i
V
Wy- / docu-
and associated leaves in the early Eocene in -wherry, is
I I i . • .
oming and in the middle Eocene of Oregon (Crane, mented in the Bridge Creek flora of Oregon (Meyer
o
.
&
common
Manchester, l<>97). In Vs,a Hrtula aheadv helnlacei'iis tms, d. !».!,, col \\\-
is ,u
«
confirmed by bracts in the Paleocene of the Tak- omiug with elongate, cone-like infriiclescences
hobe
flora in Sikhote-Alin (Akhmetiev, pers. coiiini. bearing small wingless mil-. \s this germs has not
( >'»7). In Europe. earliest example confirmed been recovered from regions outside North Ameri-
|||«-
I
by bracts is in tin- early Oligocene of Markvartice, ca, may have been endemic to North America.
it
Czech
Republic (Buzek
et al., 1978). Ostrya, which occurs in North Temperate areas
Carpinus occurs in North Temperate areas today, today, readily recognized by the persistent blad-
is
and is easily identified by the enlarged asymmet- derlike involucre which surrounds the
nutlet.
It is
rical leaflike bract attached to the ribbed nutlet. first observed in the early Oligocene of central Eu-
The genus common &
the Tertiary of Asia be- rope (Engelhardt, 1885: 320; Kvacek Walther,
i- 111
lemma
ginning in the late Eocene (Tana 1972; 1998, pi. 12, figs. 6, 7) and western North America
i,
&
&
Tanai, 1993). In Europe the genus confirmed (Meyer Manchester, 1997) and Miocene
is in the
by two kinds of bracts from the Middle Eocene of China (WGCPC, 1978) and Japan
(Tanai, 1972).
&
Eckfeld flora (Wilde Frankenhauser, 1998) and absent from northwestern North \merica
\
m
common
continues element Oligocene and
as a today, the genus occurred as late as the middle
&
m vo
later floras (Berger, 1953; Roiron Vernet, 1978; Miocene h.mcv \\elrod.
>u
)i c\
(
i !
much
Mai, 1995). It is more rare in the North 1959; Fields, 1996). In Asia, Ostrya fruits occur in
\nieiiean Tertiary. Possibly the oldest record is an the fossil record by the middle Miocene (llu/ioka.
WGCPC,
undescribed species from the middle Kocene of lb 1963; 1978; Tanai, 1961).
Washington
public, (Wehr, 1995: Crane
pi. 3, fig. 3). It I'nl, ic<>( aijiinus. is an extinct genus of fruit
cotilorms Carpinus and
to in the size rtioi ;di. with bract characters similar to Corylus and nutlet
characters similar to Carpinus. was circumboreal
It
ration of the bract, but tin brad differs from mod- during the Paleocene, with occurrences England
in
ern species in the obovate, rather than ovate, out- (Crane, 1981), France- (Crane. 1989). China (Man-
&
line. Curiously, Carpinus appears to be lacking chester Guo, 1996), and North America (Crane
&
&
from the subseqn. ill n n >rd of North Amer- et al., 1990; Sun Stockey, 1992; Manchester
I.
No Chen,
1996). Palaeocarpinus persisted into the Ko-
I
\ineiiea lo,la\ indicate-, eilhei thai the genus per- cene in western North America (Republic
flora:
sisted on this continent after the Eocene without Wehr, 1995: and eastern Asia (Buoy
pi. 3, fig. 4)
leaving a record, or that the Eocene pomdaiiM Sikhote Alin; Akhmetiev & Manchester,
tin
in
perished, followed by a later Tertiary or Quaternary progress).
recoloni/alion born Kurope or \sia. Kea\ es for mci- Caprifoliaceae. Diplodipelta from the Eocene
ly referred to Carpinus from the late Eocene and to Miocene of the western ruled States an ex-
is
I
Oligocene western North America now
are
ol
&
placed in Parararpinus, and are considered the (Manchester Donoghue, 1995). Both genera have
probable candidate
lor the foliage of ihe extinct gale fruits with persistent epigynniis sepals, but
I i
fruit Asterocarpinns (Manchester *\ Crane, 1987; the dispersal units differ. Dispersal units ol );>*'•
I I
&
Meyer
Crane, 1989; Manchester, 1997). consist of a single fruit subtended by three wings
Corylus, which occurs North America, Europe, K
in |.i from the inflorescence
lna<
'
I '
and \sia today, is first confirmed on the basis of n M \e three bract-derived
is - mi- Is,,
i-
i, i |
nuts with multilobed, both foliar and spiny bracts wings, but the involucre subtends pan
a ol fruits
in the Middle Eocene of Republic, Washington rather than just one. The occurrence Dipclta
ol
(Wehr, known
1995). Corylus-like leaves are from based upon fruits in the late Eocene of England
&
many
Paleocene and Kocene around
sites the (Reid Chandler, 1926) indicates that the genus
Northern Hemisphere, but is virtually impossible was not always confined to Asia and apparently
it
determine whether
to thev are the leaves of Corylus, crossed diredb between Europe and
Asia. In ad-
or I'alaro.arptnus. Hence, they are best placed in dition, the morpholnmi similarity between Diplo
&
genus Gardner
the fossil leaf Corylites (Boulter dipelta and Dipclta suggests early Tertian geo-
&
Kvacek, 1989; Kvacek 1994; Manchester
et
al., mtinuity.
Chen, Man-
1996). Silicified nuts of Coryloides Cerei.lipliyllaeeae. Cercidiphyllaceae
are
chester from the Kocene Oregon resemble
of Cor- represented by one living genus with two species
ylus in morphology and anatomy, but differ from native to eastern Asia. The family has an excellent
modern
species b\ being perfect 1\ spherical (Man- fossil record, with leaves and fruits extending back
chester, 1994b). the Cretaceous. Cretaceous and
to late early Tertiary
&
Cranea Manchester Chen
(1998) is an extinct leaves are usually placed in the fossil ». us ' '
>,
i
Volume Number
2
86,
Biogeographical Relationships
ndroi les Berry (Crane, 1984). Fruits of Nyssi- P) ( )(.|. In the \si Teil ar\. llie genus is confinned
i
dium Heer
(s\n h by clustered fruits and associated foliage in the
\
Miocene Kamchatka
of Zaliv Korfa, eastern (Che-
droides leaves, are pod-like follicle- sim- lebaeva, 1978).
Cornaceae. The
ilar to those of Cercidiphyllum, but they are Cornaceae sensu lato (includ-
obliquely striated, and are borne in elongate ra- ing N\ssa< Ma-M id.'a uigiaeeae] have
"•
.
cemes
Hemi-
ralher llian clusters (Crane, 1984). Nyssi- an e\c<'llcnt fossil record in the Northern
common
(i;un! - die l;ilc avlaeeous. P.d. ic.-m sphere dial uielm!. Poll] e\laul an e n, .-„
( i.i
I . : , i i ,
ami Pocenc North \mciic.a. tin- late Cretaceous Alangium distributed today from China to
<if is
Paleocene Palo, and Alangium
lo of \~ ic ei .astern Australia in tropical \frica.
i i i i i
of Europe, Greenland, and Spitsbergen. The south- leaves are striking!) convergent shape and
in llie
ernmost occurrence North America from priman. secondary, and venation
in is tertiary to the
UF
Hope, Arkansas (UF loc. 18607). leaves of some genera in the Malvales. and there
At one Paleocene Can-
locality in the of Alberta,
ada, Nyssidium fruits, Trochodendroides leaves, and lure. However, ihe eridocarps are vers .liagno-he
asso. seeds seedling Here placed |og< \ittngium foss remains
lai.'.l ;j)hI ill liai- u. iical Iriiil in
i, \
( [ 1 1 1 |
er as a single species in as. the Tertiary of North America, Europe, and Asia
.
&
Crane Stock- .vhere, the more (Eyde et al., 1969). In North America species are
I
eonser\ati\e known from Eocene Oregon (Manchester.
pi -. n n n the of
_
t
i I i
to the fruits and leaves continues. 1994b) and Miocene of Vermont (Eyde et 1969).
al.,
Detailed multiple organ investigations on Nyssi- Europe, ranges from the Eocene of England
In
it
r/n/m-related plants from Paleocene localities in (Chandler, 1961) and Germany (Mai, 1970) to the
&
anudn. ug ami, and Pat Pastern Russia uveal Pliocene in Alsace, France (Geissert Gregor,
« I'
that these plmils were verse in phvllolaxv, -hoot 1981). In Asia, fruits are known from the Pliocene
,\
&
m
growth, and u -r. n n No. and Pleistocene of Japan (Miki, 1956; Miki Ko-
.\
fl< (
i i
1985b). from Alberta had oppo- kawa, 1962; Eyde 1969). The
ey, Joffrea speirsii et al., fossil fruits all
- |e ,,:a ;pc)s> |»P also alaa aatci on long u d ml to section Marlea, which occurs in east-
;,| -
., ,;.
I I
short shoots with inflorescences in the leaf axils of ern Asia and Indonesia today (Eyde et 1969).
al.,
UK.riopodial -hoit -liool-. \ v.v>,'. /».://, ,trr!,. n::: liom Infructescences and fruits of the extinct genus
&
the Paleocem long shoots with Amersinia Manchester, Crane Golovneva
nil.ii (1999),
i.i
pseudnw
whorled. or hoik -d. leaves arid lejtnuin n relate.! lo ihe exlnni ihinese ei deiiu, - !)<tt uji.i and
«
The
florescences. plant referred to a- 7 -- mi n ,\iin a\e- for-
'
I.
i i
&
drocarpus amicus (Krassilov, 1976; Crane Stock- merly called lioth North
in
I -
1985b) has alternate phyllotaxy and bears America and Asia. The infructescences are heads
ey,
nee- in the leaf axils of long shoots, \i of tricarpellate fruits with four or five prominent
though is clear that infructescences and fruits liraet sears on ihe peduncle below the head. Per-
it
corre-potidi:i<; lo V i.x<,,v/m»> were widespread in die mineralized fruits have been sectioned lo show iri-
d
Northern Icmisphcre, more work need. ioeular endocarps lacking an
is lo ile -. -d of jiheis,
I . ( , f , i
]
lermiiM the geographic ranges of die different -p< axial iHllldle. u d n nig -u git -.-. .led locales Willi
c.
examph
cies. For nai ie.a h. ipieal valves, fhese fruits, and
I i ,i i 1 1 , i I - I II Ii
Manches-
plants with the Joffrea speirsii type of growth ar- the associated foliage, Beringiaphyllum
&
chitecture occurred outside North America. ter, Crane Golovneva (1999), occur together at
<
Rocky Mountains
,li\!l>i:!:. i,ati\. in Inn n>l lapan. eight localities in the of the U.S.
( ,<i: :. >: r. <
i
t
oguized on the basis of chisl. and Canada, in northeastern China, and in southern
r<
.
nd i— the extant spe- Primorye, Koryak Highland, and western Kam-
io-< of
1.(
i
The
cies in the lower lioth n chatka. Ifnssia. apparent absence of Amersinia
mi
i >l i i< I >i ii
I
&
central Europe (Meyer Manchester, 1997). In from the Tertian Europe, Greenland, and Spitz-
ol
North America continues through the Middle bergen suggests thai ihe genu- dispersed across
it
&
Miocene of Idaho (Smiley Rember, 1985). In Eu- Beringia but did not populate the North \llantic
oi
known
leaves are Turgai region.
< i
from the Early Oligocene Pliocene (Jahnichen Cornus occurs North Temperate regions, and
to et in
\mcnca The
al., 1980; Kovar-Eder et al., 1998). Staminate flow- e\tei d- into Soiill and \lri. a today.
found with the and leaves the Miocene Pah en North America
fruits in ins i- -ii a
i'l's
if
< i i i i
m
of Ii, :n di ,r In in tanl species by the de- both from leaves (e.g., Cornus Inpnhorea Heer.
I i i •
&
velopment of a perianth (Kvacek Konzalova, Hickey, 1977: 144, pi. 47, fig. 1) and fruits (Crane
Garden
Botanical
i
et al., 1990: fig. 31G-I) in the Paleocene of North onomic levels lo be accorded different laxa wilhin
Dakota. Leaves are readily identified to the genus the maslixioi.ls are not agreed upon. Murrell
-I
11
i
because recommended
(sciisii latoi (In- -sinootliK curving aerod- (1993) that Diplopanax be treated as
<>f
roninus -<-c.. ml. tries, lliin. widolv spaced, percur- a section within Mastixia. this proposal were to
II
rent tertiary veins, and entire margin, hut discrim- be accepted, then most the laxa now treated as
ol
genera (Mai, 1993) would need
cue subsumed
rcprodiielive male-rial. Paleo, Citrnn.-, fruits horn lo be within the extant genus Mastixia.
North Dakota, and those described from the Early \\ Inchever approach might be taken in the future.
&
Eocene England
of (Reid Chandler, 1933, as is clear that the fruit morphological diversity of
it
Dunstanea) arc auatomiealK pi-served, showing this complex was much grealei the than
in I'citiarv
resin cavities in the wall thai nidi, ate affinities w
illi
Ohem
the Cornelian group dogwoods. Thus, Langtonia one
ol of the
il,, is
i
a.rnelian iherry group s These hilo.ailar ellipsoid fruits
( (
.
North Atlantic during the a K l'ertiar\. I'm.I.h. conform to the Mastixioideae by having an endo-
< i .
dogwoods composed
of the hig-hracted provided the earp of tortuous single-seeded
is fruit fibers, loc-
1>\
of Cornus clarnensis from the Eocene of Oregon, ales, and elongate germination valves.
|'he fruit dif-
which resembles extant C. florida (Manchester. fers from other modern and loss,] mastixioids b\
1994b). the occurrence ol paired dorsal infolds in each loe-
Nyssa, with a disjunct distribution in eastern llle. giving a \\ -shaped cross section of the looiile
\sia. eastern North \iiieii. a. and Central \mciiea. and seed in contrast to the usual - or V-shape.
I
has an excellent fossil record in the Northern This genus was lust described based on specimens
Hemisphere (Eyde, 1997). Nyssa fruits have woody from the Karl) Eocene of England (Reid & Chan-
composed
stones of fibers and have one to three dler, 1933) and was subsequently recognized in the
loeules with apical-dorsal germination \al\es. middle Eocene of Oregon (Manchester, 1994b) and
Transverse sections of pcrinineralized specimens Paleocene Wyoming Haggard,
of (Tiffnev 1996).
cC
reveal that the endocarps are composed of tortuous Elaeocarpaceae. Sloanea occurs today
in
libers, lack an axial bun. and possess apical dot tropical and subtropical America, Asia, and Aus-
lie.
valves— s\ndromc
germination spun
sal a diagnostic of Fruits of four valve- correspond-
tralia. to five
\ t.v.si/. The genus is well c| lies, a led in he o< cue ing lo those o( extant Sloatira occur in the I'alen-
i i I I
and
later I'ertiaiv ol North \ineriea (Manchester, cetic ol North \iiieri. a. where llicv have gone under
1994b) and Europe (Mai, 1995). In Asia, extends the name Carpalithrs 2A-C; New-
s/>mo.we> (Kig.
it
from the Oligocene to Recent (Eyde, 1997). berry, 1898: 138, pi. 68, figs. 2, and in the
3),
Mdsti.Mu. which occurs in Asia today, is well Paleooeue of Oreenland (Kig. LM>. K: "Cistanra lin-
known
lor its excellent representation In fruit- in ger?' in Heer, 1869: 470, pi. 45, fig. 2). These re-
the European Tertiary, extending from the Eocene
&
(Reid Chandler. 1933) to the Miocene (Kirchhei- into the Kocene of the Kockv Mountain region. Slo
mer, 1957; Mai. 1993). Mastixia endocarps are an- ancaccarpum a genus based on similar spiny
is a
alomicalb miiiiI.ii to \\ssa. but the endocarps have valve from lower Olijiocene ol Hungary (Rtiskv.
1«
1 1
more pronounced
dorsal infolds, resulting loo- 1962).
in
Eucommiaceae.
Eucommia,
native only
to
central China lodav. has di-tm. live samaroid fruits
endocarp. lathei than being confined to the apical (Fig. 3A). Such fruits have an excellent record in
end Mastixia also occurs in the Kocene of Oiegon the Kocene of the western and southeastern United
&
and &
California (Manchester. 1994b; Tiffney Hag- States (Kig. 38: Call Dilcher. 1997), and extend-
gard, 1996). In addition to Mastixia itself, there is ing into the Migocene 01 Miocene of southern Mex-
(
&
complex closeb dated genera mas-
a ol cxlincl ol ico (Fig. 3C; Magall6n-Puebla Cevallos-Ferriz,
i.
Europe The
tixioids in the Tertiary of (Kirchh. imer. 1994a). generic determination of these
fruits
is
1936, 1957; Mai, 1993) distinguished by differenc- unquestionable because of the unique morphology
and venation, and the presence of latex strands ob-
sence of gum/resin servable the fossils u enriespond position
« III || III
|
tending back Cretaceous
to the late (Maastrichlian: to the laticifers of extant fruits (Szafer, 1954; Tra-
&
&
Knobloch The
Mai. 1986). living mastixioid ge- lau, 1963; Call Dilcher, 1997). In Asia, Eucom-
i.u- 'in\ i- a native of \sia today, but has mia is well documented b\ fruits in the Kocene of
fossil octillion, es in North America (Slockev et al.. Yubari, Hokkaido, Japan (Huzioka, 1961), the low-
&
1998) and Europe (Eyde Xiang, 1990). The tax- er Oligocene of Kiin Kerish. Ka/akhsian \klune-
I
Biogeographical Relationships
1991), and the Miocene Kraskino flora of lau, 1963; Mai, 1995). The North American fruits
r,
asan Basin, south Primorye, Russia (Ahlaev et are about half as large as fruits of the extant species
1993). The numerous fruit records in Kurope ami are -liuhtK rn< t a-\ nmielrieal in the place-
.
&
end from the Oligocene to the Pleistocene (Tra- merit of stigma the apex (Call Dilcher,
at fruit
;vw
—
China, A: P. C. Silvestri 7378. B. Eucommia eocenica (Bern) Brown from the Bo\a\ Cla\ Mi^i^ippi. 137.57-
— I'il. I I-
M &
Eucommia
8218. C. I, g n Puebla Cevallos-Ferriz from the Oligocene or Miocene Pie Vara For
tie
ill
=
.nation. I'uehla. Mexieo. I(,\1-I'H 237:5 t,V2. Seal.- bars cm.
]