A late Eocene wood assemblage from the Crooked River Basin, Oregon, USA
Author
Wheeler, Elisabeth A.
Author
Manchester, Steven R.
Author
Baas, Pieter
text
PaleoBios
2023
2023-11-01
40
14
1
55
http://dx.doi.org/10.5070/p9401462457
journal article
10.5070/P9401462457
0031-0298
10913330
FAGAROXYLON
VAN DER BURGH, 1964
CF.
FAGAROXYLON
SP.
FIG. 18A–I
Description—
Growth rings present, marked by radially narrow fibers, (probably also marginal parenchyma) (
Fig 18A, B
).
Wood diffuse-porous (
Fig 18A, B
). Vessels solitary (14%) and in radial multiples mostly 2–4 (-7), clusters rare; mean tangential diameter 63 (
SD
=15) µm, range 28–90 µm; 36–42–50 vessels/mm
2
. Perforation plates simple (
Fig. 18E, F
); intervessel pits crowded alternate (
Fig. 18C
), horizontal diameters small to medium 5–8.5 µm. Vessel-ray parenchyma similar to intervessel pits (
Fig. 18D
). Vessel element lengths 235–277–353 µm (n=12); vessel element end walls obliquely inclined. Helical sculpturing (grooves) in some narrow vessel elements (
Fig. 18E, F
). Gum-like deposits common.
Fibers thin-walled, non-septate, pitting not observed (
Fig. 18E–H
).
Axial parenchyma not common, scanty paratracheal (hard to distinguish from thin-walled fibers) and probably also marginal (
Fig. 18H
).
Rays 1–4 (-5)-seriate (
Fig. 17E, G
). Average ray height 497 (
SD
=203) µm, range 181–1062 µm. Homocellular, composed of procumbent cells (
Fig. 18D, F, I
), 6–9 per mm, non-storied.
Crystals common in chambered axial cells (fibers), strands sometimes adjacent to rays, one crystal per chamber, numerous (10+) crystalliferous chambers per strand (
Fig. 18E, G, I
).
Specimen—
UF
278-84907, estimated maximum diameter
10 cm
.
Occurrence—
Dietz Hill (
UF
278).
Comparisons with extant woods—
We searched the InsideWood database multiple times. One search used diffuse porosity (5p), absence of vessel patterning and exclusively solitary vessels (6-9a), exclusively simple perforation plates (13p 14a), alternate intervessel pitting that is neither minute nor large (22p, 24a, 27a), vessel-ray parenchyma pits similar to intervessel pits (30p), average vessel diameter less than 200 µm (43a), vessels per mm
2
between five and 100 (46a, 50a), non-septate fibers with simple to minutely bordered pits (61p 66p), axial parenchyma scanty paratracheal and marginal, but not accompanied by more obvious axial parenchyma (78p, 89p, 80-88a), homocellular rays not exclusively uniseriate or greater than 10-seriate (96a, 99a, 104p), storied structure absent (118–122a), oil/mucilage cells and intercellular canals absent (124–130a), prismatic crystals present in fibers (136p 143p). When not allowing any mismatches, this search yielded only
Meliaceae
(
Capuronianthus
J
.-
F
.Leroy, 1958;
Quivisianthe
Baill., 1893
, from
Madagascar
(
Rutaceae
);
Aeglopsis
Swingle, 1912
;
Afraegle
(Swingle) Engl., 1915
, from Africa; and
Sapindaceae
(one Asian
Acer
L
., 1753).
When one mismatch was allowed, results again included
Meliaceae
(three species),
Rutaceae
(12 species),
Sapindaceae
(19 species, including 11 of
Acer
). Other families retrieved were
Combretaceae R.Br.(1810)
(three
Terminalia
L
. (1767) species), which differ in having crystals in idioblasts or in having more abundant axial parenchyma),
Lamiaceae
(one
Vitex
species
with mostly septate fibers and crystals of other shapes),
Fabaceae
(five species, none have crystals in fibers, although there is the possibility crystals in fibers might have been interpreted as crystals in chambered axial parenchyma as reported for four of them).
Figure 18
. Caption on pg. 40.
Figure 18
.
Sapindales
.
Fagaroxylon
sp.
, UF 278-84907.
A, B
. Diffuse-porous wood; vessels solitary and in radial multiples, thin-walled fibers, growth ring boundary marked by radially narrow fibers, and axial parenchyma, TS.
C.
Crowded alternate intervessel pitting, TLS.
D.
Vessel-ray parenchyma pits (VRP) similar to intervessel pits, procumbent ray cells, RLS.
E.
Multiseriate rays; crystalliferous strand (C); crowded alternate intervessel pitting, vessel element end walls (E), helical thickenings (HT) in narrow vessel elements, TLS.
F
. Ray composed of procumbent cells; narrow vessel element with simple perforation plate (PP) and helical thickenings (HT), RLS.
G
. Rays 1‒4-seriate
,
crystalliferous strands adjacent to rays (C), TLS.
H
. Marginal axial parenchyma (AP); simple perforation plate (PP), RLS.
I
. Crystals in fibers, RLS. Scale bars=500 µm in A; 100 µm in B, G, 50 µm in E, F, H, I; 20 µm in C, D.
We tried other searches using slightly different combinations of features, and all of them consistently returned members of the Sapindalean families
Meliaceae
,
Rutaceae
, and
Sapindaceae
. We are confident that this wood belongs to the
Sapindales
and are inclined to think that it is most likely related to
Sapindaceae
, which is noted as the Sapindalean family most commonly having crystals in fibers (
Pace et al. 2022
). It resembles
Acer
most closely, but all present-day
Acer
consistently have helical thickenings in vessel elements, so this wood is not assigned to
Acer
.
The fruit and seed flora of Dietz Hill has an endocarp that might have affinities with
Acer
(
Manchester
and McIntosh 2007
).
Dipteronia
Oliv. (1889)
is closely related to
Acer
and fruits of it occur in the Teater Road shales that we infer to be of similar age to the Dietz Hill (
UF
278) locality (
McClain and
Manchester
2001
).
Itoh et al. (2022)
described
Dipteronia sinensis
Oliv. (1889)
(under the name
Acer dielsii
H. Lév. (1912)
. That description differs somewhat from the descriptions and observations of
D. dyeriana
A. Henry (1903)
and
D. sinensis
made by Peter Gasson (Royal Botanic Gardens, Kew) and Michael Wiemann (
Center
for Wood Anatomy Research, Madison) and available on InsideWood (insidewood.lib.ncsu.edu).
Itoh et al. (2022)
report helical thickenings in narrow vessel elements and rays commonly more than
1 mm
high, features not seen by Gasson or Wiemann. Crystals are not reported for
Dipteronia
.
Comparisons with fossil woods—
Excluding fossil wood descriptions with most features coded as unknown, the results of our searches of InsideWood’s fossil wood database returned two species of
Fagaroxylon
Van der Burgh (1964)
from the Miocene of Europe.
Fagaroxylon
is a genus created for woods with features of both
Fagara
L
. (1759), which is now subsumed into
Zanthoxylum
L
. (1753). Its diagnosis (
Van
der Burgh 1964
) and later descriptions of
Fagaroxylon bavaricum
(
Selmeier 1975
)
and
Fa. limburgense
(
Van der Burgh 1973
)
are remarkably similar to this Dietz Hill (
UF
278) wood. Similarities include most vessel features, axial parenchyma distribution, chambered crystals, ray width and cellular composition, and even the occasional occurrence of helical vessel wall thickenings. Vessel frequency is lower in
Fa. limburgense
, but this likely is because in the past vessel multiples (groups) were counted as one unit (
Wheeler 1986
).
Fagaroxylon limburgense
,
the
type
species, is reported to have fibers with distinctly bordered pits (diameter 3 µm) in their radial walls, a feature we did not observe in
UF
278-84907. Because variations in preservation and the effects thereof can make it difficult to determine whether fiber pits are>3 µm across, we choose to assign
UF
278-84907 to
Fagaroxylon
.
Fagaroxylon
’s combination of features occurs in other genera of
Rutaceae
, as well as the
Meliaceae
and
Sapindaceae
and is not unique to
Zanthoxylum
. It should be noted that seeds that correspond closely to the extant genus
Zanthoxylum
occur at Dietz Hill (
UF
278).
Other Sapindalean fruit and seed remains in the Teater Road shales include
Koelreuteria
Laxm (1771)
(
Wang et al 2013
),
Dipteronia
(
McClain and
Manchester
2001
), and several species of
Acer
leaves and fruits (
Wolfe and Tanai, 1987
, indicated as “Sheep Rock Creek”).