Systematic Placement of the Platanaceae in the Hamamelidae

8 Pages·1991·6.2 MB·

by  Robert N Schwarzwalder

PLACEMENT
SYSTEMATIC
and
Robert Schwarzwalder^
I\. Jr.^
David
THE PLATANACEAE Dilcher^
OF L.
THE HAMAMELIDAE^
IN
Abstract
A cladistic analysis of the basal Hamaiiielidae based upon 92 characters presented. The results support a close
is
phylogenetic relationship between the subfamily Altingioideae (Hamainelidaceae) and Platanaceae but suggest that
present hypotheses concerning the derivation of the Hamamelidae from the Magnoliidae need revision. Analysis of
the subfamilies of Hamamelidaceae using Platanaceae as an outgroup yields a phylogeny that corresponds to the
five
fossil record far better than does current theory. As a result of this study and recent paleobotanical and phytochemlcal
Hamamelidae
evidence, suggested that are at least as primitive as Magnoliidae.
is
it
common
Platanaceae are a monogeneric family of trees than Hamamelidaceae from a stock
lier
He
commented
with a history that extends to the Albian, of flowering plants. that the flowers
fosbil
&
Lower Cretaceous (Kutuzkina, 1974; Hickey of Platanaceae are the more primitive of the two,
&
Doyle, 1977; Dilcher Eriksen, 1983; Upchurch, but stated that ''neither family can be derived from
1984; Crane 1986; Schwarzwalder, 1986). the other/'
et
al.,
Cronquist (1981) considered Hamamelidales, which
Hama-
include Platanaceae, as an order of subclass
Procedures and Methods
melidae ancestral to other orders within the
all
A Hamamelidae
subclass except the Trochodendrales. According to cladistic analysis of the basal
this view, Hamamelidales evolved from a Magnoliid Eupteleaceae, Platanaceae, and the five subfamilies
—
or Magnoliid-derived taxon on a of evolution of Hamamelidaceae was conducted using Swof-
line
PAUP
that spawned Trochodendrales. Recognition of ford's (Phylogenetic Analysis Using Parsi-
first
Platanaceae as a well established angiosperm family mony) program (Swofford, 984) on Indiana Univ-
1
PAUP
in the Early Cretaceous raises questions concerning ersity's mainframe computer. options were
the current systematic placement of this family. chosen to consider possible trees on the basis
all
A maximum
diverse body of evidence supports a close of parsimony. Character state polarities
systematic relationship between Platanaceae and were established by outgroup analysis using Tetra-
Hamamelidaceae (Tippo, 1938; Jay, 1968; Takh- centraceae and Trochodendraceae as outgroups.
&
tajan, 1969; Hickey Wolfe^ 1975; Cronquist, Ninety-two characters are presented for each
&
1981; Zavada Dilcher, 1986). Wolfe (1973) of the nine taxa in Table 1. Character states are
suggested a relationship between Platanaceae and coded for presence or absence for each of the nine
Eupteleaceae based upon foliar characteristics. taxa surveyed. Presence of a character state in
Morphological similarities between Platanaceae any species of a given family or subfamily resulted
—
and Hamamelidaceae especially floral resem- in that taxon being scored as positive for the trait,
blances between harnamelidaceous subfamily Al- The data used in our analysis were gleaned from
—
tingioideae and Platanaceae strongly suggest the literature or generated by Schwarzwalder
al-
liance. However, there no agreement concerning (1986). The 92 characters include aspects of see-
is
the relative advanced or primitive nature of the ondary chemical compounds, wood anatomy, epi-
two families, and thus the extent of their relation- dermal characteristics, pollen structure, foliar mor-
ship remains unresolved. Tippo (1938) felt that phology, petiole anatomy, seed structure, floral
Platanaceae are ''clearly more advanced than the morphology and pollination adaptation, inflores-
Hamamelidaceae anatomically.'' Cronquist (1981) cence type, and type.
fruit
considered Platanaceae to have been derived ear- In our analysis the five subfamilies of the Hania-
Wc NSF
acknowledge the assistance of Professor Ted Barkley and grants 79-10720 and 85-16657 to D. Dilcher.
'
^ Department of Biology, Indiana University, Blooinington, Indiana 47405, U.S.A. Current address: Engineering/
308 Ann
Transportation Library, Undergraduate Library, University of Micliigan, Arbor, Michigan 48109-1185,
U.S.A.
Departments of Biology and Geology, Indiana University, Bloomington, Indiana 47405, U.S.A. Current address:
'
Museum
Florida of Natural History, University of Florida, Gainesville, Florida 32611, U.S.A.
962-969.
Ann. MissoiiHi Rot. Card. 78: 1991.
&
Volume Number 4 Schwarzwalder Dilcher 963
78.
Systematic Placement
1991 of
Platanaceae
The random
melidaceae, Altingloideae, Disanthoideae, Exbuck- suits. experimental addition of error
HaTiamelidoideae, and Rhodoleioide- data can be considered as extra ho-
landioideae, into this set
ae, were treated as taxonomic equivalents to the moplasy. Well-resolved relationships should be sta-
Platanaceae, Eupteleaceae, Tetracentraceae, and ble despite the addition of such evolutionary noise.
Trochodendraceae. This subdivision was performed Thus, this a test of the stability of proposed
is
The
to insure a meaningful analysis of these taxa. relationships. Since cladistics formulates the evo-
subfamilies of the Hamamelidaceae are phe- lutionary interrelationships of taxa considered
five all
notypically quite distinct. Unlike the other families in an analysis, an error in the data set of one
of the basal Hamamelidae, which display only mi- change in any taxon has ramifications for all taxa.
&
Hama-
nor genetic and phenotypic variation, the Rohlf Sokal's (1981) statistical tables fur-
melidaceae, with 26 genera and more than 100 nished random numbers. Three-digit random num-
species (Cronquist, 1981), are extremely hetero- bers were used to designate character states to be
The number
geneous. If the family analyzed as one taxon, changed. first digit of the specified
is
the large degree of phenotypic variability encom- the taxon (1-9), and the last two digits specified
passed by Hamamelidaceae tends to obscure in- the character (1-92). As all data are in presence-
trafamilial and interfamilial relationships. Numer- absence form rather than having three or more
ous authors have recognized the distinctness of the character states, characters could be reversed with-
hamamelidaceous subfamilies and have recom- out ambiguity. Four tests of the initial cladogram
mended one more them were conducted. In each 41 character
the elevation of or of to (Fig. 1) test
5%
828
famUial status (Blume, 1928; Hayne, 1830; Lind- states, of the total character states (92
1836; Wilson, 1905; Nakai, 1943; Chang, characters for each of nine taxa), were reversed.
ley,
A
each
1959, 1964; Skvortsova, 1960; Willis, 1966; listing of the character states reversed In
Takhtajan, 1969; Melikian, 1971, 1973a, Rao, of the four tests presented in Table 2. Figure
b; is
& 2A-D
Rao Consen-
1974; Bhupal, 1974; Wolfe, 1973; Dahl- illustrates the results of these tests.
were where more than one
gren, 1977; Jha, 1977). sus trees calculated tree
maximum
These subfamilies are considered as was determined to possess parsimony,
five sister
CONTREE
program, which
groups Platanaceae and Eupteleaceae. Tetra- Swofford's uses the
to
centraceae and Trochodendraceae, together con- methods of Adams (1972) and Rohlf (1982), was
sidered by Cronquist (1981) as members of the used in these analyses,
most primitive order within the Hamamelidae, were
used as separate outgroups to establish character
Results and Discussion
state polarities. Establishing character state polar-
using the two families phenotypically closest The cladistic analysis resulted in one tree of a
ities
Hama-
to the Magnoliidae allows us to reconstruct length of 157 steps (Fig. In this cladogram,
1).
Hamamelidaceae
melid evolution in relation to current evolutionary Platanaceae are placed within
thought (Cronquist, 1981). By using Tetracentra- sensu lato as a sister group to subfamily Altin-
ceae and Trochodendraceae as outgroups we test gioideae. Platanaceae and subfamily Altingioideae
the hypothesis of a Magnoliid-derived Hamameli- share 12 derived character states: the presence of
dae. CercidiphvUaceae and Myrothamnaceae are cyanidin (character glandular trichomes (char-
5),
not considered in this analysis, because we feel that acter 17), actinodromous to palinactinodromous
evidence suggests neither family as a hypothetical primary pattern of venation (character 42), the
group Platanaceae. union of intercostal tertiary veins to secondaries
sister to
In order to determine the stability of relation- (character 47), altingioid marginal venation (char-
ships within the cladogram generated from this acter 53), the presence of several vascular traces
study, four analyses were performed. In each in petioles (characters 56 and 57), orthotropous
test
5%
random error was introduced into the ovules (character 60), small seeds (character 64),
test,
PAUP
original data set, and was used to generate hard testas (character 69), minute petals (character
The
a cladogram(s). Using cladistics, one formulates the 74), and nonperigynous flowers (character 83).
most direct covirse of evolution possible to hypoth- position of Eupteleaceae on the cladogram and the
synapomorphy from
esize the evolutionary relationships of a given set large list separating this family
Evolutionary homoplasy, occurs Platanaceae and Hamamelidaceae suggest that
of taxa. noise,
i.e.,
more
in a data set proportional to the extent that mosaic Eupteleaceae and Platanaceae are distantly
evolution occurs in a group of related taxa. Ho- related than indicated by similarities of foliar
is
component any
moplasy an expected of data set features alone.
is
and tends to introduce ambiguity into cladistic re- The cladogram suggests that Platanaceae orig-
"
964
Annals of the
Garden
Missouri Botanical
Table 1. Characters and character states for A. Disanthoideae, B. Exbucklandioideae, C. HainameHdoideae, D.
Altiiigioideae, E. Rhodoleioideae, F. Platanaceae, G. Eupteleaceae, H. Tetracentraceae, Trochodendraceae. Su-
I.
perscripts: 1. Saupe (1981), 2. Seigler (1981), 3. Dahlgren et al. (1981), 4. Jay (1968), 5. Harborne (1967), 6.
&
&
Tang
Gornall et (1979), 7. Metcalfe Chalk (1965), 8. (1943), 9. personal observation, 10. Bailey Nast
al.
& & &
(1945), 11. Zavada Dilcher (1986), 12. Praglowski (1974), 13. Bogle Philbrick (1980), 14. Hickey Wolfe
& &
(1975), 15. Wolfe (1973), 16. Nast Bailey (1946), 17. Endress (1977), 18. Nast Bailey (1945), 19. Boothroyd
@
& =
Rao Harms
(1930), 20. Bogle (1970), 21. (1974), 22. Schwarzwalder Dilcher (1981), 23. (1930). not
=
unknown
applicable; value.
?
A B C D E F G H
I
Cyanogens
tyrosine-derived 4-
1.
-I-
Monoterpenes
2.
iridoid^'^^
-I-
+ + +
EUagic
3. acid'^
+ +
4. Myricetin*-'^
-I-
5. Cyanidin*'
+
6. Flavonols acylated'*
9 9
Leucodelphinidine*
7. ?
+
Leucocyanidine* 9 9 9
8.
t
+ +
Vessels present""
9.
Wood + + + + + + +
10. rays heterocelluli
parenchyma
11. Axial banded'
Wood >
+ + + +
1,600
12. fiber length 4-
/ini
-i-
@ @
13. Tyloses present^
> 800 + + + @
14. Vessel elements ^m"**
-I- -I-
@
Number 20-»
15. ^perforation
l)ars [)late
@
+
16. Perforation plates simple""
I I
Trichomes + +
glandular"-
1
*
+ +
Trichomes
18.
stellate'-'*
Trichomes
19. branched'-*'
+
Trichomes
20.
peltate'
+ + + + +
21. Secretory +
cells"'"
+ +
22. Crystals
solitary'-'"
+ + + + +
23. Crystals
clustered'''"
24. Stomates '" + +
laterocytic'
+
25. Stomates
paracytic*'
-I- -I-
26. Stomates anomocytic + +
.'I
+
27. Stomates
cyclocytic*'
+
28. Ej)idermis
pajtillate*
29. Pollen + + + + + +
tricolpate"-'--'-^ 4-
30. Pollen forale'"--''
-*-
+ + +
31. Pollen prolate'"^'^
+
32. Pollen spherical'"-'^
-f- -I- -i-
+ +
33. Pollen oblate'"-'^*
-I-
M,I2 + + + + + +
34. EiidcxiiH* present
+ +
35. Endexine thick at aperture" '- 4^
36. Endexine not througli aperture 1,12 + +
1
37. Pollen psilate 11,1 2, 1."I + + + + 4-
+
38. Pollen scabrate'"-"
+ +
39. Pollen rugulate 11. 2.1.1
1
+
40. Leaves opposite"-^
+ + + +
41. Primary veins pinnate""
4-
Primary +
42. veins ([>alin)actinodromous'"*
+ +
43. Primary veins acrodromous''
+ +
44. Secondary veins craspedodromous"^ ''
-f-
45. Secondary veins camptodrumous*'" '"^ + + + + +
4-
+ +
40. Tertiary veins strongly [)ercurrent"
4-
+
47. Intercostal tertiaries joining secondaries'
+ + + + + +
48. Tertiary veins forming chevrons" 4-
-i^
+
49. Teeth chloranthoid'"'
4-
-
&
Volume Number 4 Schwarzwalder 965
78, Dilcher
1991 Systematic Placement
of
Platanaceae
Table
Continued.
1,
A B C D E F G H
I
+ +
50. Teeth platanoid'"*
+ +
Secondary
51. decurrent*"*
ve^ins
+
52. Teeth
adniedially oriented**'*
+ +
53. Altingioid marginal venation^
54. Teeth hamanielidoid'*
-i-
+
Nodes
55. unilac jnar"'^
Number + + +
56. petiole traces
-I- -i-
Number > 1"'^ + +
57. petiole traces
+
58. Petiole encloses axillary bud' ^ '"
+ + +
59. Stipules present^*^
-I-
+ +
60. Ovules
anatropous'**'**'"'-**
-I-
Embryo + + + + + +
61. narrow^'
+
Raphe
62. bundle
enclosed'*'^'
Embryo + +
63.
luinute'^'^^'
+ + +
64. Seeds
small'^^'«-'
H-
-I-
+ +
65. Subchalazal area vascularized'"'^-'
+
Tegmen
66. two-layered-'
+ + + +
Tegmen
67. persistent in mature seed-
+ + + +
Raphe
68. bundle ending
at chalaza'"-'
69. Testa hard, conspicuous"'^'
'^'^'^^« + + + + +
70. Carpels fused
+ +
71. Carpel wall cpen'*'«"*^
+ +
72. Flowers
imperfect'*'-^^''*^**
'«'''-" + + + +
73. Flowers perfect'^
74. Petals minute or absent"""'**-''
-i-
+ +
75. Sepals minute or absent 16,18.19.20.22
+ + + +
76. Carpels '"'«• "*-''
sessile
Stamens
77.
valvate'"''-^"'"-'"
>
78. Ovules l'"-i^-i^-2"
-I-
+ + +
79. Anemophilous'^'^'^
-I- -I-
+ + +
80. Entomophilous'^''-'"
+
81.
Epigynous"*-'*''^-^**'"
+ + + + +
82. Hypogynous'^'^'^-2'^2^
-I-
+ +
83.
Perigynous'^'«'^-2*>-2^
+ + +
84. Flowers solitary or in pairs^*^'*'^ '" 20
+ +
'"^'"'^o
85. Flowers
in spikes'^
+ +
cyme cymose 20
86. Inflorescence a or raceme'^- 17, 18. |().
'«"* + +
^«
87. Flowers in heads'"'
+
88. Flowers
in pseudanthia'"'^'^'^'^*''^'
+ +
89. Fruit a follicle or follicetum'" .l7.IB.10.2n.21
+ + +
90. Fruit
capsuhtr^"'^''^'''^*^-^' H-
-i-
+
91. Fruit an
achene^"'-'^'^-^"--'
+
92. Fruit a samara'^'''^-'^--^--'
Members known
inated late in a series of evolutionary events that of the Platanaceae are first from
gave rise to the subfamilies of Hamamelidaceae. fossil leaves and reproductive material from the
The position of Disanthoideae in the cladogram Albian, Lower Cretaceous (Kutuzkina, 1974; Hick-
is
& &
corroborated y Cronquist 968, 1981) and ey Doyle, 1977; Dilcher Eriksen, 983;
t
1 1
(
Takhtajan (1969); however, this is not unexpected Upchurch, 1984; Crane at al., 1986; Schwar-
The
since their placement of Disanthoideae based zwalder, 1986). megafossil records of
first
is
upon the Magnoliidae. However, Altingioideae are leaves from the genus Liquid-
their similarity to
ambar
the positions of Hamamelidoideae, Altingioideae, L. in the Cenoinanian, lowermost Upper
and Platanaceae as the most recently derived taxa Cretaceous, (Newberry, 1898; Chesters et
al.,
of the group do not agree with the record. 1967) and Upper Cretaceous (Brown, 1933; En-
fossil
966 Annals of the
Garden
Missouri Botanical
Table Character state reversals for the tests of
2.
Character
cladistic relationshijKs in the basal Haiiiamelidae.
state reversals are identified, by taxon, for each of the
Numbers
four tests presented in the body of the paper.
given after each taxon correspond to the characters listed
Table
in 1.
MAMA
Random One 2A)
Error Test
(Fig.
89
Disanthoideae: 29. 33, 34, 54, 70,
80
Exbucklandioideae: 30, 38, 60,
ALT 57
Hamamelidoideae: 4, 14, 23, 27, 38,
75
Altingioideae: 68,
86
Rhodoleioideae: 23, 27, 31, 48, 52, 69, 82,
2, 4,
PLAT
85
Platanaceae: 76,
5,
71
Eupteleaceae: 23, 56, 59, 64, 68,
76
Tetracentraceae: 59, 61,
Trochodendraceae: 65
Hamamelidae
Figure Cladogram with
of the basal
1
Two
. Random 2B)
Error Test (Fig.
Tetracentraceae and Trochodendraceae as outgroups.
88
Numbers
Synapomorphies are designated by rectangles. Disanthoideae: 30, 49, 71,
make numbers 34
rectangles reference to character in Exbucklandioideae: 25,
in
=
TETR
above
Table 1. Abbreviations are as follows: Hamamelidoideae: 23, 45, 52, 91
=
TROC = EUPT
Trochodendraceae,
Tetracentraceae, 86
Altingioideae: 15, 18, 20, 45, 51, 54,
5,
= RHOD =
DISA Rhodo-
Eupteleaceae, Disanthoideae,
20
Rhodoleioideae:
HAMA =
EXBU = Hama-
Exbucklandioideae,
leioideae,
78
Platanaceae: 59, 66,
5,
^ PLAT =
ALTI and
melidoideae, Altingioideae, Plat-
89
Eupteleaceae: 16, 28. 43, 44, 46, 84,
anaceae.
73
Tetracentraceae: 18, 32, 42, 67,
Trochodendraceae: 27, 40, 55, 64, 73
1,
Random
Error Test Three 2C)
(Fig.
The
dress Cronqulst, 1981). records of
in first
59
mcgafossils from the Hamamelidoideae are foliar Disanthoideae: 3, 31, 38,
53
Exbucklandioideae:
19,
from the lower Paleoccne (Mathiesen, 1932; 4,
fossils
79
Hamamelidoideae:
18, 22, 34, 61, 77,
1936; Koch, 1963; Wolfe, 1966, 1973),
Rollick,
* Altineioideae: 48, 52, 70, 80, 86, 91
r J r *u J
Trr-hi e earl1 y 1 records of these put^ativel1 y ad-
fossil fi i^ io on ^o ^o co co
t,, j 7-t6a
Rhodoleioideae: 16, 18, 30, 43, 48, 52, 59,
-^
, r f
vanced taxa and, t,he ,lac,k of earl1 y lossii1 represen-
qq
^ r>, c 5trzo
^ Plat^ anaceae: 33,
5,
cladogram suggest
tation of the other taxa in the
68
E^p^^i^-^eeae: 32, 43, 66,
may
that the analysis not be valid.
91
Tetracentraceae: 81,
The discrepancy between these results and the 69
Trochodendraceae: 19, 34, 56, 63,
record prompted us to lest the validity of our
fossil
Random Four 2D)
Error Test (Fig.
Four were performed by
analysis. tests in-
initial
92
Disanthoideae: 75, 77, 84, 87,
7,
5%
troducing error mlo the data set and conducting
50
Exbucklandioideae: 39,
4, 6,
PAUP
analyses on each with the software.
cladislic
78
Hamamelidoideae:
39,
3,
R
Its are presented in Figure 2. 88
Altingioideae: 10, 73,
The between and
relationship Altingioideae Plat- 81
Rhodoleioideae: 19, 58, 60, 65, 68, 71,
anaceae, and the position of Eupteleaceae are pre- Platanaceae: 26, 28, 37, 62
served in the four consensus trees in Figure 2. The Eupteleaceae: 23, 24, 54, 91
81
other subfamilies of Hamamelidaccae, however, are Tetracentraceae: 4, 12, 24, 25, 48, 66, 67,
Trochodendraceae: 55
40,
subject to extensive rearrangement with the ad-
random Thus, phylogenetic
dition of error. the
Hamamelidae
presented
reconstruction of the basal
Figure questionable we assume that there Eriksen, 1983; Upchurch, 1984; Crane et aL,
In 1 is if
5%
homoplasy data 1986; Schwarzwalder, 1986), predates the occur-
the original
at least in set.
is
member
A possible source of error in the hypothetical rence of any other of the Hamamelidae.
&
phylogeny presented Figure the use of the Hickey Wolfe (1975) noted the early occurrence
in
1 is
them
two families of Trochodcndrales as outgroups in of the platanoids and suggested as ''possible
evidence of Platana- early members of the trend toward the hamamelid
the cladistic analysis. Fossil
ceae the Albian, upper Lower Cretaceous (Ku- line.'' If we assume that their early occurrence
in
& &
members
1974; Hickey Doyle, 1977; Dilcher and morphological similarity to other of
tuzkina,
.
&
Volume Number 4 Schwarzwalder Dilcher 967
78.
1991 Systematic Placement of
Platanaceae
A)
1
3
DISANTHOIDEAE
6
EXBUCKLANDIOIDEAE
10
ALTINGIOIDEAE
14
HAMAMEUDOIDEAE
15
c
ALTINGIOIDEAE
16
5
PUVTANACEAE
19 17 HAMAMEUDOIDEAE
21 42
RHODOLEIOIDEAE
26
47
39
53
EUPTELEACEAE
50
56 EXBUCKLANDIOIDEAE
5B
57
TETRACENTRACEAE
67
60
TROCHOOENDRACEAE
70
64
DISANTHOIDEAE
71
69
90
74
RHODOLEIOIDEAE
91
83
PLATANACEAE
[
B)
Adams
Figure 3. consensus tree of two equally par-
simonious trees where the Platanaceae serve as the out-
DISANTHOIDEAE
group. Synapomorphies are designated by rectangles.
EXBUCKLANDIOIDEAE
Numbers numbers
in rectangles refer to character in
HAMAMELIDODEAE
Table
ALTINGIOIDEAE 1
PLATANACEAE
RHODOLEIOIDEAE
EUPTELEACEAE
the group demonstrate an early evolution from a
TETRACENTRACEAE
hamamelid prehamamelid and
or lineage, use the
TROCHODENDRACEAE
Ham-
Platanaceae as an outgroup for analyzing the
we
amelidaceae, obtain an alternative hypothesis
C)
two
of evolution in these families (Fig.
3).
DISANTHOIDEAE
Figure 3 suggests that Altingioideae were the
c
EXBUCKLANDIOIDEAE hamamelidaceous
earliest-derived of the subfami-
HAMAMEUDOIDEAE
Later evolution in the family, according to the
lies.
RHODOLEIOIDEAE
analysis, resulted in the evolution of the other four
ALTINGIOIDEAE
and
subfamilies with Disanthoideae Rhodoleioideae
PLATANACEAE
being the latest-derived. While the consensus tree
EUPTELEACEAE
(Fig. 3) demonstrates the difficulty in resolving the
TETRACENTRACEAE
TROCHODENDRACEAE
node connecting Hamamelidoideae, Exbucklan-
and Disanthoideae-Rhodoleiuideae
dioideae, the
some
group, there support for of the relationships
is
The
proposed by this cladogram. position of Altin-
D)
Hama-
gioideae as the earliest-derived subfamily of
melidaceae supported by 2 synapomorphies,
is 1
DISANTHOIDEAE
EXBUCKLANDIOIDEAE
which include features of phytochemistry, epider-
HAMAMELIDOIDEAE mal anatomy, venation, petiole anatomy, ovule and
ALTINGIOIDEAE seed morphology, and floral structure (Fig. 3).
PLATANACEAE
There corroborative evidence from the
fossil
is
RHODOLEIOIDEAE
scheme As
record phylogenetic
for this (Fig. 3).
EUPTELEACEAE
mentioned previously, Platanaceae can be traced
TETRACENTRACEAE
Lower Up-
TROCHOOENDRACEAE to the Cretaceous, Altingioideae to the
FiGURE 2. Adams consensus trees of the basal Hama- per Cretaceous, and Hamamelidoideae to the Low-
5%
melidae with randomly induced error; Tetracentra- er Paleocene. Foliar megafossil records of Ex-
—
ceae and Trocho<lendraceae are outgroups. A. Consen-
bucklandioideae date from the Upper Oligocene
parsimonious (172
sou^so tvrxe^e^ of three equally trees steps).
^. ....^^ ^^^^.y ^^.^....^>..^^^
^l^^^y ' 'l^^^/i AA (Lakhanpal, 1958) and Lower Miocene of Oreeon
r
B. Consensus tree ot two equally parsimonious trees (ioo ^ t>
'
"^
—
(Brown, 1946) and the Middle Miocene of Wash-
steps). C. Cons(;nsus tree of three equally parsimonious
—
We
trees (174 steps). D. Consensus tree of four trees (174 ington State (Brown, 1946). could find no
steps).
records of fossil leaves, flowers, or fruits of the
However,
Disanthoideae or Rhodoleioideae. TifTney
Rho-
(1986) reviewed records of dispersed seeds of
&
Champ, from Upper Eocene (Mai
dolela the
Walther, 1985) and Disanthus Maxim, from the
968 Annals of the
Garden
Missouri Botanical
& A
Lower Oligocene (Mai Walther, 1978), These Brown, R, 1933. cretaceous sweet gum. Bot. Gaz.
611-615.
(London)
may 94:
data suggest that Platanaceae have been the
II
_
1946. some and
Alterations in iossil living
,. r 1- .
, hneage gave
earhest oflfshoot of a that later rise to
344-355
^^^^^_ ^^^^^ ^^^^^ ^^.^ 3^^
j^
Hamamelidoideae and
Ahingioideae and,
still later, Chang, C. T. 1959. The pollen morphology of Liquid-
&
other subfamilies of the Ilamamelidaceae. ambar and Altingia Nor. Bot. Zurn, (Moscow
L.
& 1375-1380.
Leningrad) 44:
Ilickey Wolfe's (1975) suggestion that ^'Cer-
1964. The pollen morphology of the families
and Trochodendrales form a clade .
cidiphyllales"
Hamamelidaceae and Ahingiaceae. Trudy Bot. Inst.
separate from the remainder of the subclass that
173-
Akad. Nauk SSSR, Ser. 4, Eksper. Bot. 13:
was derived from a prehamamelid or hamamelid
232.
&
lineage need of elaboration. Although our data Chestfrs, K. L, F. R. Gnauck N. F. Hughes. 1967.
in
is
W.
The
Angiospermae. Harland
In B. et (editors),
al.
suggest that the Eupteleaceae might be better placed
Record. Burlington House, London.
Fossil
Hamamelidales,
Trochodendrales ihan the
in the &
Crane, P. R., E. M. Frus K. R. Pedersen. 1986.
they lend some support for the derivation of the
Lower Cretaceous angiosperm flowers: fossil evidence
852-
Hamamelidales from the "[)latanoids" proposed by on early radiation of dicotyledons. Science 232:
854.
Wolfe occurrence of
rly
Platanaceae along wi.h clad.,ic proximity to '''"T:^±J'Sh.:'tI:^!:!!'S.T
tl.eir 'i'!!*"'™
of Flowering Plants. Houghton Mifflin, Boston.
Ahingioideae and distance from rochodendrales
I jgg^ An System
Integrated of Classification
_
does not support inclusion of Trochodendrales in Columbia New York.
of Flowering Plants. llniv. Press,
A
the lineage that led direcUy to the evolution of Dahlgren, R. 1977. commentary on a diagrammatic
presentation of the angiosperms in relation to the
Hamamelidaceae.
Platanaceae and
listribution of character states. PI. Syst. Evol., Su[)pl.
I
Hamamelidaceae
analysis of the
Tlie cladistic
—
253-283.
1:
using the Platanaceae as an outgroup results in a &
Rosendal-Jensen Nielsen. 1981.
S. B.
J.
,
cladogram from current con- A angiosperms with com-
that differs drastically revised classification of the
ments on between chemical and other
correlation
ceptions of evolution the family (Cronquist 1968,
in
&
Young
characters. In D. A. D. S. Seigler (editors),
1981; TakhlajaUj 1969) but agrees overall with
Phytochemlstry and Angiosperm Phylogeny. Praeger
We
the record. are not suggesting that this
fossil New
York.
Publ.,
one phylogeny
reconstruction represents the true DiLCHER, D. L. 1979. Early angiosperm reproduction:
Platanaceae and Hamamelidaceae. Yet, we pro- an introductory report. Rev. Paleobot. Palynol. 27:
of
291-328.
Hama-
pose that the current practice of rooting the
&
Eriksen. 1983. Sycamores are ancient
L.
meUdae the Magnoliidae need of reevalu-
in is in W. 8-
Mus. Colorado Quart. Spring 1983:
trees.
&
Paleobotanical (Dilchcr, 1979; Rctallack
ation.
11.
Hama
Dilcher, 1981) and phylochemical data (Giannasi, Enuress, P. K. 1977. Evolutionary trends in the
1986) suggest that Hamamelidae are at least as melidales-Fagales-group. PI. Syst. Evol., Suppl. 1:
321-347.
^^^ ^'*''
Maenoliidae. Examination of the Ha- ..-^
primitive as
^ ^ F^.elsenstein, 1982. N.,umeric. al, methi odjs rfor inrterr*ing
J.
mamolidae without assumptions as to their pre-
379-404.
evolutionary trees. Quart. Rev. Biol. 57:
may
sumed ancestors our understanding
facilitate Giannasi, D. E. 1986. Phytochemical aspects of phy-
of subclass and of angiosperm evolution. logeny in Hamamelidae. Ann. Missouri Bot. Card.
tlie
417-437.
73:
&
BoHN
GoKNAix, R. B. A. R. Dahlgren. 1979.
J.,
angiosperms.
Tlie distribution of flavonoids in the
LiTERATrRE CiTED
1-30.
Bot. Not. 132:
Adams, E. N. 1972. Consensus techniques and the HaRBORNE, B. 1967. Comparative Biochemistry of
J.
390- New
Academic York.
comparison of taxonomic trees. Syst. Zool. 21: the Flavonoids. Press,
&
397. Harms, H. 1930, Hamamelidaceae. In: A. Engler
&
W. 2nd
Bailey, L C. Nast. 1945. Morphology and K. Prantl, Die Naturlichen Pflanzenfamilien, edi-
Trorhodrndron and Tctracentron, L Band 18a.
relationships of tion.
stem, root, and leaf. Arnold Arbor. 26: 143-153. Hayne, F. G. 1830. Abbildungen von Storaxbaumen.
J.
171-172.
Bllme, K. 1928. Florae Javae nee non Insularum Flora (Stuttgart) 13:
L.
&
Adjacentium. Frank, Brussels. HiCKEY. L. A. Doyle. 1977. Early Cretaceous
J. J.
J.
Bo(;LE, A. L. 1970. Floral morphology and vascular fossil evidence for angiosperm evolution. Bot. Rev.
3-104.
anatomy of the Hamamelidaceae: the apetalous gen- (Lancaster) 43:
&
310-
Hamamelidoideae. Arnold Arbor. 51: Wolee. 1975. The bases of angiosperm
era of
J.
J.
phylogeny: vegetative morphology. Ann. Missouri Bot.
366.
& A 538-589.
C. T. PiiiLBRlGK. 1980. generic atlas of Card. 62:
The
liamamelidaceous pollens. Contr. Gray Herb. 210: HoLLIGK, A. 1936. Tertiary floras of Alaska. U.S.
1-185.
29-103. Geol. Sur. Prof. Pap. 182:
BooTMROYi), E. 1930. The morphology and anatomy Jay, M. 1968. Distribution des flavonoides chez les
L.
136-
Hamamelidacees Taxon
of the inflorescence and flower of the Platanaceae. et families affines. 12:
678-693.
Amer. Bot. 17: 147.
J.
&
Volume Number Schwarzwalder 969
4 Dilcher
78,
Systematic Placement
1991 of
Platanaceae
&
Jha, V. 1977. Chemotaxonomy of the Hamamelida- Retallack, G. D. L. Dilcher. 1981. Arguments for
ceae. Indian Bot. Soc. 56: 44-48. a Glossopterid ancestory of angiosperms. Paleobiol-
J.
54-67.
Koch, B. E. 19(»3. Fossil plants from the lower Paleo- ogy 7:
cene of the Agatdalen (Angmartussut) area, central RoHLF, F. 1982. Consensus indices for comparing
J.
131-144.
Nugssuaq peninsula, northwest Greenland. Meddel. classifications. Math. Bioscience 59:
&
Gr0nland 172: 1-120. R. SOKAL. 1981. Statistical Tables, 2nd
W.
KuTHZKiNA, E. 1974. Platanaceae. In: A. Takhtajan edition. H. Freeman, San Francisco.
(editor), Fossil Flowering Plants of the U.S.S.R., Vol- Saupe, S. 1981. Cyanogenic compounds and angio-
&
ume Magnoliaceae-Eucommiaceae. Nauka, Len- sperm phylogeny. In D. A. Young D. S. Seigler
1.
Phytochemistry and Angiosperm Phyloge-
ingrad. (editors),
New
Lakhanpal, R. N. 1958. The Rujada flora of west ny. Praeger Publ., York.
central Oregon. Univ. Calif, Publ. Geol. Sci. 35: 1- Schwarzwalder, R. N. 1986. Systematics and Early
66. Evolution of the Platanaceae. Ph.D. Dissertation.
A
LiNDLEY, 1826. Natural System of Botany, 2nd Indiana Univ., Bloomington, Indiana.
J.
&
Lon^;man, London. D. Dilcher. 1981. Bisexuality the
edition. L. in
&
Mai, D. H. IL Walther. 1978. Die Floren der pistillate inflorescence of Platanus occidentalis L.
89-98.
Haselbacher im Weisselster-Becken (Bezirk Proc. Indiana Acad. 90:
Serie Sci.
Leipzig, DDR). Abh. Staatl. Mus. Mineral. Geol. Seigler, D. S. 1981. Terpenes and plant phylogeny.
&
Dresden 28: 1-200. In D. A. Young D. S. Seigler (editors), Phyto-
&
1985. Die Obereozanen Floren chemistry and Angiosperm Phylogeny. Praeger Pub!.,
New
des Weisselsler-Beckens und seiner Randgebeite. Abh. York.
Mus. Mineral. Geol. Dresden 33: 1-260. Skvortsova, N. T. 1960. The anatomical structure of
Staatl.
Mathiesen, 1932. Notes on some plants the conducting system of leaf petioles of represen-
F. fossil
J.
from east Gieenland. Meddel. Gr0nland 85: 1-62. tatives of families Hamamelidaceae and Altingeaceae.
Melikian, a. p. 1971. Anatomical structure of the Dokh Akad. Nauk S.S.S.R. 133: 1231-1234.
PAUP:
spermodermis the members of the genera Liquid- SWOFFORD. D. 1984. Phylogenetic Analysis Us-
in
ambar L. and Altingia Nor. in relation to their ing Parsimony. Version 2.2. Illinois Natural History
taxonomy. Nauki 24: 50-55. Survey, Champaign-Urbana, [Photocopy.]
Izv. Biol. Illinois.
1973f.. Seed coat types of Hamamelidaceae Takhtajan, A. 1969. Flowering Plants, Origin and
.
&
and allied families in relation to their systematics. Dispersal. Oliver Boyd, Edinburgh. [Translated by
&
350-359.
(Moscow
Zurn. Leningrad) 58: C. Jeffrey.]
Bot.
anatomy wood
19731). The anatomy of seed coat and sys- Tang, Y. 1943. Systematic of the of
.
Mem.
tematics of the family Hamamelidaceae. Izv. Biol. the Hamamelidaceae. Bufl. Fan. Inst. Biol. 14:
104-105. 8-62.
Nauki
26:
&
Metcalfe, C. R. L. Chalk. 1965. Anatomy of the TiFFNEY, B. H. 1986. Fruit and seed dispersal and the
Dicotyledons, 2nd edition. Oxford Univ. Press, Ox- evolution of the Hamamelidae. Ann. Missouri Bot.
394-416.
Card. 73:
ford.
Nakai, T. 1943. Ordines, Familiae, Tribi, Genera, Sec- TiPPO, 0. 938. Comparative anatomy of the Moraceae
1
tiones, Species, Varietates et Combinationes Novae. and their presumed allies. Bot. Gaz. (London) 100:
1-99.
Tokyo.
Imperial Univ.,
&
W.
Nast, C. G. Bailey. 1945. Morphology and Upchiirch, G. R. 1984. Cuticle evolution in early Cre-
I.
relationships of Trochodendron and Tetracentron. taceous angiosperms from the Potomac Group of
Inflorescence, flower, and Arnold Arbor. Virginia and Maryland. Ann. Missouri Bot. Gard. 71:
II. fruit. J.
267-275. 522-550.
26:
&
A
1946. Morphology of Eiiptelea WiLLlS, C. 1966. Dictionary of the Flowering
J.
and comparison with Trochodendron. Arnold Ar- Plants and Ferns, 7th edition. Cambridge Univ. Press,
J.
186-191. Cambridge.
bor. 27:
&
Newberry, S. 1898. The later extinct floras of North Wilson P. 1905. Altingiaceae. In: N. L. Britton L.
J.
New
America. U.S. Geol. Survey Mon. 35: 1-295. [A M. Llnderwood (editors). North American Flora.
New
posthumous work edited by A. Rollick.] York Botanical Garden, Bronx, York.
Pkaglowski, 1974. The pollen morphology of the Wolfe, A. 1966. Tertiary plants from the Cook Inlet
J. J.
Trochodendraceae, Tetracentraceae, Cercidiphylla- Region, Alaska. Prof. Pap. U.S. Geol. Sur. 398-B:
B1-B32.
ceae and Eupteleaceae with reference to taxonomy.
&
449-467.
Pollen Spores 16: 1973. Fossil forms of the Amentiferae. Brit-
.
Rao, p. R. M. 1974. Seed anatomy in some Hama- tonia 25: 334-355.
&
melidaceae and phylogeny. Phytomorphology 24: Zavada, M. S. D. L. DnxHER, 1986. Comparative
113-139. morphology and phylogeny
pollen relationship to
its
&
Rao, T. a. O. P. Bhupal. 1974. Typology of foliar of pollen in the Hamamelidae. Ann. Missouri Bot.
348-381.
sclereids in various taxa of Hamamelidaceae. Proc. Gard. 73:
127-138.
Indian Acad. Sci., Sect. B, 79: