ELSEVIER

Review of Palaeobotany and Palynology 103 (1998) 235–251

Revision of the so-called ‘Reevesia’ fossil woods from the Tertiary inJapan — a proposal of the new genus Wataria (Sterculiaceae)
Kazuo Terada Ł , Mitsuo Suzuki
Botanical Garden, Faculty of Science, Tohoku University, Kawauchi, Aoba, Sendai 980, Japan
Received 17 April 1997; accepted 22 January 1998

Abstract
The Tertiary so-called ‘Reevesia’ fossil woods from the Oligocene and Miocene in Japan are characterized by distinct
ring porosity and tile cells in rays. Comparison of these ‘Reevesia’ fossil woods with extant woods showed that only one
specimen has characteristics of Reevesia, while the others differ from all extant genera of the Malvales, and so are regarded
as belonging to an extinct genus of Sterculiaceae, most similar to the extant genus Triplochiton Schumann and the fossil
genus Triplochitioxylon Manchester, and the new genus Wataria K. Terada et M. Suzuki, gen. nov. is proposed for thosefossil woods. Reevesia miocenica Watari and R. oligocenica M. Suzuki are transferred into Wataria as W. miocenica K.
Terada et M. Suzuki, comb. nov. and W. oligocenica K. Terada et M. Suzuki, comb. nov., respectively, and one new species,
Wataria parvipora, is described here. The specimen with characteristics of Reevesia is described as a new species Reevesia
japonoxyla K. Terada et M. Suzuki, sp. nov.  1998 Elsevier Science B.V. All rights reserved.
Keywords: fossil wood; Reevesia; Sterculiaceae; Tertiary; tile cells; Triplochiton; Wataria

1. Introduction
Tile cells are a distinctive wood anatomical feature.
They are defined as a special type of apparently empty
upright (or rarely square) ray cells occurring in horizontal series usually interspersed among the procumbent cells of rays (IAWA Committee, 1989). Tile cells
are restricted to the order Malvales (Chattaway, 1933;
Carlquist, 1988; IAWA Committee, 1989). Tile cells
are classified into two extreme types, the Durio andPterospermum types, and an intermediate of the twois also present (Chattaway, 1933).
Ł Corresponding author. Present address: Fukui Pref. Board of

Education, Cultural Affairs Division, Dinosaur Museum Project,33-17-1 Ohte, Fukui 910-8580, Japan. Tel.: C81-776-20-0581;
Fax: C81-776-20-0672; E-mail: k-terada@pref.fukui.jp

Watari (1952) discovered three fossil woods thatare characterized by distinct ring porosity and thepresence of tile cells in rays from the Mioceneof Yamagata Prefecture, northern Honshu (Fig. 1,a–c). Because Kanehira (1921), who worked on
Formosan woods, indicated that only the extant
species Reevesia formosana Sprague of Sterculiaceae had ring porosity and intermediate tile cells,
he assigned the fossil woods to Reevesia. Watari(1952) named the fossil woods Reevesia miocenica
Watari and considered them closely related to R.
formosana.
Subsequently, Reevesia oligocenica Suzuki, which
is very similar to Reevesia miocenica, was describedfrom the Oligocene of Fukuoka Prefecture, northern
Kyushu (Fig. 1, i; Suzuki, 1976). Additional specimens of R. miocenica were reported from the Lower

0034-6667/98/$19.00 c 1998 Elsevier Science B.V. All rights reserved.
PII: S 0 0 3 4 - 6 6 6 7 ( 9 8 ) 0 0 0 3 9 - 6

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Fig. 1. Locality map of so-called ‘Reevesia’ fossil woods in Japan. a D Iragawa, Atsumi-machi, Nishitagawa-gun, Yamagata Prefecture;b D Isagodani, Tsuruoka City, Yamagata Prefecture; c D Tamugimata, Higashi-mura, Yamagata Prefecture; d D Nametaki, Wajima City,
Ishikawa Prefecture; e D Nigoriike, Monzen-machi, Ishikawa Prefecture; f D Minamihora, Kawabe-machi, Kamo-gun, Gifu Prefecture;h D the river bed of Kiso River at a border of Minokamo and Kani Cities, Gifu Prefecture; i D Koinoura, Tsuyazaki-machi, Fukuoka
Prefecture.

Miocene of Noto Peninsula, Ishikawa Prefecture, central Honshu (Fig. 1, e; Suzuki and Watari, 1994).
Recently, Lower Miocene fossil forests composedof many erect stumps were found on the river bed of
Kiso River, Gifu Prefecture, central Honshu (Fig. 1,
h; Shikano, 1995). A preliminary observation of thewood structure of those stumps revealed that mostof them had ring porosity and tile cells, and werequite similar to Reevesia miocenica (Terada, 1995).
Furthermore, two fossil woods with ring porosity andtile cells were also found from the Lower Miocene
Hachiya Formation of Kawabe-machi near the fossil
forests (Fig. 1, f, g).

The so-called ‘Reevesia’ fossil woods characterized by distinct ring porosity and tile cells were
known from various areas and ages in the Tertiaryof Japan (Fig. 1). This fact suggests that ‘Reevesia’trees had flourished during the Tertiary of Japan,
although they are now extinct in Japan.
During a study on Tertiary fossil woods in Japan,
it was noticed that most ‘Reevesia’ fossil woods
differed in some features from the genus Reevesia
and from other extant genera of Malvales. Therefore,
we re-examined all ‘Reevesia’ fossil woods fromJapan and revised them systematically.

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2. Materials and methods
Nine specimens were examined in detail (Table 1). The specimen of no. 35252 is the syntype ofReevesia miocenica Watari, which is one of the three
specimens reported by Watari (1952) and depositedin the University Museum, University of Tokyo(TI). It was collected from the Lower Miocene
at Iragawa, Atsumi-machi, Nishitagawa-gun, Yamagata Prefecture (38º400 N, 139º380 E; Fig. 1, a).
No. 71189 is the holotype of Reevesia oligocenica
Suzuki, which is one of the two specimens reported by Suzuki (1976), and collected from theLower Oligocene Tsuyazaki Formation at Tsuyazaki,
Fukuoka Prefecture (33º490 N, 130º270 E; Fig. 1, i;
Sakai, 1994). No. 53108 is one of the five specimens
reported as Reevesia miocenica from the Lower
Miocene Nawamata Formation of Noto Peninsula,
Ishikawa Prefecture (37º200 N, 136º480 E; Fig. 1,e; Suzuki and Watari, 1994). No. 53219 is collected from the Lower Miocene Nawamata Formation at Nametaki, Wajima City, Ishikawa Prefecture (37º210 N, 136º480 E; Fig. 1, d). No. 59019
is collected in the volcanic breccia of the Lower
Miocene Hachiya Formation on the right riverside of
Hida River at Kawabe Dam, Kawabe-machi, Kamogun, Gifu Prefecture (35º280 N, 137º040 E; Fig. 1,
g; Nomura, 1986, 1992). No. 59044 is also collected from the Hachiya Formation at Minamihora, Kawabe-machi, Kamo-gun, Gifu Prefecture(35º290 N, 137º030 E; Fig. 1, f), about 3 km north-237

west of the Kawabe Dam. Nos. 59061, 59082 and
59098 are the representatives of 27 fossil stumps
which were collected from the fossil forests on theriver bed of Kiso River at a border of Minokamo
and Kani Cities, Gifu Prefecture (35º260 N, 137º020 E;
Fig. 1, h; Terada, 1995). The horizon of the forests is
the Lower Miocene Nakamura Formation (Shikano,1995).
All the fossil specimens examined are silicified
wood. Ground cross, tangential and radial thin sections were prepared by the conventional technique.
Excepting for the loaned specimens of Reevesia
miocenica (nos. 35252, 53108) from TI, all theother fossil specimens examined in this paper aredeposited in the Wood Collection of the Herbarium,
Faculty of Science, Tohoku University, Sendai, Japan(TUSw).
A comparison of wood structure of those fossils
with extant species of the Malvales was made using samples deposited in TUSw and loaned fromFU (Forest Products Department, Kyushu University, Fukuoka, Japan), Kw (Royal Botanic Garden, Kew, England), Lw (Rijksherbarium, Leiden,
Netherlands), MADw or SJRw (U.S. Forest Products
Laboratory, Madison, USA), Tw (Royal Museum
for Central Africa, Tervuren, Belgium) and TWTw
(Forestry and Forest Products Research Institute,
Tsukuba, Japan). For searching candidates which
may have affinity with the fossil woods, the Fossil Dicotyledonous Wood Database (Wheeler, 1991)
was used.

Table 1
List of the examined specimens of the so-called ‘Reevesia’ fossil woods
Specimen No.

Locality

Formation

Horizon e

35252
71189
53108
53219
59019
59044
59061
59082
59098

Iragawa, Atsumi, Yamagata Pref. (Fig. 1a) a
Koinoura, Tsuyazaki, Fukuoka Pref. (Fig. 1i) b
Nigoriike, Monzen, Ishikawa Pref. (Fig. 1e) c
Nametaki, Wajima, Ishikawa Pref. (Fig. 1d)
Kawabe, Kamo, Gifu Pref. (Fig. 1g)
Minamihora, Kamo, Gifu Pref. (Fig. 1f)
Kiso River, Minokamo-Kani, Gifu Pref. (Fig. 1h)Kiso River, Minokamo-Kani, Gifu Pref. (Fig. 1h)Kiso River, Minokamo-Kani, Gifu Pref. (Fig. 1h)

Atsumi F. d
Tsuyazaki F.
Nawamata F.
Nawamata F.
Hachiya F.
Hachiya F.
Nakamura F.
Nakamura F.
Nakamura F.

L. Mio.
L. Oligo.
L. Mio.
L. Mio.
L. Mio.
L. Mio.
L. Mio.
L. Mio.
L. Mio.

a Syntype specimen of Reevesia miocenica Watari (1952) in TI.
b Holotype specimen of Reevesia oligocenica Suzuki (1976) in TUSw.
c R. miocenica in Suzuki and Watari (1994) in TI.
d The formations are deduced by us from the locality.
e L. Mio D Lower Miocene; L. Oligo.D Lower Oligocene.

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2.1. Taxonomic treatments
MALVALES
STERCULIACEAE

Wataria K. Terada et M. Suzuki, gen. nov.
Type: Wataria miocenica (Watari) K. Terada et
M. Suzuki, comb. nov.
Diagnosis of genus: Wood distinctly ring porouswith wide vessels at beginning of growth rings incross section. Growth rings distinct. Wide vessels inearlywood mostly solitary, round. Narrow vessels inlatewood solitary, and in groups of 2–5 with vasicentric parenchyma; very thick-walled. Vessel elements
short; perforation plates exclusively simple; intertracheary pitting alternate; helical thickenings absent;
thin-walled tyloses often present in wide vessels;
gum-like deposits rarely present in narrow vessels.
Axial parenchyma abundant, apotracheal and paratracheal; apotracheal in uni- or biseriate tangential
bands alternating with 1–3 rows of fibers in latewood; paratracheal abundant, vasicentric forming 1–
3-seriate sheath; conspicuously storied; fusiform andin strands of 2–10 cells or more; crystals absent.
Rays heterocellular; uni- or multiseriate. Multiseriate rays 2–10-seriate, composed of tile cells andprocumbent cells. Tile cells Pterospermum type orintermediate to the Durio type; brown-colored substances rarely observed in the tile cells; crystals
rarely present.
Etymology: The genus Wataria is named for the late
Dr. Shunji Watari for his pioneer work on this taxon.
Wataria miocenica (Watari) K. Terada et M. Suzuki,comb. nov.
Basionym: Reevesia miocenica Watari, 1952, J. Fac.
Sci. Univ. Tokyo, Sect. 3 Bot. 6, 126–129 photos
8C–D, 9A–E.
Typification: Although Watari (1952) originally described three specimens as Reevesia miocenica andindicated no holotype, we designated a lectotype inthis paper according to the article 9 of ICBN (Greuter
et al., 1994) as Watari’s description is chiefly basedon the specimen of no. 35252.
Lectotype: No. 35252; Slides: 35252-1 to -6.
Repository: University Museum, University ofTokyo, Japan.

Type locality: Iragawa, Atsumi-machi, Nishitagawa-gun, Yamagata Prefecture (38º400 N, 139º380 E;
Fig. 1, a).
Type stratum and stratigraphic horizon: Atsumi Formation (deduced from the locality), Lower Miocene(Sato et al., 1989; Yamaji, 1989).
Diagnosis: This species is distinguished from otherspecies in the genus by: (1) wide vessels arranged
in 1–3 layers in earlywood; and (2) very wide earlywood vessels of about 270 µm in mean tangential
diameter (Table 2).
Comments: Although Watari (1952) described
Reevesia miocenica as having faint helical thickenings in some vessels, our re-examination of R.
miocenica including the type specimens, revealed
that helical thickenings were absent. The character
described by Watari (1952) is mutated herein.
Syntypes: No. 35008 in TI, collected from the Lower
Miocene at Isagodani, Tsuruoka City, Yamagata Prefecture (38º390 N, 139º460 E; Fig. 1, b). No. 35213 in
TI, collected from the Lower Miocene at Tamugimata, Higashi-mura, Yamagata Prefecture (38º340 N,
139º560 E; Fig. 1, c).
Additional specimens: Nos. 53103, 53108, 53110,
53117 and 53123 in TI, collected from theLower Miocene Nawamata Formation at Nigoriike, Monzen-machi, Ishikawa Prefecture (37º260 N,
136º480 E; Fig. 1, e) (Suzuki and Watari, 1994).
No. 53219, a new specimen reported in this paper,collected from the Lower Miocene Nawamata Formation at Nametaki, Wajima City, Noto Peninsula,
Ishikawa Prefecture (37º210 N, 136º480 E; Fig. 1, d),
and deposited in TUSw.
Wataria oligocenica (M. Suzuki) K. Terada et
M. Suzuki, comb. nov.
Basionym: Reevesia oligocenica M. Suzuki, 1976,
Bot. Mag. Tokyo 89, 65–68, fig. 4A–G.
Holotype: No. 71189; Slides: 71189-1 to -6.
Paratype: No. 71208; Slides: 71208-1 to -3.
Repository: Herbarium, Faculty of Science, Tohoku
University, Sendai, Japan.
Type locality: Koinoura, Tsuyazaki-machi, Fukuoka Prefecture, northern Kyushu Japan (33º490 N,
130º270 E; Fig. 1, i).
Type stratum and stratigraphic horizon: Tsuyazaki
Formation, Lower Oligocene (Sakai, 1994).

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239

Table 2
Anatomical characters of the so-called ‘Reevesia’ fossil woods
Specimen(No.)

Wide vessels in earlywood
TD a (µm)

Narrow vessels in latewood

RD a (µm)

LV

NV

WT (µm)200–350 (294)300–450 (376)
250–320 (288)

2–3
2–3
1–3

solitary
solitary
solitary

Wataria oligocenica comb. nov.
71189
140–260 (210)
150–250 (199)

3–5

Wataria parvipora sp. nov.
59044
100–250 (176)59061
125–220 (167)59082
110–275 (171)59098
90–200 (150)

105–285 (187)
125–250 (187)
100–250 (185)
125–275 (196)

Reevesia japonoxyla sp. nov.
59019
110–210 (162)

170–290 (235)

Wataria miocenica comb. nov.
35252
200–350 (287)53108
220–350 (265)53219
200–300 (257)

HT b

Rays
MRH a (µm)

MRW a (µm)

TT

5.0
5.8
6.8

150–1150 (621)
175–1950 (870)
100–2200 (1280)

70–150 (105)
20– 75 (50)
40– 70 (54)I
I
P

solitary

4.2

200–1000 (688)30– 75 (59)

I

1–2
1–3
1–2
1–2

solitary
solitary
solitary
solitary

8.3
7.1
6.8
6.7

100–2000 (716)unmeasurable
550–1200 (803)unmeasurable

10–100 (36)

P
I
I
I

2–5

cluster

2.9

550–2700 (1392)

40– 70 (54)C

70–140 (112)P

TD D tangential vessel diameter; RD D radial vessel diameter; LV D layers of wide vessels in earlywood.
NV D narrow vessels; WT D mean wall thickness of narrow vessel; HT D helical thickenings on vessel walls.
MRH D multiseriate ray height; MRW D multiseriate ray width.
TT D type of tile cells: P D Pterospermum-type; I D intermediate of Durio- and Pterospermum-types.
a Range (mean). b
D absent; C D present.

Diagnosis: This species is distinguished by following features from other species in the genus: (1)wide vessels arranged in 3–5 layers in earlywood;and (2) earlywood vessels of about 210 µm in meantangential diameter (Table 2).
Wataria parvipora K. Terada et M. Suzuki, sp. nov.
Holotype: No. 59044; Slides: 59044-1, 2, 3, 4,5, 6
and 7. This hand-sample is a stem wood about 5 cm
in diameter, 11 cm long.
Repository: Herbarium, Faculty of Science, Tohoku
University, Sendai, Japan.
Type locality: Minamihora, Kawabe-machi, Kamogun, Gifu Prefecture (35º290 N, 137º030 E; Fig. 1, f).
Type stratum and stratigraphic horizon: Hachiya
Formation, Lower Miocene (Nomura, 1986, 1992)
Paratypes: 27 specimens, nos. 59058–59066,
59077–59084, 59089–59095, 59097–59099, inTUSw, were collected from the fossil forests of theLower Miocene Nakamura Formation on the river
bed of Kiso River at a border of Minokamo and Kani
cities, Gifu Prefecture (35º260 N, 137º020 E; Fig. 1, h;
Shikano, 1995). Most of them were compressed anddeformed during burial.

Etymology: The specific epithet, parvipora, is derived that this species has much narrower vessels inearlywood than that of Wataria miocenica.
Description: Wood distinctly ring porous (Plate I, 1,
2). Growth rings distinct; 0.5–1.5 mm wide (Plate I,1, 2).
Wide vessels arranged in one or two layers in earlywood in cross section, mostly solitary and sometimes in radial or oblique multiples of two to three;solitary vessels round, 100–250 (mean 176) ð105–
285 (mean 187) µm in tangential ð radial diameters;
walls slightly thickened (about 4 µm) (Plate I, 1, 2).
Narrow latewood vessels solitary in cross section, notin multiples nor in clusters; forming groups of 2–5
vessels with vasicentric parenchyma in latewood; circular or angular especially at the end of growth rings,
15–100 (mean 50) µm in tangential diameter; walls
8.3 µm thick (Plate I, 1, 2). Vessel elements short,
65–290 (mean 184.8) µm long in earlywood, 145–
210 (mean 175.6) µm long in latewood; end walls
horizontal or somewhat oblique in earlywood, andoblique in latewood (Plate I, 3). Perforation plates
exclusively simple (Plate I, 3). Intertracheary pits alternate, dense, circular or polygonal in outline, about
5 µm in diameter, with horizontally elongated aper-

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PLATE I

Wataria parvipora K. Terada et M. Suzuki, sp. nov. Holotype: no. 59044. Scale bars indicated in 1. The bar is 250 µm in 1, 100 µm in
2–4, 50 µm in 5 and 6.
1. Cross section showing ring porous wood with distinct growth rings.
2. Cross section showing wide vessels in earlywood, thick-walled solitary narrow vessels in latewood and uniseriate banded parenchyma
in latewood.
3. Radial section showing wide vessel with simple perforation and alternate intertracheary pitting.
4. Tangential section showing both uniseriate rays and multiseriate rays with tile cells, and storied structure of fibers, axial parenchyma
and narrow rays.
5. Tangential section showing multiseriate rays with tile cells and densely pitted vaisentric parenchyma around vessels.
6. Radial section showing tile cells and procumbent cells of a multiseriate ray.

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tures (Plate I, 3). Helical thickenings not observed.
Thin-walled tyloses often present (Plate I, 1). Gumlike deposits rarely observed only in narrow vessels.
Fibers consisting ground mass of the latewood
with axial parenchyma; square or polygonal in crosssection, 7–25 (mean 15.5) µm in diameter; walls
about 3.5 µm thick; not septate; storied (Plate I, 2).
Axial parenchyma abundant, apotracheal andparatracheal; conspicuously storied (Plate I, 4);fusiform and in strands of 2–10 cells, up to 18
cells (Plate I, 5), 225–325 (mean 261) µm in length(Plate I, 2, 4); polygonal to round in cross section,
10–40 (mean 21.5) µm in diameter. Apotracheal
parenchyma in uni- or biseriate tangential bands alternating with 1–3 rows of fibers in latewood (Plate I,
2). Paratracheal parenchyma abundant and vasicentric forming 1–3-seriate sheaths (Plate I, 2). Vessel-parenchyma pits small circular, alternate, dense(Plate I, 5). Crystals absent.
Rays heterocellular, 1–7-seriate (Plate I, 4, 5).
Uniseriate rays 5–12.5 (mean 8.3) µm wide and
37.5–275 (mean 146.5) µm tall. Multiseriate rays
mostly 3–5 cells and 10–100 (mean 36.1) µm wide,
and 100–2000 (mean 715.9) µm tall; composed oftile cells and procumbent cells (Plate I, 6). Tile
cells Pterospermum type to intermediate (Plate I, 5,6); 7.5–20 (mean 13.5) ð15–45 (mean 25.5) ð5–
27.5 (mean 15.8) µm in tangential ð vertical ðradial diameters. Procumbent cells 3–10 (mean 6)
ð5–17.5 (mean 10.4) ð15–37.5 (mean 26.3) µm intangential ð vertical ð radial diameters. Ray-vesselpits alternate and dense. Brown-colored substances
sometimes observed in tile cells (Plate I, 6). Crystals
not observed, but rarely with swollen cells which
may suggest the occurrence of crystals.
Genus Reevesia Lindley, 1827
Reevesia japonoxyla
K. Terada et M. Suzuki, sp. nov. (Plate II)
Holotype: No. 59019; Slides: 59019-1 to -5. This
sample is a trunk with estimated diameter of 30 cm.
Repository: Herbarium, Faculty of Science, Tohoku
University, Sendai, Japan.
Type locality: the right riverside of Hida River at
Kawabe Dam, Kawabe-machi, Kamo-gun, Gifu Prefecture (35º280 N, 137º040 E; Fig. 1, g).

241

Type stratum and stratigraphic horizon: Hachiya
Formation, Lower Miocene (Nomura, 1986, 1992)
Etymology: From Japan.
Description: Wood distinctly ring porous (Plate II,
7). Growth rings distinct 0.5–1.5 mm wide (Plate II,
7).
Wide vessels arranged in two to five layers inearlywood, mostly solitary and sometimes in radialor oblique multiples of two to three in cross section; solitary vessels radially elongated oval, 110–210 (mean 162) ð170–290 (mean 234.5) µm intangential ð radial diameters; walls thin (about 3µm) (Plate II, 1, 2). Narrow vessels in latewoodcommonly in clusters of 5–20 vessels; each vessel polygonal in cross section, 10–50 (mean 24.1)µm in diameter; walls 2.9 µm thick (Plate II, 2).
Latewood vessels distributed in ulmiform or in tangential bands (Plate II, 1, 2). Vessel elements 150–
330 (mean 263.8) µm long; end walls horizontal orsomewhat oblique in earlywood, and oblique in latewood (Plate II, 3–5). Perforation plates exclusively
simple (Plate II, 3–5). Intertracheary pits alternate,
dense, circular or polygonal in outline, about 5 µm
in diameter, with horizontally elongated apertures(Plate II, 3). Helical thickenings distinct on narrowvessel walls (Plate II, 4). Tyloses not observed. Gumlike deposits often present in vessels (Plate II, 1).
Fibers forming the ground mass of the latewood;square or polygonal in cross section (Plate II, 2),
10–20 (mean 15.1) µm in diameter; walls about 4µm thick; not septate.
Axial parenchyma apotracheal and paratracheal(Plate II, 2); conspicuously storied; fusiform andin strands with 2–4 cells (Plate II, 5). Apotracheal parenchyma diffuse-in-aggregates in latewood
and paratracheal vasicentric scanty (Plate II, 2).
Parenchyma cells 10–25 (mean 16.4) µm in diameter in cross section (Plate II, 2) and 250–375(mean 322.6) µm in length (Plate II, 5). Crystals
absent.
Rays heterocellular, 1–7-seriate (Plate II, 5).
Uniseriate rays 15–20 (mean 17.5) µm wide, 92.5–500 (mean 205.5) µm tall. Multiseriate rays mostly
3–5 cells and 40–70 (mean 54.3) µm wide and
550–2700 (mean 1392) µm tall; composed of tile
cells and procumbent cells (Plate II, 5, 6). Tile cells
Pterospermum type (Plate II, 5, 6); 10–25 (mean
16.7) ð15–57.5 (mean 32.6) ð4–20 (mean 11.2)

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PLATE II

Reevesia japonoxyla K. Terada et M. Suzuki, sp. nov. Holotype: no. 59019. Scale bars indicated in 1. The bar is 250 µm in 1, 100 µm in
2, 3 and 5, 50 µm in 4 and 6.
1. Cross section showing ring porous wood with 3–5 layers of wide vessels and sporadic gum-like dark contents in vessels inearlywood.
2. Cross section showing wide earlywood vessels and narrow latewood vessels in clusters.
3. Radial section showing wide vessels with simple perforation and alternate intertracheary pitting.
4. Radial section showing helical thickenings on vessel walls.
5. Tangential section showing multiseriate rays with tile cells and storied structure.
6. Radial section showing Pterospermum-type tile cells in multiseriate rays.

K. Terada, M. Suzuki / Review of Palaeobotany and Palynology 103 (1998) 235–251µm in tangential ð vertical ð radial diameters.
Procumbent cells 7.5–15 (mean 10.3) ð7.5–17.5(mean 12.9) ð35–85 (mean 60.6) µm in tangential
ð vertical ð radial diameters. Ray-vessel pits alternate, minute and dense. Brown-colored substanceand crystals invisible.

3. Affinities and discussions
3.1. Comparison with extant Reevesia
All ‘Reevesia’ fossil woods examined have incommon the following characteristics: (1) distinct
ring porosity; (2) exclusively simple perforation
plates; (3) alternate intertracheary pitting; (4) storied structure; and (5) heterocellular multiseriaterays with tile cells. They can be divided into twogroups by features of the latewood vessels (Table 2). One group has mostly solitary and thickwalled latewood vessels without helical thickenings.
Specimens reported as Reevesia miocenica Watarior R. oligocenica Suzuki, and newly obtained specimens, nos. 53219, 59044, 59058–59066, 59077–
59084, 59089–59095, 59097–59099, belong to thisgroup. The other group is represented by only one
specimen, no. 59019, which has latewood vessels inclusters and helical thickenings on vessel walls. As
discussed below, the former group is designated asa new fossil wood genus, Wataria, of Sterculiaceae,
and the latter group is a new species of Reevesia, R.
japonoxyla (Table 2).
When Watari (1952) described Reevesia miocenica (D Wataria miocenica comb. nov.), he foundonly extant species Reevesia formosana Sprague hasring porosity and tile cells based on a description ofwood anatomy by Kanehira (1921). As tile cells of
his fossil woods and Reevesia formosana are intermediate between the Durio and the Pterospermum
type, he described his fossils as a new species ofReevesia closely related to R. formosana (Watari,1952).
Although Kanehira (1921) described Reevesia
formosana as ring porous wood, its ring porosity
is not well developed and is quite different from thatof Watari’s fossils. Distinct ring porosity is one of
the common features common to all fossil woods of
Wataria. We re-examined Kanehira’s wood sample

243(FU-A50) of Reevesia formosana (Plate III, 1–3), an
additional wood sample (TUSw-10584) of R. formosana obtained from Taiwan, and fossil woods of
Wataria including the syntype specimen of Watari’s
Reevesia miocenica (Table 3).
As a result, we found many wood anatomical differences between Reevesia formosana and Wataria
fossil woods (Table 3; Plate III, 1–3): (1) Wataria
has distinct ring porosity while R. formosana does
not; (2) latewood vessels of Wataria are mostly
solitary while those of R. formosana are usuallyin clusters; (3) helical thickenings on vessel walls
are not observed in Wataria but distinct in R. formosana; (4) vessel occlusions are thin-walled tyloses
in Wataria and gum-like deposits in R. formosana;(5) vasicentric parenchyma is abundant in Wataria,
while only scanty in R. formosana; (6) apotracheal
parenchyma in Wataria is distinct and in uni- orbiseriate tangential bands while that of R. formosana
is in diffuse-in-aggregates; (7) storied structure isdistinct in Wataria, while indistinct in R. formosana;and (8) the number of cells in a parenchyma strand
is 2–10 cells or more cells in Wataria, while only
2–4 cells in R. formosana. These anatomical differences suggest that Wataria has no direct affinity to
Reevesia formosana.
There are about 18 extant species (Hsue, 1984)of Reevesia distributed in the subtropical to warm–
temperate regions of Taiwan (Li and Lo, 1993),
southern China (Hsue, 1984), eastern Himalaya(Masters, 1875) and Java (Backer and Bakhuizen
van den Brink, 1963). In addition, Monachino (1940)
reported a new genus with only one species, Veeresia clarkii Monachino et Moldenke, from Hidalgo,
Mexico, as being closely related to Reevesia. Solheim (1987) placed Veeresia in synonymy withReevesia and reported a new species from Chiapas and Nicaragua, central America. However, he
has not yet effectively published about it because it
is only in an abstract. So in this paper we treat these
two species as Veeresia.
Of about 18 species of Reevesia, only a few
descriptions of wood anatomy are available. Our observations and the descriptions by Tang (1932, 1936)
of Chinese Reevesia pubescens Masters [D syn. R.
cavaleriei Léveille (Hsue, 1984)] indicate that it has
semi-ring or indistinct ring porosity, latewood vessels in clusters, and helical thickenings in latewood

244

Table 3
Anatomical comparison of ‘Reevesia’ fossil woods with extant woods and fossil records
Botanic nameSpecimen

Porosity‘Reevesia’ fossil woods
Wataria miocenica comb. nov.
Wataria oligocenica comb. nov.
Wataria parvipora sp. nov.
Reevesia japonoxyla sp. nov.

Vessels

Parenchyma

MTD VEL aocclusion NV

HT b PL

Rays
NS

storiedCP b TT CR b

250–425 (332)
225–360 (291)
225–325 (261)
250–375 (322)

2–10
2–10
2–10
2–4

distinct
distinct
distinct
distinct

IP
I
IP
P

2–4
2–4
2–4
2–4
2–4
2–4
2–4
2–4
2–4
2–4
2–4
2–4
2–4

indistinct
indistinct
distinct
distinct
distinct
indistinct
indistinct
indistinct
indistinct
distinct
distinct
distinct
distinct

C
C
C
C
C
C
?

I
I
P
I
P
I
I
I
I
I
I
I
I

C
C
C
C
C
C
C

distinct

?

I

C

35252
71189
59044
59019

ring
ring
ring
ring

350
260
250
210

70–350 (234)75–325 (219)65–290 (185)
150–330 (264)tylosistylosistylosisgum

solitary
solitary
solitaryclusterring
semi-ringring
semi-ringring
semi-ring
ring
ring
ring
semi-ring
diffusesemi-ring
diffuse

250
170
210
100
200
140
170
200
190
325
300
270
130

225–450 (347)
220–440 (320)320–500 (403)
280–540 (457)500–740 (624)230–590 (443)
200–470 (371)
230–520 (388)
240–500 (390)75–335 (230)
100–350 (272)
100–350 (247)
129–230 (163)

gum
gum
gum
gum
?
gum
?
?
?
tylosistylosistylosistylosisclusterclusterclusterclusterclustercluster
multiple of 2–3
multiple of 2–3
multiple of 2–3
solitary
solitary
solitary
solitary

C
C
C
C
C
C
C
C
C

Triplochiton zambesiacus c

FU-A50TUSw-10584
SJRw-21786
SJRw-21895
Kw-38115
TUSw-10585
MADw-42819
MADw-42820
MADw-44043
TWTw-2772
TWTw-4265
TWTw-5622
PRFw-26601

250–475 (380)
270–400 (343)
320–480 (417)
350–520 (463)
500–750 (625)430–600 (497)
330–450 (412)
310–480 (406)
300–600 (421)
225–385 (298)210–360 (302)
250–360 (304)
130–201 (165)

Fossil records
Triplochitioxylon oregonensis c

NBW-50semi-ring 287
to diffuse

solitary?

431–660 (532) 2–4

Extant species
Reevesia formosana
Reevesia pubesens
Reevesia thyrsoidea
Reevesia wallichii
Veeresia clarkii

Triplochiton scleroxylon

273–502 (403) tylosisC

MTD D maximum tangential vessel diameter; VEL D vessel element length; NV D narrow vessels; HT D helical thickenings.
PL D axtial parenchyma length; NS D number of cells in a parenchyma strand; CP D crystal in axial parenchyma.
TT D type of tile cells; I D intermediate of Durio and Pterospermum types; P D Pterospermum type; CR D crystal in ray cells.
a Range (mean).
b
D absent; C D present.
c Data from Manchester (1979).

=C
=C

C

K. Terada, M. Suzuki / Review of Palaeobotany and Palynology 103 (1998) 235–251(No.)

K. Terada, M. Suzuki / Review of Palaeobotany and Palynology 103 (1998) 235–251

245

PLATE III

Scale bars indicated in 1. The bar is 250 µm in 1 and 4, 100 m in 2, 3, 5 and 6.
1–3. Microphotographs of extant Reevesia formosana (FU-A50).
1.
Cross section showing semi-ring porous wood and thin-walled narrow vessels in clusters.
2.
Tangential section showing multiseriate rays with tile cells, helical thickenings in vessels and unstoried structure of axial elements.
3.
Radial section showing tile cells.
4–6. Microphotographs of extant Triplochiton scleroxylon (TWTw-2772).
4.
Cross section showing semi-ring porous wood with thin-walled tyloses in vessels.
5.
Tangential section showing multiseriate rays with tile cells, vasicentric parenchyma and storied structure of fibers, axial
parenchyma and rays.
6.
Radial section showing rays with tile cells and crystals in both rays and axial parenchyma.

246

K. Terada, M. Suzuki / Review of Palaeobotany and Palynology 103 (1998) 235–251

vessels (Table 3). Reevesia thyrosidea Lindley ofChina is diffuse to ring porous wood with latewood
vessels in clusters and distinct helical thickenings(Table 3; Cheng et al., 1992; our observations). Moll
and Janssonius (1906), cited in Metcalfe and Chalk(1950), described Reevesia wallichii Brown fromwestern Java as it did not have vessels in clusters,
helical thickenings on vessel walls or tile cells. Our
study of the specimen (SJRw-30221) described byMoll and Janssonius (1906) indicates, however, thatit has typical tile cells and vessels in multiples of
2–5. Metcalfe and Chalk (1950) suggested that thedifferences in wood anatomy of R. wallichii fromother Reevesia species might be due to the difference of habitat. However, our observation on an
additional specimen of R. wallichii (TUSw-10585)
reveals that the wood anatomy of the species is quite
different from the Moll and Janssonius’ description
as follows: it has semi-ring to diffuse porosity withlatewood vessels in clusters, distinct helical thickenings and typical tile cells. Solereder (1908) also
reported that Reevesia wallichii has helical thickenings in vessels. Furthermore, Reevesia wallichii
occurs only in the eastern Himalaya region (Masters,
1875; Stainton, 1988; Long and Rae, 1991), and does
not reach to Java where Moll and Janssonius’ specimen was collected (Backer and Bakhuizen van den
Brink, 1963). This suggests that the description ofMoll and Janssonius (1906) was based on incorrectly
identified material.
There is no description of the wood anatomyof Veeresia. According to our observation on three
specimens (MADw-42819, 42820, 44043) of Veeresia clarkii from Mexico, it shows more distinct ring
porosity than the above mentioned Reevesia species
and has latewood vessels in radial multiples of 2–3,
helical thickenings on vessel walls, tile cells in rays
and numerous crystals in both axial parenchyma andrays (Table 3). Although there are some differencesof wood anatomy between Veeresia and Reevesia,
Veeresia is quite similar in gross anatomical features
to Reevesia.
In conclusion, the wood structure of Reevesia including Veeresia is quite uniform, i.e., (1) the wood
is mostly semi-ring or rarely ring porous, (2) latewood vessels are usually in clusters or in multiples,(3) helical thickenings are distinct on latewood vessel
walls, and (4) tile cells are present (Table 3). Thus it

is apparent that Wataria does not belong to Reevesia
or to Veeresia, because Wataria has thick-walled solitary latewood vessels and lacks of helical thickenings on the vessel walls. Other differences are (1)Wataria has thin-walled tyloses in wide vessels, while
Reevesia has gum-like deposits; and (2) Wataria hasmuch shorter vessel elements and axial parenchyma
strands than Reevesia and Veeresia (Table 3).
3.2. Comparison within the Malvales
The most outstanding feature of Wataria is thepresence of tile cells in rays. Species with tile cells
are restricted to species in four families of Malvales: Sterculiaceae, Tiliaceae, Bombacaceae andMalvaceae (Table 4). In the Sterculiaceae, species

Table 4
Tile cells occurrence among extant genera
Durio type
Sterculiaceae
Reevesia a
Triplochiton a
Guazuma a
Leptonychia
Scaphopetalum
Kleinhovia
Tiliaceae
Microcos
Mortoniodendron a
ColonaLueheaLueheopsis
MolliaVinticena
Bombacaceae
Durio
BoshiaNeesiaCoelosstegia
Malvaceae
KydaiaIntermediate

Pterospermum type

Reevesia a
Triplochiton a
Guazuma a

Reevesia a
Triplochiton a
Pterospermum
Melochia

Grewia a

Grewia a
Mortoniodendron a
Duboscia
Trichospermum

HampeaOchroma

Malvaviscus
Montezuma
Pavonia
Urena

Drawn based on Chattaway (1933), Metcalfe and Chalk (1950),
Manchester and Miller (1978), Carlquist (1988) and IAWA Committee (1989).
a Genera with different types of tile cells within the genus.

K. Terada, M. Suzuki / Review of Palaeobotany and Palynology 103 (1998) 235–251

with tile cells are restricted to four tribes, Theobromaeae, Helictereae, Hermannieae and Mansonieae,
in the Tiliaceae to the tribes Grewieae and Tilieae,
in the Bombacaceae to the tribes Durioneae and Matisieae, and in the Malvaceae, to the tribes Kydieae
and Ureneae. In these four families, tile cells are
known in 29 genera of ten tribes in total (Table 4).
In these genera with tile cells, no species has thedistinctly ring porosity. The only species in the Malvales that has such distinct ring porosity as Wataria
is Firmiana simplex (L.) Wight in Japan, Taiwan and
China, but it does not have tile cells.
Chattaway (1933) recognized two extreme typesof tile cells, the Durio and Pterospermum types, although there is intermediate between the two types.
The most obvious difference between the two types
is the height of tile cells. The tile cells of the Durio type have the same height as the procumbent
cells and the tile cells of the Pterospermum type
are higher than the procumbent cells. Most genera
have either of the two types of tile cells. In theBombacaceae and the Malvaceae, the tile cell type isthe same within the tribes, e.g. in the Bombacaceae
the Durio type is restricted to the tribe Durioneae
and the Pterospermum type to the tribe Matisieae.
However, three genera of the Sterculiaceae and two
genera of the Tiliaceae have intermediate tile cells
and=or both types (Table 4). Reevesia has both types
and intermediate one, and Guazuma of the Sterculiaceae has the Durio type and also intermediateone(Chattaway, 1933). Triplochiton of the Sterculiaceae is considered as Durio type by Chattaway(1933), but Metcalfe and Chalk (1950) and Manchester (1979) regarded it as intermediate while Carlquist (1988) considered it as Pterospermum type(Table 4). Grewia of Tiliaceae has either the intermediate or the Pterospermum type (Chattaway, 1933;
IAWA Committee, 1989). Mortoniodendron of the
Tiliaceae also has either of the two types (Carlquist,
1988). As Wataria has both the intermediate and
the Pterospermum type (Table 2), Wataria may haveaffinities with the above mentioned genera.
Axial parenchyma is an important feature inassessing taxonomic relationships among dicotyledonous woods (Metcalfe and Chalk, 1983). Axial
parenchyma of Wataria is apotracheal banded andparatracheal vasicentric. Apotracheal parenchyma isin uni- or biseriate tangential bands alternating with247

1–3 rows of fibers. Paratracheal parenchyma is abundant and in vasicentric 1–3-seriate sheaths around thevessels. The distribution pattern of axial parenchyma
is usually constant throughout genera in Sterculiaceae(Chattaway, 1932) as well as among dicotyledonous
woods (Metcalfe and Chalk, 1983). Among the 29
genera with the tile cells (Table 4), only Triplochiton has similar parenchyma to Wataria, i.e., apotracheal banded of uni- or biseriate and paratracheal vasicentric 1–3-seriate sheaths (Plate III, 4). Therefore
Triplochiton may have affinity with Wataria becauseof the similarity of axial parenchyma.
Triplochiton is an endemic genus in tropical
Africa. There are two species, Triplochiton scleroxylon Schumann in western Africa (Hutchinson
and Dalziel, 1958), and Triplochiton zambesiacus
Milne-Redhead in southeastern Africa, Rhodesia(Wild, 1963). Although Schumann (1901) originallyproposed Triplochiton as a member of a new family of the Malvales, the genus is now generally
regarded as a member of the Sterculiaceae (Hutchinson, 1967; Takhtajan, 1983). Hutchinson (1967) regarded Triplochiton Schumann as belonging to thetribe Tarrietieae, along with Heritiera Aiton, Hildegardia Schott et Endlicher and Mansonia Drummond
ex Prain, but Edlin (1935) regarded Triplochiton tobelong to the tribe Mansonieae, along with Mansonia. In addition to the similarity in axial parenchyma,
Triplochiton resembles Wataria in the following features (Table 3; Plate III, 4–6): (1) narrow latewood
vessels are mostly solitary and thick-walled; (2) helical thickenings are absent on vessel walls; (3)thin-walled tyloses occur in wide vessels; and (4)tile cells are intermediate between the Durio andPterospermum types. However, there are differences
between Wataria and Triplochiton as follows (Table 3; Plate III, 4–6): (1) Wataria is distinctly ring
porous while Triplochiton is semi-ring to diffuse
porous; (2) Wataria has 2–10 cells in a parenchyma
strand, while Triplochiton has only 2–4 cells; and (3)Wataria has no crystals in axial parenchyma and few
crystals in the rays, while Triplochiton has abundant
crystals in both axial and rays parenchyma. Therefore, although Wataria shares more characterization
with Triplochiton than any other member of the Malvales, it is distinguishable as an extinct genus of
Sterculiaceae.

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K. Terada, M. Suzuki / Review of Palaeobotany and Palynology 103 (1998) 235–251

3.3. Comparison with other fossil wood records
According to the Fossil Dicotyledonous Wood
Database (Wheeler, 1991), eleven species of fossil
woods where reliable tile cells were reported: Coloxylon dechampsii Lemoigne, 1978, Pterospermoxylon
kutchensis Awasthi et al., 1980, Triplochitioxylon oregonensis Manchester, 1979 and Chattawaya paliformis Manchester, 1980 of the Sterculiaceae, andseven species of Grewioxylon of the iliaceae; G. intertrappea Shallom, 1963, G. malhurzariense Prakashet Dayal, 1963, G. indicum Prakash et Dayal, 1964,
G. sp. cf. malhurzariense Lakhanpal et al., 1976,
G. fontanesii Vozenin-Serra, 1981, G. neumaieri
Selmeier, 1985, and G. ortenburgense Selmeier, 1985.
Among the above 11 species, Coloxylon dechampsii
should be excluded, cells because Lemoigne (1978)
mentioned in the original description that it has nottile cells but sheath cells. Pterospermoxylon kutchensis from the Pliocene of western India is quite different from Wataria as it is diffuse porous woodand has scanty paratracheal parenchyma and apotracheal diffuse and diffuse-in-aggregates parenchyma.
All seven species of Grewioxylon differ from Wataria
because they are diffuse porous woods and have indistinct and=or scanty apotracheal parenchyma. Chattawaya paliformis from the Middle Eocene of Oregon, USA, differs from Wataria as it has 4–14-seriate
wide multiseriate rays, shows prominent feature oflarge and irregular-shaped tile cells, and lacks noticeable storied structure. Triplochitioxylon oregonensis
from the Middle Eocene of Oregon, USA, resembles extant Triplochiton (Manchester, 1979). Triplochitioxylon oregonensis is also similar to Wataria as
well as to Triplochiton in these anatomical features,e.g.: (1) distribution pattern of axial parenchyma; (2)mostly solitary and thick-walled vessels in latewood;(3) lack of helical thickenings on vessel walls; (4)thin-walled tyloses in wide vessels; and (5) intermediate tile cells (Table 3). However, Triplochitioxylon
oregonensis differs from Wataria in these features:(1) diffuse porosity; (2) small number of cells in aparenchyma strand; and (3) occurrence of crystals inaxial parenchyma and rays. Furthermore vessel elements and axial parenchyma strands of Triplochitioxylon oregonensis are much longer than in Wataria(Table 3). Therefore Wataria does not belong to
Triplochitioxylon.

We conclude that Wataria is an extinct genus of
the Sterculiaceae but with likely affinity with extant
Triplochiton and fossil Triplochitioxylon.
3.4. Variation of wood structure in Wataria
All Wataria fossil woods examined are quite similar in gross anatomical features, but there is considerable variation in some quantitative characters andin a few qualitative ones (Table 2).
Tangential diameter of vessels at the beginningof annual rings is fairly stable within a species
and often employed as a diagnostic characters. The
mean tangential diameters of earlywood vessels of
Wataria miocenica are 257–287 µm, those of W.
parvipora are 150–176 µm, and of W. oligocenica
are 210 µm (Table 2). Earlywood vessel diameters are significantly larger in Wataria miocenica
than in W. parvipora and W. oligocenica (Fig. 2).
The number of layers of wide vessels in earlywood
also differs Wataria miocenica and W. parvipora
have 1 to 3 layers of wide vessels, while W.
oligocenica has 3 to 5 layers (Table 2). Therefore
Wataria oligocenica differs from W. miocenica in
its narrower earlywood vessels and more layers ofwide vessels in earlywood, as Suzuki (1976) originally described. Wataria parvipora differs from W.
miocenica in its narrower earlywood vessels, andfrom W. oligocenica in fewer layers of wide vessels
in earlywood (Table 2).
Height and width of multiseriate rays are variableboth interspecifically and intraspecifically in extant
Sterculiaceae (Chattaway, 1937). Ray height and ray
width are much altered in most species during the
ontogeny of the xylem cylinder (Carlquist, 1988).
Ray size of Wataria species is highly variable (Table 2), and we did not find any diagnostic value forthese characters.
Among eight specimens of Wataria examined,
two specimens, no. 53219 of W. miocenica and no.
59044 of W. parvipora, have Pterospermum-type tile
cells, an other six specimens are intermediate between Durio and Pterospermum types. As discussed
previously, there are some extant genera and species
that have two types of tile cells and=or intermediatesof them. Therefore, it is not strange that there are one
extreme type and intermediate one within the speciesof Wataria.

K. Terada, M. Suzuki / Review of Palaeobotany and Palynology 103 (1998) 235–251

249

Fig. 2. Size variation of wide vessels in earlywood on Wataria woods, showing mean and standard deviation of diameters.

3.5. Fossil wood of Reevesia
Among ‘Reevesia’ fossil woods that have distinct
ring porosity and tile cells, one specimen, no. 59019,
appears similar to extant Reevesia and Veeresia andis not assigned to Wataria. This specimen has latewood vessels in clusters and helical thickenings on
vessel walls (Table 2; Plate II, 2, 4). These features
agree with those of extant Reevesia and Veeresia as
mentioned previously. In addition, (1) no. 59019 has
longer vessel elements and axial parenchyma strands
than Wataria species, but has similar length to thatof Reevesia and Veeresia, (2) the number of cells
in a parenchyma strand of no. 59019, Reevesia and
Veeresia is 2–4 cells, while in Wataria it is 2–10
cells, and (3) Wataria has tyloses while no. 59019
and Reevesia have gum-like deposits in wide vessels(Table 3).
However, no. 59019 differs from extant Reevesia
species by being distinctly ring porous and having more layers, 2–5, of wide vessels in earlywood. Veeresia clarkii differs from no. 59019 as
follows: (1) vessel diameter of no. 59019 decreases
abruptly from earlywood to latewood while that of
V. clarkii decreases gradually; (2) latewood vesselsof no. 59019 are in clusters of 5–20, which are

distributed in ulmiform or tangential bands, while
those of V. clarkii are in radial multiples of only
2–3 vessels; (3) no. 59019 does not have crystals inaxial parenchyma or in ray cells, while V. clarkii has
crystals abundantly in axial parenchyma and in ray
cells (Table 3).
Therefore the present fossil is regarded as amember of Reevesia, but differs from any extant
Reevesia and Veeresia species examined, and isnamed Reevesia japonoxyla. Distinct ring porosity
shown in Reevesia japonoxyla may be regarded as aphysiognomic adaptation to distinct seasonal change
in the Miocene age (Suzuki and Watari, 1994).
Although there are many fossil pollen records ofReevesia or its allies from Paleocene to Pliocene ofEurope (Bertoldi et al., 1994), this is the first reliable
report of fossil wood of Reevesia.

Acknowledgements
We would like to express our thanks to Mr. K.
Shikano, Gifu Museum, for his help of collecting
fossil woods and his advice for the stratigraphyof the Nakamura and Hachiya Formations, to Professor H. Ohba, the University of Tokyo (TI), for250

K. Terada, M. Suzuki / Review of Palaeobotany and Palynology 103 (1998) 235–251

his courtesy to loan us type specimens of Reevesia
miocenica Watari, to Dr. S.R. Manchester, University
of Florida, for his courtesy to loan us type specimensof Triplochitioxylon oregonensis Manchester. Thanks
are also due to Professor J. Tsutsumi and Dr. K.
Takata, Kyushu University, Fukuoka (FU), to Drs.
P. Gasson, P. Rudall and D.F. Cutler, Royal Botanic
Gardens, Kew, England (Kw), to Dr. P. Baas, Rijksherbarium, Leiden, Netherlands (Lw), to Dr. R.B.
Miller, U.S. Forest Products Laboratory, Madison,
USA (MADw, SJRw), to Dr. H. Beeckman, Royal
Museum for Central Africa, Belgium (Tw), to Drs.
T. Fujii, S. Noshiro and Miss Y. Kondo, Forestry
and Forest Products Research Institute, Tsukuba(TWTw), to Dr. C.I. Peng, Academia Sinica, Taiwan, for their courtesy to loan us wood sample ofMalvales; to Mr. M. Shishido, Tohoku University, for
his advice of making thin sections. Special thanks to
Dr. E.A. Wheeler, North Carolina University, and
Dr. S.R. Manchester for their critical comments on
our manuscript and their useful suggestions. This
study was partly supported by a Grant-in-Aid for Encouragement of the JSPS Fellowships for Japanese
Junior Scientists (to K. Terada) and Fujiwara Natural
History Foundation (to K. Terada).

References
Awasthi, N., Guleria, J.S., Lakhanpal, R.N., 1980. A dicotyledonous wood from the Pliocene bed of Mothala, District
Kutch, Western India. Palaeobotanist 26, 199–205.
Backer, C.A., Bakhuizen van den Brink, Jr., R.C., 1963. Flora ofJava, vol. 1. Noordhoff, Groningen, 648 pp.
Bertoldi, R., Binotti, A., Castello, M.F., 1994. Reevesia and Itea
in the pollen flora of the upper Neogene continental deposit atSarzana (lower Magra Valley, northern Italy). Rev. Palaeobot.
Palynol. 80, 159–172.
Carlquist, S., 1988. Comparative Wood Anatomy. Springer,
Berlin, 436 pp.
Chattaway, M.M., 1932. The wood of the Sterculiaceae I. Specialisation of the vertical wood parenchyma within the subfamily Sterculieae. New Phytol. 31, 119–132.
Chattaway, M.M., 1933. Tile-cells in the rays of the Malvales.
New Phytol. 32, 261–273.
Chattaway, M.M., 1937. The wood anatomy of the family Sterculiaceae. Philos. Trans. R. Soc. London B 228, 313–366.
Cheng, J.-Q., Yang, J.J., Liu, P., 1992. Anatomy and Propertiesof Chinese woods. China Forestry Publ. House, Beijing, 621
pp. (in Chinese).
Edlin, H.L., 1935. A critical revision of certain taxonomic groupsof the Malvales. New Phytol. 34, 1–20 and 122–143.

Greuter, W. et al. (Eds.), 1994. International Code of Botanical Nomenclature (Tokyo Code). Reg. Veg. Vol. 131. Koeltz
Scientific Books, Germany, 389 pp.
Hsue, H., 1984. Sterculiaceae. In: Feng, K. (Ed.), Flora Republicae Popularis Sinicae, vol 49=2. Academic Press, Beijing, pp.
114–189 (in Chinese).
Hutchinson, J., 1967. Family Sterculiaceae. In: The Genera of
Flowering Plants, vol. 2. Oxford Univ. Press, London, pp.
497–522.
Hutchinson, J., Dalziel, J.M., 1958. Flora of West Tropical
Africa, vol. I, part 2. Crown Agents, London, 828 pp.
IAWA Committee 1989. IAWA list of microscopic features for
hardwood identification. In: Wheeler, E.A., Baas, P., Gasson,
P.E. (Eds.), IAWA Bull. 10, 219–332.
Kanehira, R., 1921. Anatomical features and identification of
Formosan Woods with critical remarks from the climatic pointof view. Bur. Prod. Ind. Gov. Formosa, Taihoku, 317 pp.
Lakhanpal, R.N., Prakash, U., Bande, M.B., 1976. Fossil dicotyledonous woods from the Deccan Intertrappean beds ofMandla Distinct in Madhya Pradesh. Palaeobotanist 25, 190–
204.
Lemoigne, Y., 1978. Flores Tertiares de la Haute Vallée de
I’Omo (Ethiopie). Palaeontographica B 165, 89–157.
Li, H., Lo, H., 1993. Sterculiaceae. In: Editional Committee ofFlora of Taiwan (Ed.), Flora of Taiwan, 2 ed., vol. 3. Taipai,
pp. 756–736.
Long, D.G., Rae, S.J., 1991. Sterculiaceae. In: Grierson, A.J.C.,
Long, D.G. (Eds.), Flora of Bhutan, vol. 2=1. Royal Botanical
Garden, Edinburgh, pp. 195–208.
Manchester, S.R., 1979. Triplochitioxylon (Sterculiaceae): A new
genus of wood from the Eocene of Oregon and its bearing on
xylem evolution in the extant genus Triplochiton. Am. J. Bot.
66, 699–708.
Manchester, S.R., 1980. Chattawaya (Sterculiaceae): A new
genus of wood from the Eocene of Oregon and its implication for xylem evolution of the extant genus Pterospermum.
Am. J. Bot. 67, 59–67.
Manchester, S.R., Miller, R.B., 1978. Tile cells and their occurrence in Malvalean fossil woods. IAWA Bull. 2–3, 23–28.
Masters, M.T. 1875. Sterculiaceae In: Hooker, J.D. (Ed.), Floraof British India, vol. 1. L. Reeve, England, pp. 353–379.
Metcalfe, C.R., Chalk, L., 1950. Anatomy of the Dicotylodons,
vol. 1. Clarendon Press, Oxford, 724 pp.
Metcalfe, C.R., Chalk, L., 1983. Anatomy of the Dicotylodons,
2 ed., vol. 2. Wood Structure and Conclusion of the General
Introduction. Clarendon Press. Oxford, 297 pp.
Moll, J.W., Janssonius, H.H., 1906. Mikrographie des Holzes derauf Java vorkommenden Baumarten. E.J. Brill, Leiden, 568
pp.
Monachino, J., 1940. A new genus and species of Sterculiaceae.
Bull. Torr. Bot. Club 67, 621–622.
Nomura, T., 1986. Preliminary report of the Miocene Hachiya
Formation and its K-Ar ages. J. Geol. Soc. Jpn. 92, 75–78 (inJapanese).
Nomura, T., 1992. Stratigraphy of the Miocene Hachiya Formation, Gifu Prefecture, Central Japan; An instance of the
volcanism took place in the Setouchi Geologic Province dur-

K. Terada, M. Suzuki / Review of Palaeobotany and Palynology 103 (1998) 235–251
ing lower Miocene. Bull. Mizunami Fossil Mus. 19, 75–101(in Japanese).
Prakash, U., Dayal, R., 1963. Fossil wood resembling Grewia
from the Deccan Intertrappean beds of Muhurzari near Nagur,
India. Curr. Sci. 32, 315–316.
Prakash, U., Dayal, R., 1964. Fossil wood of Grewia from theIntertrappean Series, India. Palaeobotanist 13, 17–24.
Sakai, H., 1994. Koinoura pyoclastic deposits in the Lower
Oligocene Tsuyazaki Formation, North Kyushu. J. Geol. Soc.
Jpn. 100, 692–708, in Japanese.
Sato, H., Yamaji, A., Tsuchiya, N., Ohguchi, T., Taguchi, K.,
1989. Shonai district. In: Editorial Committee of Tohoku (Ed.),
Regional Geology of Japan Part 2 Tohoku. Kyoritsu Shuppan,
Tokyo, pp. 169–175 (in Japanese).
Schumann, K., 1901. Eine neue Familie der Malvales. Bot. Jahrb.
Syst. Pflanzengesch. Pflanzengeogr. 28, 330–331.
Selmeier, A., 1985. Jungtertiäre Kieselhölzer (Bombacaceae,
Tiliaceae, Meliaceae) aus dem Ortenburger Schotter vonRausheröd (Niederbayern). Münchner Geowiss. Abh., Reihe
A, Geol. Paläntol. 6, 89–140.
Shallom, L.J.A., 1963. Fossil dicotyledonous wood with tile
cells, from the Deccan Intertrappean beds of Mahurzari. J.
Indian Bot. Soc. 42, 170–176.
Shikano, K., 1995. Fossil forests of Nakamura Formation. In:
Minokamo city Educational Boards (Ed.), Strata and fossilsof Nakamura Formation in Minokamo Basin — Reports forfossil footprints of mammals and fossil forests. pp. 30–45 (inJapanese).
Solereder, H., 1908. Systematic Anatomy of the Dicotyledons,
vol. 1. Ajay Book Service, New Delhi, 644 pp.
Solheim, S.L., 1987. Reevesia (Sterculiaceae, Helictereae): systematics and biogeography of an Asian–eastern America disjunction. Am. J. Bot. 74, 745 (abstract).
Suzuki, M., 1976. Some fossil woods from the Palaeogene ofnorthern Kyushu. Bot. Mag. Tokyo 89, 59–71.

251

Suzuki, M., Watari, S., 1994. Fossil wood flora of the lower
Miocene Nawamata Formation of Monzen, Noto Peninsula,central Japan. J. Plant Res. 107, 63–76.
Stainton, A., 1988. Flowers of the Himalaya, A Supplement.
Oxford Univ. Press, London, 86 pp.
Takhtajan, A., 1983. The systematic arrangement of dicotyledonous families. In: Metcalfe, C.R., Chalk, L. (Eds.), Anatomyof the Dicotylodons, 2 ed., vol. 2. Clarendon Press, Oxford,
pp. 180–201.
Tang, Y., 1932. Identification of some important hardwoods ofsouth China by their gross structures I. Bull. Fan. Mem. Inst.
Biol. 3, 256–338.
Tang, Y., 1936. Timbers of China. Shanghai, 680 pp. (in Chinese).
Terada, K., 1995. Fossil woods of the fossil forests. In: Minokamo City Educational Boards (Ed.), Strata and fossils ofNakamura Formation in Minokamo Basin — Reports for fossil footprints of mammals and fossil forests. pp. 46–47 (inJapanese).
Vozenin-Serra, C., 1981. Les structures ligneuses néogénes duplateau de Di Linh (Sud-Viêtnam). Palaeontographica B 177,
136–161.
Watari, S., 1952. Dicotyledonous woods from the Miocene along
the Japan-sea side of Honshu. J. Fac. Sci. Univ. Tokyo Sect.
III (Bot.) 6, 97–134.
Wheeler, E., 1991. Fossil dicotyledonous wood database, C.A.
LaPasha (programmed), Computer Assisted Wood Identification, ver. 1.1. North Carolina State Univ.
Wild, H., 1963. Sterculiaceae. In: Exell, A.W., Fernandes, A.,
Wild, H. (Eds.), Flora Zambesiaca. Crown Agents, London,
pp. 517–564.
Yamaji, A., 1989. Geology of Atsumi area and Early Miocene
rifting in the Uetsu district, northern Japan. Mem. Geol. Soc.
Jpn. 32, 305–320, in Japanese.