The Wood of Rellimia from the Middle Devonian of New York

Author(s): Joanne M. Dannenhoffer and Patricia M. Bonamo

Source: International Journal of Plant Sciences, Vol. 164, No. 3 (May 2003), pp. 429-441
Published by: The University of Chicago Press
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Int. J. Plant Sci. 164(3):429–441. 2003.
䉷 2003 by The University of Chicago. All rights reserved.
1058-5893/2003/16403-0010$15.00

THE WOOD OF RELLIMIA FROM THE MIDDLE DEVONIAN OF NEW YORK

Joanne M. Dannenhoffer1 and Patricia M. Bonamo
Department of Biology, Central Michigan University, Mount Pleasant, Michigan 48859, U.S.A.; and Department of
Biological Sciences, Binghamton University, Binghamton, New York 13902, U.S.A.

Rellimia is one of the best known of the progymnosperms, although the anatomy of its wood and tree
structure has yet to be fully described. The occurrence of Rellimia in Devonian strata makes it one of the
earliest plants known to have developed wood. This study examines pyrite permineralizations from the Catskill
region of New York and elucidates anatomical details of the secondary xylem using light and scanning electron
microscopy. The wood is pycnoxylic with tall, narrow rays that are typically uniseriate, although biseriate
and multiseriate rays can be present. Ray cells are slightly rectangular, and the rays are homocellular. All cells
appear to have thin walls and appear to be ray parenchyma. Tracheids have crowded, circular-bordered pits
on all walls, including the long, tapering end walls. Bordered pits are in alternate spirals across the face walls
and are often hexagonal in outline. Fractures through the bordered pit pair reveal details of the wall and
pyrite casts that fill the pit canal and pit chamber during fossilization. Bordered pit pairs have crossed pit
apertures, with the pit canal being longer than the shallow pit chamber. Preserved pit membranes and secondary
walls show microfibrillar arrangement. Anatomical characteristics of the known wood of progymnosperm
taxa are compared and discussed in light of the group’s position in lignophyte evolution.

Keywords: progymnosperms, Aneurophytales, secondary xylem, Givetian.

Introduction preservation of the New York material allows many more of

The progymnosperms are an extinct group of plants that
occupy an important place in the phylogeny of lignophytes
(Nixon et al. 1994; Rothwell and Serbet 1994). The most
completely known genus is Archaeopteris, an abundant com-
ponent of worldwide Upper Devonian floras. It was the dis-
covery of its combined morphological and anatomical char-
acters, pteridophytic reproduction and gymnospermous
anatomy, that led to the formation of this extinct group (Beck
1960a, 1960b). Archaeopteris is characterized by development
of a secondary anatomy that represents the first evidence for
a plant body that resembles that of modern trees (Meyer-
Berthaud et al. 1999, 2000). However, in the aneurophytes,
an older Middle to Upper Devonian group of progymno-
sperms, there is the first evidence for a bifacial cambium and
the production of wood (Beck 1957; Scheckler and Banks
1971a; Dannenhoffer and Bonamo 1989).
Within the aneurophyte group, Rellimia is considered well
known by its reproductive morphology but lacking in the de-
scription of its anatomy (Beck and Wight 1988). Branching
systems consist of at least four orders of spirally arranged axes.
The primary xylem is a three-lobed protostele in each axis
order (Bonamo 1977). Secondary xylem surrounds the primary
xylem in the larger axis orders, and the largest of these contain
growth increments within the wood (Dannenhoffer and Bon-
amo 1989). The wood has been briefly described as containing
radially aligned tracheids and tall narrow rays (Kräusel and
Weyland 1938; Dannenhoffer and Bonamo 1989). The pyrite
1
Author for correspondence; e-mail danne1jm@cmich.edu.
Manuscript received August 2002; revised manuscript received January 2003.
the details of the secondary xylem and the tracheids to be
discerned.
Grierson (1976) first used the scanning electron microscope
(SEM) to elucidate the tracheid anatomy of the pyritized fossil
lycopod Leclercqia. Using his work as the foundation, we here
expand on his interpretation of bordered pits by showing the
more complicated anatomy of the tracheids and bordered pit
pairs in Rellimia wood. Previous discussions of the bordered
pits of fossil tracheids use diagrams of sectional views of the
bordered pit pairs combined with face views (Schmid 1967;
Grierson 1976; Beck et al. 1982). The three-dimensional model
presented here is a more complete representation of what can
be seen with the SEM after material has been acid treated. We
describe here the characteristics of the tracheids and secondary
xylem of Rellimia wood, one of the earliest records of sec-
ondary growth from a bifacial cambium.
Material and Methods
The pyritized material of Rellimia was collected from the
construction site of the Blenheim-Gilboa pumped storage
power plant in Blenheim, Schoharie County, New York. The
material belongs to a large group of mostly compression spec-
imens of Rellimia (Bonamo 1977; Dannenhoffer and Bonamo
1989). The morphology and a brief description of the sec-
ondary anatomy of two specimens, one with abundant wood,
have been previously published (Dannenhoffer and Bonamo
1989). Here, we describe the details of the wood anatomy of
the pyritized axes of the N and N + 1 order preserved in spec-
imen 349.1. This material is housed in the paleobotanical col-
lection of Binghamton University (SUNY Binghamton).

429

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 Fig. 1 Light and scanning electron micrographs of Rellimia wood. l p lumen cast, r p ray . Bar p 100 mm in C–E; bar p 50 mm in A;
bar p 10 mm in B, F. A, Light micrograph of a transverse section with three rays all about the same width but of different seriation. The top
ray is multiseriate, the middle ray is uniseriate, and the bottom ray is biseriate. B, SEM micrograph of transverse section shows tracheids and
a ray. Pits are visible on both tracheids and rays cells (arrows), but it is impossible to determine if they are bordered or simple when preserved
this way. The tracheid lumen cast is composed of large pyrite crystals. C, Oblique radial section showing a partly preserved ray and elongate
tracheids. D, SEM micrograph of cellularly preserved rays (arrows) in radial section that may be one single large ray broken during the specimen
preparation. E, Tangential section with a uniseriate ray and a tracheid with pits on both face and sectioned walls (arrows). F, SEM micrograph
of tangential section of ray. Ray cell lumen casts have smooth walls and no evidence of ray cell pits.

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 DANNENHOFFER & BONAMO—FOSSIL WOOD OF RELLIMIA 431

Table 1
Ray and Tracheid Measurements of Rellimia Wood
Number
measured Range Mean SD
Tracheids:
Radial diameter 84 22.8–104.5 60.8 16.1
Tangential diameter 80 28.5–81.7 42.9 9.8
Length 1 … 1520 …
Cell wall thickness 50 3.8–15.2 9.5 2.4
Pit diameter 43 9.5–20.9 14.5 2.5
Rays:
Width (in cells) … 1–3 cells … …
Width 42 13.3–72.2 29.5 11.3
Height (in cells) … 2–20 cells … …
Height 41 161.5–864.5 488.3 186.2
Ray cells:
Length 45 47.5–153.0 87.4 26.0
Height 55 24.7–60.8 40.9 8.7
Width 60 7.6–57.0 22.4 12.7
Width (uniseriate rays) 31 13.3–57.0 28.8 2.9
Width (biseriate, multiseriate rays) 29 7.6–39.9 15.6 7.6
Ray wall thickness 7 1.9–3.8 2.2 0.04
Note. Cells and cell walls were measured as the two contiguous walls of adjacent cells and one lumen pyrite
cast to avoid the difficulty of separating the wall at the middle lamella. Measurements other than width and height
of rays in number of cells are in micrometers.

The fossils collected from Blenheim-Gilboa occur in gray
mudstone, within the Panther Mountain Formation of the
Hamilton Group (Rickard and Fisher 1970). This is locality
2 of the Catskill clastic wedge described by Banks et al. (1985).
The formation belongs to the Tioughniogan Stage, Erian Series
(American Standard), which is equivalent to the Givetian, Mid-
dle Devonian of the European Standard (Rickard 1964). In a
comparison of the in situ spores of Leclercqia from the Blen-
heim locality with dispersed spores from Emsian through lower
Givetian localities, Richardson et al. (1993) suggest the locality
is no older than late Eifelian and is probably lower Givetian.
For light microscopy, materials were prepared by standard
sectioning and grinding techniques, were subsequently ex-
amined with a petrographic microscope, and were photo-
graphed with a Leitz Arisotphot. To examine the details of
wood structure with SEM, split and sectioned specimens were
etched with 10% nitric acid for various times from 30 s to 30
min, following the techniques of Grierson (1976). This etching
technique dissolves the carbonaceous wall and thereby exposes
the pyrite cell casts, although wall fragments often remain.
Etched pieces were neutralized with water or Kodak Photoflo,
air-dried, and mounted on SEM stubs, followed by sputter
coating with 20 nm of gold palladium. Specimens were viewed
and photographed with an ETEC Autoscan B1 at 10–20 kV
with Polaroid P/N 55 film.
Description
Wood Anatomy
The secondary xylem of Rellimia shows the dual wood char-
acteristics of radial alignment of cambial derivatives and the
presence of both axial and radial systems that have a char-
acteristic appearance in transverse, radial, and tangential
planes of section (fig. 1). The wood is compact, with narrow
rays that give it a pycnoxylic or “typically gymnospermous”
appearance. Rays are quite abundant, sometimes occurring as
often as between every one or two files of tracheids (fig. 1A).
Often the rays have no preserved cell walls, but when preserved
they are uniseriate, biseriate, and multiseriate (fig. 1A). Uni-
seriate rays are commonest. The mean width of rays is 29.5
mm (table 1). Biseriate and multiseriate rays are about the same
width or only slightly wider than uniseriate rays, appearing as
if a uniseriate ray has been divided. Rays ranged from two to
more than 20 cells high. The mean ray height is 488.3 mm
(table 1).
Tracheid Structure
Tracheids are circular to rectangular to polygonal in trans-
verse shape and most often have four to six walls (fig. 1A,
1B). They are larger in the radial dimension, with a mean radial
diameter of 60.8 mm compared to a mean tangential diameter
of 42.9 mm (table 1). Tracheids are very long, more than 20
times longer than they are broad (fig. 1C–1E). One tracheid
measured along its full length was more than 1500 mm long
(table 1). The end wall of tracheids can be slightly to very long
and tapering. As it narrows, the number of bordered pits across
the face decreases (fig. 2A). Some tracheids have branched tips
covered with bordered pits (fig. 2B). The branched tracheid
tips and the long tapering end walls suggest apical intrusive
growth during cell development.
Tracheid walls have crowded bordered pits completely cov-
ering the radial and tangential walls (fig. 2C). Pits occur in an
alternate series of two to five across the tracheid (fig. 2C, 2D).
The number of pits across a tracheid face correlates with the
width of the tracheid; wider tracheids have more pits. Pits are
typically circular to elliptical in shape, although sometimes

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 Fig. 2 Details of the structure of tracheids and bordered pit pairs between adjacent tracheids. c p pit -canal cast, h p pit-chamber cast,
i p inner pit aperture, l p lumen cast, m p pit membrane, o p outer pit aperture, r p ray, w p secondary wall. Bar p 100 mm in A, B, D;
bar p 10 mm in C, E–G. A, SEM view of long, tapering end walls of several tracheids (arrows). As the walls narrow, they have fewer rows of
pits. B, Tangential section showing branched tracheid with sectioned bordered pits on the radial walls. C, SEM view of tracheids with pits on
both radial and tangential walls. Pits are crowded on all wall faces. D, Radial section with tracheids in face view showing pits are hexagonal,
crowded, and in alternate series. E, Transverse section with a nonpreserved ray and tracheids with bordered pits. The pit canal and pit-chamber
casts have been preserved with small pyrite crystals. The pit membrane is also preserved between two pit-chamber casts. F, Two adjacent tracheids
show the anatomy of bordered pit pairs. Remnants of secondary wall remain among the pyrite casts of the lumens, pit canals, and pit chambers.
The pit canal is long and has an inner and an outer pit aperture. The pit chambers are rather shallow. G, SEM view of tracheids, showing the
granular nature of the pyrite cast and the smoother texture of preserved cell wall. At places, the tracheid fracture has occurred to leave no
evidence of pits (l), but in other places the fracture reveals appearance 1 pits. These pits show the pyrite pit-canal cast attached to the lumen
cast.

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 DANNENHOFFER & BONAMO—FOSSIL WOOD OF RELLIMIA 433

Fig. 3 Model of the possible fracture planes through two adjacent pyritized tracheids. The three-dimensional model represents the wall and
pyrite casts of two adjoining tracheids. Pyrite is white, the secondary wall is gray, and the compound middle lamella is black. The front of the
model at right shows the sectional view with wall material and pyrite casts of the space system of the cells similar to that shown in fig. 2E, 2F.
The pyrite casts include part of the lumen cast, the pit-canal casts (c), and pit-chamber casts (h). The organic components include the compound
middle lamella (m) and secondary wall (w). The secondary wall around the elongate pit canal has two openings, the inner pit aperture (i) and
the outer pit aperture (o). The model is cut through the two cells much like the tracheids fracture during acid treatment. Nine representations
of the pits, appearances 1–9, are shown, and the appearance depends on the different combinations of pyrite casts and cell wall. If the fracture
is along the pyrite cast of the lower cell, two pit appearances occur: (1) the pit-canal casts attached to the lumen cast of the lower cell, and (2)
the pit-canal and pit-chamber casts attached to the lumen cast of the lower cell. If the fracture is within the secondary wall of the lower cell,
two pit appearances occur. (3) The secondary wall with unfilled pit chambers of the lower cell, pyrite casts of the pit canals may or may not
be visible, and (4) the secondary wall surrounding pyrite-filled pit chambers of lower cell. If the fracture is along the compound middle lamella
(pit membrane) of both cells, three pit appearances occur: (5) the pit membrane, (6) the compound middle lamella and attached pit-chamber
casts of the upper cell, and (7) the compound middle lamella and attached pit-chamber and canal casts of the upper cell. If the fracture is within
the secondary wall of the upper cell, two pit appearances occur: (8) the secondary wall of the upper cell fractured through the pit border resulting
in the pit chamber covered with a layer of secondary wall and (9) the secondary wall of the upper cell with slitlike inner pit apertures, with or
without visible pit-canal casts.

they are so closely appressed they have flattened contacts, giv-
ing the pit a hexagonal shape (fig. 2C, 2D). Pits like these in
Rellimia—bordered, crowded, hexagonal, and alternate—are
said to be of the “Araucarian” type (Greguss 1955). Pits have
a mean diameter along the horizontal axis of 14.5 mm (table
1).
Interpretation of the different appearances of bordered pit
pairs. Tracheids studied with both incident light and scan-
ning electron microscopy show many different appearances of
the bordered pit pairs. These appearances reflect the many
possible fracture planes that could occur through a three-
dimensional bordered pit pair, the presence or absence of pyrite
casts, and the presence or absence of preserved cell wall. The
pyrite next to the wall is much finer grained than the pyrite
within the lumen and often faithfully reproduces the cell spaces
of the bordered pit as pyrite casts (fig. 2E, 2F). In SEM mi-
crographs, the preserved wall (w) can be distinguished from
the granular pyrite (l) of the cast system (fig. 2G).
In order to understand the anatomy of the bordered pit pairs,
a model of the cell walls between two pyritized tracheids shows
all the pit appearances we observed in the secondary xylem of
Rellimia (fig. 3). A bordered pit pair connects two cells that
share common intercellular cement. The secondary wall is not
a complete wall; it has holes and an uneven thickness around
the bordered pits. If the wall is sectioned and the material is
viewed with incident light (fig. 2B, 2E), preserved pit pairs
give the wall a beaded appearance as the bright pyrite casts
alternate with the light-absorbing wall. In material etched with
acid and then viewed with the SEM, the secondary wall is
often dissolved and the bordered pit is seen as pyrite casts of
the space system (fig. 2F).
The space system in these tracheids is filled with pyrite and
consists of the cell lumen, the pit canal (c), and the pit-chamber
(h) casts (figs. 2F, 3). The pit canal and pit chamber are part
of the bordered pit. The pit canal is long so there are two pit
apertures, the inner pit aperture (i) and the outer pit aperture
(o). The inner pit aperture is a hole in the secondary wall
through which the cell lumen is connected to the pit canal.

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 Fig. 4 SEM micrographs of the nine appearances of the bordered pits (1–9), corresponding to those depicted in the model. b p pit border,
c p pit-canal cast, h p pit-chamber cast, l p lumen cast, w p secondary wall. Bar p 10 mm in all except E, where bar p 1 mm. A, Tracheid
end wall showing bordered pits in side view and in face view. All secondary wall is etched away, and the fracture has left pyrite casts of the pit
canal and the pit chambers. In face view, the pits of appearances 1 and 2 are apparent. B, Tracheid whose plane of fracture exposes both
preserved wall and pyrite casts. Pits are of four appearances. At the top right, the fracture shows the lumen cast has attached pit-canal casts,

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 DANNENHOFFER & BONAMO—FOSSIL WOOD OF RELLIMIA 435

The outer pit aperture is a hole in the secondary wall through
which the pit canal is connected to the pit chamber. The pit
border is the portion of the secondary wall that overarches
the pit chamber from inner pit aperture to outer pit aperture
and surrounds the pit canal.
Pyrite filling of the cell space system and the preservation
of wall entombed within these pyrite casts provide many pos-
sible fracture planes through bordered pit pairs of adjacent
tracheids. These fractures yield different appearances of the
bordered pit pair that are depicted in the model as pit ap-
pearances 1–9 and are described below as seen in SEM mi-
crographs (fig. 3; figs. 2G–4I). Often, a fracture through a
Rellimia bordered pit pair occurs at the level of the lumen cast
of one tracheid, with the acid treatment removing all or almost
all of the cell wall material. When this happens, the tracheid
will have one of three appearances depending on where the
fracture occurs in the pyrite cast system (fig. 3). If the fracture
occurs along the interface of the inner pit aperture, between
the lumen cast and the pit-canal cast, the result may be a pyrite
cast without any evidence of pit chambers or canals (figs. 2G,
3). Often, the fracture in the pyritized space system will occur
within the pit canal or between the pit canal and the pit cham-
ber at the outer pit aperture. Then, appearance 1 pits occur
on the lumen cast as slitlike protrusions that are pit canals of
various sizes depending on the fracture plane (figs. 2G, 3; fig.
4A, 4B). When a break occurs between the pit-chamber cast
and the compound middle lamella, all three pyritized portions
of the space system of the tracheid remain intact as pyrite casts.
The pits, appearance 2, are either seen in section or from the
surface of the pit-chamber cast (fig. 3; fig. 4A, 4B).
Many times, a fracture occurs within these same places of
the pyrite cast system in a tracheid, but the acid treatment has
not removed all the secondary wall material. Then, the pits
have a different appearance. In some samples, the pyrite cast
is fractured at the outer pit aperture between the pit canal and
pit chamber, with the pit chamber removed and the secondary
wall remaining. The pit, appearance 3, is viewed along the
surface of the pit border and appears as a depression in the
secondary wall (fig. 3; fig. 4B, 4C). Usually, the pyritized pit-
canal cast (c) can be seen through the pit aperture, but oc-
casionally the cast is missing. When the pit-chamber cast of
the lower cell is not broken and the wall is separated at the
secondary wall–compound middle lamella interface, the tra-
cheid has intact pit chambers surrounded by secondary wall,
for pits of appearance 4 (fig. 3; fig. 4B, 4C, 4F). The pit-
chamber cast when viewed from this side is flat from its contact
with the pit membrane. This view of the pits is often observed
with light microscopy (fig. 2D).
A fracture between two tracheids along their compound
middle lamella and pit-chamber cast is common, but preser-
vation of the compound middle lamella after acid treatment
is rarer (fig. 4D). Sometimes, the compound middle lamella is
so well preserved that the pit membrane is apparent and ap-
pears as pit appearance 5 (figs. 3, 4E). More often, when the
tracheid walls are fractured in this plane, the compound middle
lamella is not visible because pyritized pit-chamber casts of
the adjacent tracheid cover it. If the split in the pyrite is along
the outer pit aperture, it removes the pit-canal casts and leaves
the pit-chamber casts, for pit appearance 6 (figs. 3, 4F). With-
out the secondary wall of the tracheid, the pyrite casts of a
pit chamber viewed from this side have a rounded shape com-
pared with the flat shape when viewed from the other side, as
in appearance 4 pits. At times, the break in the pyrite occurs
at the inner pit aperture. In these instances, the pit-chamber
cast with an attached pit-canal cast remains but all the sec-
ondary wall of this tracheid is removed, for pit appearance 7
(figs. 3, 4G). The pit canal is approximately the full diameter
of the pit chamber.
In less common circumstances, a fracture occurs within the
secondary wall of the tracheid, leaving the pit border but re-
moving the rest of the secondary wall, for pit appearance 8
(figs. 3, 4H). The pit border either overlies the pit-chamber
cast that has been broken from the pit-canal cast or, in some
cases, portions of the pit-canal cast may remain. Here, the pit-
border surface that faces the rest of the secondary wall and
the cell lumen of the tracheid is exposed, as opposed to the
surface that faces the pit chamber (fig. 4H, cf. 8 and 3). More
often, the fracture is along the surface of the secondary wall

appearance 1 pits. Below that, the pit-chamber cast is still attached, but the secondary wall is gone, showing pit appearance 2. At the bottom,
pit-chamber casts have been removed by a fracture at the inner pit aperture, and the pit canal is visible at the aperture surrounded by the
preserved pit border of the secondary wall, pit appearance 3. To the left, preserved wall around the pit-chamber casts reveals appearance 4 pits.
C, Etched tracheids showing pits of appearance 3 and 4. Most of the pits are appearance 3 and show the secondary wall of the pit border
without a pit-chamber cast, but the pit canal shows through the outer pit aperture. Some pits have the pit-chamber cast in part, and others have
the entire pit-chamber cast, pit appearance 4. The secondary wall is apparent between the appearance 4 pits. D, Fracture of the tracheid mostly
at the level of the compound middle lamella. In some areas, the compound middle lamella is etched away, and the bordered pits of the cell
below have secondary wall with fractured pit-canal casts, appearance 3 pits. In some places, it appears the compound middle lamella is thin
enough to transmit the electron beam and visualize the pyrite cast below (arrowhead). E, Fracture at the compound middle lamella shows
appearance 5 pits that consist of the pit membrane. The wall is without a parallel fibrillar nature that is apparent in preserved surfaces of the
secondary wall. F, Pit-chamber casts exposed along the outer pit aperture, showing appearance 6 pits. Scars of the pit canal are evident on most
casts. Tracheid to the left shows appearance 4 pits, which are the pit-chamber casts viewed from the other side, the surface situated next to the
compound middle lamella. G, Tracheids etched with the pit-chamber casts mostly retaining pit-canal casts showing pit appearance 7. The view
is from the lumen of the upper tracheid that is broken away, with the secondary wall and lumen cast removed. The pit canal extends horizontally
across the entire pit-chamber cast. H, Parts of two tracheids etched to show the pit border but from opposite sides. The tracheid to the left has
appearance 8 pits, showing the pit border facing the lumen. The right tracheid shows the pit borders from the side facing the pit chamber, pit
appearance 3. A comparison of 8 and 3 on the left tracheid shows the oppositely inclined, low-angled, crossed pit apertures of contiguous cells.
I, Portion of a tracheid showing the inner surface of the secondary wall with the inner pit apertures, appearance 9 pits. The wall has fibrillar
components in helical arrangement. The lumen cast of the tracheid is visible in the upper unbroken portion of the tracheid.

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436 INTERNATIONAL JOURNAL OF PLANT SCIENCES

Fig. 5 Features of tracheid walls and ray cells. b p pit border, c p pit-canal cast, h p pit-chamber cast, i p inner pit aperture, m p pit
membrane, o p outer pit aperture, r p ray, w p secondary wall. Bar p 10 mm in B, C, E; bar p 1 mm in A, D. A, High magnification of
tracheid walls that shows a pit membrane and the pit borders that face the secondary wall. Pit borders have a circular pattern of fibrillar
components; the pit membrane does not. B, Tracheid etched to reveal pits of cell below, appearance 3, and spiral bands of secondary wall
between the rows of pits (arrows). C, Part of three tracheids broken so they appear sectioned and show the anatomy of bordered pit pairs. The
pits have the pit-canal casts combined with the pit-chamber casts, and the secondary wall is partly etched away. The long pit-canal cast indicates
a thick secondary wall with both inner and outer pit apertures. The secondary wall has an inner portion that is mostly degraded and an outer
portion that is preserved. D, Higher magnification of fig. 5C showing bordered pit pairs where the pit membrane is located between the pit-
canal cast and the combined pit-chamber casts. It thus appears that this bordered pit was aspirated. E, SEM view of ray cells showing procumbent
appearance of some cells and lack of preserved pits on wall faces.

and the inner pit aperture between the pit-canal cast and the
lumen cast. The secondary wall is preserved, and the pits are
slitlike openings, with or without pit-canal casts protruding
through the inner pit aperture, for pit appearance 9 (figs. 3,
4I). When fractured in this way, if some of the tracheid stays
untouched by the acid treatment, the lumen cast can remain
and be seen filling the cell space.
Observations of bordered pit pair structure and tracheid cell
wall. Detailed analysis of tracheids in SEM shows charac-
teristics of wood structure that would have remained unknown
if the pyritized material had only been examined by incident
light. For example, observations of the pyrite casts show that
the pit canal is typically long and the pit chamber is rather
shallow (fig. 2F). The length of the pit canal is a reflection of
the thickness of the secondary wall. The wall between adjacent
tracheids has a mean thickness of 9.5 mm (table 1). However,
the compound middle lamella portion of the wall is quite thin
(figs. 2E, 4D). The wall is thin enough so that the SEM electron

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 DANNENHOFFER & BONAMO—FOSSIL WOOD OF RELLIMIA
beam may penetrate it and reveal the pyrite cast below (fig.
4D).
The fortuitous fracture that occurred through the wall ma-
terial of the secondary wall separating the pit border from the
rest of the wall allowed observation of the structure of the pit
aperture in adjacent cells lying one on top of the other (fig.
4H). These pits, appearance 8, occur on the upper tracheid of
the bordered pit pair and can be compared with appearance
3 pits that are revealed on the adjacent lower tracheid, al-
though they appear to be located on the same tracheid face.
When these pit apertures are compared, the bordered pits have
slitlike apertures that are horizontally oriented and slightly
inclined but in opposite directions (fig. 4H). Thus, the pit ap-
ertures within a bordered pit pair are crossed at a low angle.
The preserved secondary wall, revealed along the surface
that faces the lumen, shows parallel fibrillar components (fig.
4I). The secondary walls of the pit borders show a parallel
arrangement of the fibrillar components in a nearly elliptical
to circular pattern (fig. 5A). Additionally, fractures within the
tracheid wall reveal other features of the secondary wall struc-
ture. Sometimes, these fractures show portions of secondary
wall material that appear like spiral bands of wall between
the rows of pits (fig. 5B).
When the pit membrane is preserved, its fibrillar pattern has
no apparent parallel organization (figs. 4E, 5A). In one sample,
three cracked tracheids were broken through the pyrite casts
and wall material, as if the material had been sectioned (fig.
5C, 5D). The secondary wall is mostly degraded, but the pe-
rimeter portion of the secondary wall remains. The pit cham-
bers have been preserved as a single unit because the pit
membrane is displaced to the outer pit aperture (fig. 5D). This
may indicate that the pit was aspirated when preserved. There
is no apparent differentiation of the pit membrane into torus
and margo commonly found in extant species that have as-
pirated pits.
Ray Cell Structure
Ray cells are square to slightly rectangular in shape when
viewed in radial sections (figs. 1D, 5E). When rectangular, the
long axis is horizontally oriented and cells are procumbent.
The mean length of the ray cells, measured in radial and trans-
verse section, is 87.4 mm (table 1). The length of the ray cell
spans the width of two to three tracheids. The next largest
dimension of the ray cell is its height (measured in both tan-
gential and radial sections), with a mean value of 40.9 mm.
The smallest dimension of the ray cell is its width, with a mean
width of 22.4 mm for all ray cells, 28.8 mm for uniseriate ray
cells, and 15.6 mm for biseriate and multiseriate ray cells. The
size of uniseriate ray cells is almost the same as whole ray
width (table 1). These differences of cell sizes in rays of dif-
ferent seriation correlates with the observation that ray width
is not significantly different with additional seriation (fig. 1A;
table 1).
The type of ray cell present, whether ray parenchyma and/
or ray tracheids, could not be determined by the nature of the
wall pitting. Either the pits present cannot be evaluated as to
whether they are simple or bordered (fig. 1B) or the pits are
not preserved in the ray cell walls (fig. 5E). However, the ray
cell walls are thin, with a mean thickness of 2.2 mm (table 1).
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437
These walls are much thinner than those of the axial tracheids
and may suggest that the rays are made up of ray parenchyma
(fig. 1A). The common lack of preservation of the cell walls
of the rays also suggests ray parenchyma. We believe the rays
are homocellular and consist solely of parenchyma cells.
Discussion
The pyritized material of Rellimia allows a fuller description
of the structure of the wood and secondary xylem tracheids
of this early wood-producing progymnosperm. A previous de-
scription of the anatomy of Rellimia by Kräusel and Weyland
(1938) was accompanied only by a text figure and three poor
illustrations of petrified material (his plate XXIV, figs. 4–6).
Our description of the wood of Rellimia mostly agrees with
the few details provided by Kräusel and Weyland (1938). In
both, the wood surrounds an irregularly three-lobed protostele
with protoxylem areas in the arms. The secondary cells radiate
from the tips of the protostele. In the German material, the
tracheids in transverse section are square to rectangular to
irregular in shape and up to 120 mm in diameter. The tracheids
in the New York material have the same shape and roughly
the same size but may be slightly smaller. The wall sculpture
in the German material is reported as scalariform to reticulate,
with possible bordered pits. The superior preservation of the
New York material shows that all the secondary xylem tra-
cheids have bordered pits and never scalariform wall sculpture.
Ray structure of the German material is virtually unknown
and is described as uniseriate, probably reflecting the narrow-
ness of the ray. The rays in New York Rellimia are also narrow,
but uniseriate to multiseriate rays are present in the wood.
Overall, the anatomical descriptions of the German and New
York material are similar.
Dannenhoffer and Bonamo (1989) correlated the occurrence
and abundance of secondary xylem with the axis size and axis
order. This wood in the largest axis order contains three
growth increments. An area containing radially aligned, thick-
walled cells of the secondary phloem surrounds the secondary
xylem. When preserved, the outer portion of the axis con-
tains thin-walled parenchyma of the cortex. No periderm is
apparent.
The fortuitous preservation of the Rellimia fossils allows an
unprecedented description of the pyritized ancient wood and
tracheid features in an early aneurophyte. This includes study-
ing the material in cross, radial, and tangential sections, much
like studying less recalcitrant fossil material that can be thin-
sectioned or extant woods. In addition, using the SEM enables
a detailed description of the geometry of the bordered pits.
Although this is the first explanation of the SEM views of
bordered pits using a three-dimensional model, other workers
have used electron microscopy to look at and interpret the
bordered pits of fossils (Schmid 1967; Grierson 1976; Leo and
Barghoorn 1976; Beck et al. 1982). Schmid (1967) was the
first to employ electron microscopy to study the secondary
xylem of Devonian fossils. He studied the wood of Callixylon
and Cordaites with the TEM. Using replicas of the wood, he
examined the face views of bordered pits. In some cases, the
pit appeared to show a pit border with a slitlike mineral cast
of the pit chamber. In other instances, the pit border was frac-
tured in a different plane, revealing the much larger pit-
 Table 2
Comparison of Wood Characteristics in the Wood-Producing Aneurophytes and Callixylon
Triloboxylon Aneurophyton Tetraxylopteris Triloboxylon
Rellimia thomsonii arnoldii germanicum schmidtii Proteokalon petryi ashlandicum Callixylon spp.
Age Middle Devonian Middle Devonian Middle–Upper (?Middle) Upper Upper Devonian Upper Devonian Upper Devonian
Devonian Devoniana
Growth layers Yes Probably In some Yes No Yes In most
Tracheids:
Araucarian pits Yes Yes Yes Yes Yes Yes In most
Apertures Inclined, crossed Inclined, crossed Inclined, crossed Inclined Inclined Inclined Inclined, crossed
Pits on all walls Yes Yes Yes Yes Yes Yes No (pits may occur
on tangential wall
in late wood)
Pits reduced on
tangential walls No No No Yes, smaller and Yes Yes, smaller and Yes
fewer fewer
Diameter (mm) R 23–105; T 29–82 30–78 R 23–52; T 21–46 R 26–113; T 33–80 R 50; T 40 … …
Rays:
Width (no. cells) 1 common, also 2 1 common, also 2 1 1 common to 1 1 Mostly 1, or 2,
or 3 multiseriate some multi
Ray tracheids present None seen None seen None seen Yes Yes Yes In most
Sources Dannenhoffer and Stein and Beck 1983 Kräusel and Weyland Beck 1957; Scheck- Scheckler and Banks Scheckler and Banks Lemoigne et al.
Bonamo 1989 1929; Serlin and ler and Banks 1971b 1971a; Scheckler 1983; Chitaley
Banks 1978; Schweit- 1971a 1975 and Cai 2001
zer and Matten 1982
a
Middle Devonian specimens of Tetraxylopteris schmidtii exist if one accepts Sphenoxylon as synonymous and the fragments of Mustafa (1975).
 DANNENHOFFER & BONAMO—FOSSIL WOOD OF RELLIMIA
chamber cast. He showed that appearances of the pit are de-
pendent on the plane of fracture of the material.
Grierson (1976) showed that the pit structure of the primary
tracheids of Leclercqia could be analyzed by SEM interpre-
tation of the entombing pyrite casts. Like Schmid, he inter-
preted the pits based on the fracture plane. In this study of
Rellimia wood, interpretation of the anatomy of the bordered
pit pair incorporates fracture plane and both the carbonaceous
walls that remain and the entombing pyrite casts. Features such
as length and size of the pit canals and pit chambers and the
composition of the pit membrane can be elucidated by un-
derstanding the fracture plane and what remains after etching:
pyrite, wall, or pyrite and wall. For example, it was possible
to determine that the pits of adjacent tracheids have crossed
pit apertures even though the entombing pyrite is opaque. This
understanding allows us to look back at other aneurophytes
and determine that the tracheids also have crossed pit apertures
(Schweitzer and Matten 1982, Aneurophyton pl. 6, figs. 29,
31; Stein and Beck 1983, Triloboxylon arnoldii fig. 12). In
aneurophytes, where the SEM has been used to study pyrite
material, the tracheids have crossed pit apertures. If pyritized
material of the other aneurophytes were studied like this, they
would probably also show this feature, as they are reported
to have inclined pit apertures (table 2).
Our preparation of Rellimia tracheids for the SEM yielded
splitting within the secondary wall that may suggest that the
secondary wall is not homogeneous in structure. At times,
differential preservation occurs such that the outer portion of
the wall is preserved and the inner part is degraded (fig. 5C,
5D). This may indicate that the thin outer portion is degra-
dation resistant, while the inner portion is degradation prone.
In addition, a heterogeneous secondary wall could explain why
we see some of the unusual fracture planes in the tracheids.
For instance, if the wall is fractured at the junction of the
supposed degradation-prone and degradation-resistant layers,
a thin layer of resistant secondary wall remains visible, as in
pit appearance 8.
A heterogeneous secondary wall with degradation-resistant
and degradation-prone portions in Rellimia secondary xylem
tracheids may be comparable to the decay-resistant and decay-
prone secondary wall layers in primary xylem g-type tracheids
of early fossil Lycophytina and p-type tracheids of early fossil
Euphyllophytina (Kenrick and Crane 1997; Friedman and
Cook 2000). Recent work showing enzyme degradation pat-
terns of tracheids of Huperzia and Equisetum indicates that
the common ancestor of the Lycophytina and Euphyllophytina
had secondary wall thickenings with degradation-prone and
degradation-resistant layers (Cook and Friedman 1998; Fried-
man and Cook 2000). Rellimia, within the Euphyllophytina,
indicates that the degradation-prone and degradation-resistant
wall layers may have also been a characteristic of basal lig-
nophytes. Degradation-prone and degradation-resistant layers
are apparently lacking in extant lignophytes (Friedman and
Cook 2000), indicating that this wall characteristic was lost
within the lignophyte clade.
Sometimes, fractures within the secondary wall reveal a ba-
sic spiral pattern that we cannot equate to degradation layer
(fig. 5B). The pattern is similar to the first phase of two-phase
secondary wall deposition described by Bierhorst (1960) and
Bierhorst and Zamora (1965). However, this spiral wall pat-
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439
tern could also be interpreted as reduced scalariform wall
thickenings between which bordered pits have formed, remi-
niscent of the scalariform thickenings of the p-type tracheids
that characterize early fossil euphyllophytes (Kenrick and
Crane 1997; Friedman and Cook 2000). These p-type tra-
cheids have hollow scalariform bars between which are bor-
dered pits, and across the pit aperture is a decay-resistant por-
tion of the secondary wall that has holes or pitlets (Gensel
1979; Hartman and Banks 1980; Kenrick and Crane 1997;
Friedman and Cook 2000). The nature of the tracheid pitting
in Rellimia is quite different from the p-type tracheid. The
bordered pits are circular and no secondary wall is present in
the pit aperture; therefore, there are also no pitlets. Addition-
ally, the pit border is thick, so there are a long pit canal and
two pit apertures instead of a single pit aperture. The sub-
stantial difference in the pitting of Rellimia secondary xylem
tracheids compared to p-type tracheids within primary xylem
of basal euphyllophytes suggests that pitting features may have
been modified rapidly within basal euphyllophytes.
Our addition of details to the description of the wood of
Rellimia furthers knowledge of anatomical features present in
early aneurophytes. It is interesting that the organization of
the first woods is so complex, very much like what we know
in modern plants. One can easily recognize the diagnostic
planes of section: cross, radial, and tangential section. Our
evidence indicates that wood at its inception had the complex,
three-dimensional organization that characterizes later mem-
bers of the lignophyte clade.
Understanding the wood of aneurophytes is interesting in
light of the antiquity and ancestral status of the aneurophytes
in the lignophyte clade. Using information of the known wood
of other members of the Aneurophytales, we characterize the
wood structure of the group (table 2). All have dense pyc-
noxylic wood with square to polygonal tracheids that are sim-
ilar size. Branched tracheids occur in all genera. The tracheids
have elliptical or circular bordered pits arranged alternately in
multiseriate rows on all tracheid walls. The pit apertures are
inclined and slitlike. The rays are typically uniseriate, although
biseriate and multiseriate rays occur in several species. Ray
height is generally tall, with rays at least 20 cells high occurring
in all taxa. Of interest, most are reported to have growth layers.
Aneurophytes have secondary xylem that is remarkably alike
in the details of the structure. The minor differences in the
wood descriptions of the taxa might exist within a single genus
or even a single specimen. The similarity of these wood char-
acters strengthens the unity of the aneurophytes as a group.
However, the slight differences in the wood of the Aneu-
rophytales may indicate early trends in the evolution of wood
structure. In particular, the type and nature of the pitting on
the tracheid walls and the presence of ray tracheids appear to
be different in Middle Devonian versus Upper Devonian taxa.
“Araucarian” pits are alternately arranged, multiseriate, and
hexagonal or angular in shape from crowding (Greguss 1955).
In contrast, coniferous pits are circular in shape, larger in size,
and separate from each other and, when more than uniseriate,
they are normally oppositely arranged. Araucarian pitting is
generally considered the more primitive pit type (Bailey 1933;
Erasmus 1976). Within the progymnosperms, a modification
of araucarian pits by reduction in pit seriation, decrease in the
number of pits, and restriction of pits to the radial face can
440 INTERNATIONAL JOURNAL OF PLANT SCIENCES
be demonstrated (table 2). Rellimia and Triloboxylon arnoldii
have araucarian pits that have never been observed to be other
than multiseriate and densely packed on all walls. However,
in the Upper Devonian taxa, Tetraxylopteris, Proteokalon, and
Triloboxylon ashlandicum, there is evidence for pit modifi-
cation. In all three, although pits occur on all walls, there is
a difference in pitting on the tangential walls. In Tetraxylop-
teris and T. ashlandicum, the pits are reported to be smaller
and less numerous on the tangential walls (Beck 1957; Scheck-
ler and Banks 1971a). In addition, in Tetraxylopteris, the pit-
ting is sometimes alternate but other times opposite. In the
younger genus Callixylon, more specialized pitting is present,
as the pits are grouped on only portions of the wall. In ad-
dition, the pits are generally restricted to the radial walls, ex-
cept in some specimens they may also be present on the tan-
gential wall of late-wood tracheids (Beck 1970). In most
species, there is a reduction in pit seriation, and in Callixylon
arnoldii the pitting is uniseriate. Overall, the genus shows
trends in specialization by reduction both in pit numbers and
in pit seriation and the drifting of pits into pit groups, achieving
a “coniferous”-type pit in some cases.
The wood of Middle Devonian aneurophytes has been re-
ported to have rays with only ray parenchyma cells, whereas
Upper Devonian taxa are reported to have both ray paren-
chyma and ray tracheids (table 2). Aneurophyton, which spans
the Middle to Upper Devonian boundary, has only ray paren-
chyma. Previously, the absence of ray tracheids in older taxa
could be attributed to a difference in how well the wood has
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