Bor.J. Linn. SOC.,68: 303-318. With 4 plates and 2 figures
,June 1 9 7 4
The Givetian flora from Cairo, New York:
Rhacophyton, Triloboxylon and Cladoxylon
LAWRENCE C. MATTEN, F.L.S.
Department of Botany, Southern Illinois University, Carbondale, Illinois 62901,
U.S.A .
Accepted f o r publication September 1973
petrified Rhacophyton, Triloboxylon and Cladoxylon are described from the Givetian of
eastern New York State. In cross section, specimens referable t o Rhacophyton cerafungium
have a mesarch primary xylem strand in the shape of a bar with swollen ends. Pycnoxylic
secondary xylem surrounds the primary xylem. Vascular strands, interpreted as traces, are also
present. Triloboxylon has a three-fluted primary xylem strand surrounded by secondary xylem.
Several mesarch protoxylem areas are present in cross sectional view. The specimens of
Cladoxylon, the first of this genus in the Middle Devonian of North America, show t h e typical
polystelic pattern in cross section. Obvious secondary xylem and peripheral loops are absent.
Lateral appendages were observed o n two of the specimens. A comparison of the Cairo and
Gilboa floras indicates that they represent different ecological niches during the late Middle
Devonian.
CONTENTS
Introduction
. . . . . . . . . . .
Materials and methods
. . . . . . . .
Rhacophyronceratangiurn
. . . '. . . .
Taxonomic considerations
. . . . .
Description of Cairo specimens
. . .
Discussion
. . . . . . . . .
Triloboxylon amoldii
.
. . . . . .
Taxonomic considerations
. . . . .
Nomenclature
. . . . . . . .
Description of Cairo specimen
. . . .
Discussion
. . . . . . . . .
Disposition of Dawson's Dadoxylon hallii
Cladoxylon hue ben
. . . . . . . . .
Taxonomic considerations
. . . . .
Nomenclature
. . . . . . . .
Description of Cairo specimens
. . .
Discussion
. . . . . . . . .
Comparison of t h e Cairo flora with t h e Gilboa flora
. . . . . . . . .
Acknowledgements
References
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INTRODUCTION
This paper is part of a series on a Middle Devonian flora from near Cairo,
New York. To date, the following plants have been described: Actinoxylon
303
304
L. C. MATTEN
banksii Matten (Matten, 1967, 1968b), Reimannia aldenense Arnold, Cairoa
lamanekii Matten and terete axes (Matten, 1973). The current paper describes
three additional members of this flora: Rhacophyton ceratangium Andrews &
Phillips, Triloboxylon arnoldii sp. nov. and Cladoxylon hueberi sp. nov. The
study of the Cairo flora is important because of its size, location and
stratigraphic horizon. Next to the Gilboa flora, this is the largest Givetian flora
in North America and is without doubt the largest petrified Givetian flora. Its
geographic proximity to Gilboa and lower Frasnian floras may stimulate
thought on paleo-ecology and evolutionary trends in the Devonian floras of
New York.
MATERlALS AND METHODS
The specimens studied, iron sulphide petrifications, were collected during
the summers of 1964, 1966 and 1967. The fossils came from a quarry (Fig. 1)
south of New York State Route 145, approximately two miles north-east of
Cairo, New York (Freehold, N.Y., 7% min Sheet 6168 lSE, AMS Series V821,
1945; 905,700 yd E. x 2,221,800 yd N.). The petrifications were collected
from a small, unnamed, local, fine-grained, black, thinly-bedded, shale lens. The
exposure of the lens was no more than 9 m wide and 0.9 m high at its thickest.
The lens is now covered up. The horizon of the fossils is the Kiskatom
Formation (Fisher et al., 1962). Fletcher (1963) placed the Kiskatom, in part,
in the Plattekill Formation. The Kiskatom (Plattekill) is equivalent in age
(Rickard, 1964) to the marine Skaneateles Formation, Hamilton Group,
Gazenovia Stage, Erian Series (= Givetian).
Figure 1. Part of Freehold, New York, 7%min Quadrangle map showing location of Cairo flora
(arrow). The grid represents squares of 1000 yd/side.
THE CAIRO FLORA FROM EASTERN NEW YORK
305
The petrifications were prepared by embedding in palstic, sectioning with a
trim saw and a Gillings-Brownwill thin-sectioning machine, polishing, etching in
nitric acid, and mounting (Heard, 1927; Matten, 1968b). Cellulose acetate peels
were made from some of the specimens, and the peel technique using polyvinyl
chloride (Jennings, 1972) was employed with varying degrees of success.
Photography was accomplished using standard Vickers stereo and high-powered
microscopes, a high intensity light source, and a Nikkormat FTN camera with
microscope adapter.
R HACOPH YTON CERA TANGIUM
Taxonomic considerations
This is a genus based primarily upon information obtained from compression
material. Only R. ceratangium (Andrews & Phillips, 1968; Krausel & Weyland,
1941) and R. zygopteroides Leclercq (Leclercq, 195 1) reveal anatomical data.
The relative importance of anatomy in circumscribing R. ceratungium is
exemplified by the presence of anatomical information in the last two lines of a
22-line diagnosis (Andrews & Phillips, 1968: “All axes observed in T.S. have a
very slender bar of primary wood swollen at each end; this is surrounded by
scalariform tracheids aligned in regular radial rows.”). Should isolated petrified
axes conforming to the description of Andrews and Phillips be placed in the
genus? At present, no other known Devonian plant has anatomy like that of
Rhacophyton, but the occurrence of clepsydropsoid anatomy amongst several
genera in the Carboniferous indicates the possibility of several genera in the
Devonian having Rhacophyton-like anatomy.
Andrews and Phillips also made a nomenclatural change in their paper on
R. cerutangium (Andrews & Phillips, 1968). Dawson (1862) had described
some fragmentary material from the Hamilton Group of New York as
Cyclopteris incerta Dawson. Krausel & Weyland (1941) described Rhacophyton
incertum from West Virginia, using Dawson’s specific epithet. Andrews &
Phillips (1968) collected and described new material of Rhucophyton from
West Virginia that they judged as being conspecific with Krausel & Weyland’s
material, but they excluded Dawson’s material, description and illustrations
(Andrews & Phillips, 1968: “Thus, while the possibility may exist that the
fossils described by Dawson and by White belong to Rhacophyton, we regard it
as highly doubtful and incapable of proof.”). Therefore, since they did not
accept Dawson’s material as being the same as theirs and because they excluded
Dawson’s material from their circumscription of the species, Andrews and
Phillips established the new species Rhacophyton ceratungium.
Description of Cairo specimens
Two specimens (SIPC CQ 60.1, CQ 69) have been determined as conforming
to the anatomy of Rhacophyton cerutangium (Andrews & Phillips, 1968).
Specimen CQ 60.1 is the larger, having dimensions of 3 x 4 mm in cross section
(Plate 1A); the smaller axis (CQ 69) being 1.5 x 2.5 mm in cross section. A
total of 26 transverse and two longitudinal sections was prepared for study.
The primary xylem, in cross section, exhibits a bipolar configuration with a
306
L. C. MATTEN
mesarch protoxylem area at each pole. The primary xylem strand of CQ 60.1 is
3.5 mm long in cross section and the poles appear to be slightly dilated
(Plate 1B). Peripheral loops are absent. The centre of the primary xylem strand
connecting the poles is usually somewhat crushed; however, in section
CQ 60.1-10d a circular protoxylem area occurs midway along the primary
xylem strand. I t is possible that other areas of protoxylem occur along the
primary xylem strand, but definitive proof is lacking due to poor preservation.
Surrounding the primary xylem is a wide band of radially aligned cells,
interpreted as secondary xylem, consisting exclusively of tracheids and rays(?).
The tracheids are rectangular to hexagonal in cross section and range in
diameter from 60-75 pm along the radial plane and from 45-60 pm along the
tangential plane. Pits appear to be distinctly bordered (Plate 1C). In cross
section, radially elongate gaps, about one cell wide, between radial rows of
tracheids are interpreted as being vascular rays. N o evidence of vascular rays
was found in longitudinal section.
In several sections of CQ 60.1 there are two or three terete strands of
primary xylem around the periphery of the secondary xylem (Plates 1D and
2A-C). The origin and fate of these strands can not be determined from the
material available, although they are strongly suggestive of being traces to one
or more lateral appendages.
Discussion
Krausel & Weyland ( 1941) described petrified specimens of Rhacophy ton
from the Upper Devonian of West Virginia. The Cairo specimens conform
remarkably to their description in all critical details. Two of their illustrations
are reproduced here for a direct comparison (Plate2D,E). The ends of the
primary xylem strands are only slightly inflated in both Krausel & Weyland’s
material (Plate 2E) and the Cairo specimens (Plate lA,B). Krausel & Weyland
illustrated a specimen with two terete vascular strands in the cortex (Krausel &
Weyland, 1941: pl. 8, fig. 5 , textabb. 7 ; Plate 2D). The terete strands in the
Cairo specimen (Plates 1D and 2B) compare favourably with those in the West
Virginia specimen (Plate 2D). The vascular strands in the Cairo specimen
(Plate 2A,C) occur within one quadrant of the axis. It is tempting to speculate
on the number of additional traces (if there are any) not preserved, the origin
of these traces, and the organ to which they are connected. A possible
explanation is that there may have been four traces and that the sections of
CQ 60.1 represent a node on a fertile frond, with two of the traces going to a
pair of sterile pinnae and the other two going to the fertile pinnae.
Andrews & Phillips (1968) describe and illustrate ‘peripheral loops’ in their
West Virginian material. The absence of ‘peripheral loops’ in Krausel &
Weyland’s specimens and the Cairo specimens may be explained in several
ways. The ‘peripheral loops’ may appear only at particular stages in trace
formation or in the formation of only certain kinds of traces. Leclercq (1951)
states that the stem stele of Rhacophyton zygopteroides lacks ‘peripheral
loops’. The specimens described by Krausel & Weyland may thus be stems. The
‘peripheral loops’ of Andrews & Phillips may be just an artifact of preservation
due to the breakdown and disintegration of the soft tissues of a trace and the
subsequent separation of the metaxylem from around the area. Andrews &
THE CAIRO FLORA FROM EASTERN NEW YORK
307
Phillips (1968: fig. 42,46) illustrate several specimens supporting this interpretation. In all cases, in Andrews & Phillips’ illustrations, an obvious
‘peripheral loop’ is present at only one end of the bipolar stele. Lastly, the
presence or absence of ‘peripheral loops’ may indicate a speciation problem
that will not be solved until more and better material (compression and
petrification) is found.
The exclusion of Dawson’s material from the Hamilton Group of New York
(Andrews & Phillips, 1968) restricted the genus to the Upper Devonian. The
present report again extends the range of the genus into the Middle Devonian.
It also strengthens the possibility that Dawson’s material may have been
R hacophy ton.
The systematic position of Rhacophyton is still uncertain. Andrews &
Phillips ( 1 968) discuss its affinities with the Aneurophytales, the Coenopteridales and the Trimerophytina and conclude that R hacophyton lies somewhere
between the Trimerophytina and the Aneurophytales in evolutionary
development, but is not a member of either group. Bierhorst (1971) places
Rhacophyton in the Aneurophytales and Andrews (1970), in the Traitk, places
Rhacophyton in the Coenopteridales.
The anatomy of Rhacophyton superficially resembles that of the Aneurophytales, but several differences seem to preclude a close relationship. The
ornamentation of the secondary xylem tracheids is almost exclusively
multiseriate circular bordered pitting on all walls in the Aneurophytales (those
of Rellitnia being the exception). The secondary xylem tracheids are scalariformly bordered in Rhacophyton. Leclercq & Bonamo (1971) suggest that the
secondary xylem of Rhacophyton ceratangium could represent an orientated
metaxylem. They argue that Andrews & Phillips ( 1 968) did not document the
presence of vascular rays in the description, illustrations or diagnosis and that
the occurrence of radially aligned tracheids is not a sufficient enough criterion
to establish the presence of secondary xylem. If Leclercq & Bonamo (1971) are
correct, the lack of true secondary xylem is a major difference between
Rhacophyton and the Aneurophytales. Secondary phloem, periderm, and a
Sparganium-like outer cortex are found in the Aneurophytales but are not
known in Rhacophyton. Branch traces are single and lobed (3-4-lobed) in cross
section in the Aneurophytales and there are several terete traces in the
Rhacophyton material from New York and West Virginia (Krausel & Weyland,
1941) and a single clepsydroid trace in the Belgian material (Leclercq, 1951).
The Aneurophytales lack catadromic fertile pinnae and a pair of sterile axes
(petioles) attached to the same side of the stem above the fertile axes.
Although both the Aneurophytales and Rhacophyton possess unwebbed
dichotomizing ultimate appendages, this feature is found amongst several
diverse Devonian plants such as Archaeopteris, Pseudosporochnus, Hyenia and
Diplo trnema.
In the organization of its fronds, Rhacophyton is at least as specialized as, if
not more so than, the Aneurophytales. Rhacophyton possesses certain
anatomical characters, such as protostely, mesarchy and radially aligned
metaxylem or secondary xylem, that are found amongst diverse, apparently
unrelated plants. Such characters are more indicative of a general morphological level of development in the late Middle and Upper Devonian flora and
should be approached cautiously when trying to determine phylogenetic or
3 08
L. C. MATTEN
systematic relationships. I t is quite likely that Rhacophyton is a Devonian
pteridophyte that parallels the Aneurophytales in the development of mesarch
protosteles, secondary xylem, three-dimensional branching, unwebbed,
dichotomizing ultimate appendages, and the production of large masses of
elongate sporangia on special fertile parts and that it is not closely related to
the Aneurophytales. Leclercq & Bonamo (1971) review the features that the
Coenopteridales and Rhacophyton have in common: radially symmetrical
stellate stem stele, clepsydroid bundles in the fronds, peripheral loops,
repetition of the clepsydroid configuration in the second order of branching of
the fertile frond, etapteroid mode of emission of traces, scalariform tracheids,
organization of fertile parts, and elongate terminal sporangia. On the basis of
the above features, they supported the assignment of Rhacophyton to the
Zygopteridaceae of the Coenopteridales.
Finally the similarity of certain features of Rhacophyton and Stenokoleos
should be pointed out. Stenokoleos bifidus Matten & Banks (Matten & Banks,
1969) from the Frasnian of New York had an axis that bore pairs of
appendages on one side of the axis at each node in a distichous fashion, the
main axis stele was bilateral, mesarch and had peripheral loops, and it had
clepsydroid traces and mostly scalariformly bordered pits. Matten & Banks
(1969) suggested that Stenokoleos was an early representative of the Lower
Carboniferous protostelic pteridosperms exemplified by Tristichia and
Tetrastichia.
TRILOBOXYLON ARNOLDII
Taxonomic considerations
A nomenclatural situation analogous to that of Rhacophyton ceratangium is
found in Triloboxylon. Dawson (1862) described and illustrated several wood
fragments from the “Tully pyrite”, Hemlock Creek, Ontario County, New
York as Dadoxylon halli. Arnold (1940) described and illustrated several new
specimens (including the first to show primary tissues) from western New York
and included them in Dawson’s species, transferring it to the genus Aneurophyton as A. hallii (Dawson) Arnold. Scheckler & Banks (1971a) described in
detail two specimens lent to them by Dr Arnold, one of which was described in
Arnold’s 1940 paper. These specimens were collected from Spring Creek,
.4lden, Erie County, New York. Scheckler & Banks (1971a) then made the
incorrect combination Triloboxylon hallii (Arnold) Scheckler & Banks, instead
of Triloboxylon hallii (Dawson) Scheckler & Banks, since the original species
was established by Dawson, not Arnold.
However, it is believed that the intent of Scheckler & Banks was to exclude
from the new combination and circumscription of the species the material
described by Dawson (i.e. analogous to the Rhacophyton ceratangium
situation). The reasons for this belief are: 1. they list the type locality of
Triloboxylon hallii as Spring Creek, Alden, Erie County, New York (Arnold’s
locality not Dawson’s); 2. they refer to the holotype as being in the University
of Michigan Museum of Paleontology (no. 23848) and they illustrate the
specimen (Scheckler & Banks, 1971a: fig. 23, 25-28) showing features of the
phloem, cortex, periderm, traces and primary xylem (none of which otcur in
THE CAIRO FLORA FROM EASTERN NEW YORK
3 09
Dawson’s original specimens); 3. they omit any mention of Dawson’s material
or his publication. The writer believes, as apparently do Scheckler & Banks,
that the material described by Dawson (1862) as Dadoxylon halli may possibly
belong to Triloboxylon, but that such an assignment is incapable of proof. The
exclusion of Dawson’s material from the circumscription of a species based
upon Arnold’s specimens requires that a new species name must be proposed
for Arnold’s and Scheckler & Banks’ material, and for the new Cairo material.
Triloboxylon arnoldii is proposed.
Nomenclature
Triloboxylon arnoldii sp. nov.
Triloboxylon hallii (Arnold, 1940) Scheckler & Banks, 1971 (non Dawson,
1862). Scheckler, S. E. & Banks, H. P., 1971. Anatomy and relationships of
some Devonian progymnosperms from New York. Am. J. Bot., 58: 737-5 1,
fig. 22-28.
Aneuvophyton ha& (Dawson) Arnold (in part). Arnold, C. A., 1940. Structure
and relationships of some Middle Devonian plants from western New York.
Am. J. Bot., 27: 57-63, fig. 2,3.
Diagnosis-As in Scheckler & Banks (1971a: 747-8).
Illustrations of species-Arnold (1940: figs 2,3); Scheckler & Banks (1971a:
figs 22-28).
Holotype-Museum of Paleontology, University of Michigan no. 2 3 848.
Type locality-Bed of Spring Creek where it crosses State Route 20 near Alden,
Erie County, New York (Corfu 7% min. Quadrangle, 1950, N 4252.5 x
W 7822.5).
Horizon-According to Arnold (1940), this is the Ledyard Member of the
Ludlowville Formation, which is in the Hamilton Group of the Tioughnioga
Stage, Erian Series, Givetian (Rickard, 1964).
Description of Cairo specimen
A single specimen, SIPC CQ 20, from the Cairo flora is referable t o
Triloboxylon arnoldii. The specimen was approximately 15 mm long and 3 mm
wide and was cut into eight transverse and five longitudinal sections. Only
primary and secondary xylem was preserved in the specimen.
In cross section, the primary xylem, consisting of hexagonal tracheary
elements, has a three-armed configuration (Plate 3A). Protoxylem areas are
mesarch, terete and discrete from one another. At least seven distinct
protoxylem areas can be seen in some sections. The protoxylem elements’ are
distinctly smaller than the metaxylem elements. The former range in diameter
from 14 to 25 pm, the latter gradually increase in size from 25 pm next to the
protoxylem to 75 pm farthest from the protoxylem. In longitudinal section the
protoxylem elements have helical and scalariform-reticulate thickenings
(Plate 3D). The metaxylem elements have multiseriate circular bordered pits.
Secondary xylem composed of tracheids and vascular rays surrounds the
primary xylem. The tracheids are rectangular to hexagonal in cross section
(Plate 3B). The cells are slightly larger in the radial dimension, averaging 35 pm
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L. C. MATTEN
radially and 3 1 pm tangentially. The tracheids have 2- to 6-seriate circular
bordered pits on both the tangential and radial walls (Plate 3E). The pits are
9-12 pm wide and their apertures are orientated horizontally to slightly
obliquely. In tangential sections, several tracheids appear to have branched ends
(Plate 3F). Only uniseriate rays were observed, attaining a height of over 15
cells (Plate 3C).
Discussion
Scheckler & Banks (1971a) distinguish Triloboxylon arnoldii (formerly
T. hallii) from Triloboxylon ashlandicum Matten & Banks (Matten & Banks,
1966) by the presence of sclerotic bundles in the inner cortex and the absence
of parenchyma in the primary xylem of Triloboxylon arnoldii and the presence
of a parenchymatous inner cortex and strands of xylem parenchyma near the
tips of the arms of primary xylem in Triloboxylon ashlandicum. I t is
questionable whether these differences warrant separation at the species level.
Beck (1970) discussed the separation of genera within the Archaeopteridales
based upon differences that may not be good generic characters. He suggested
that Svalbardia, Actinoxylon, Archaeopteris and Eddya may well be part of the
same genus and perhaps represent variations that may occur within one
individual plant. The same type of argument may be used in respect of the two
species of Triloboxylon. The distribution and occurrence of sclerenchyma in
the cortex and the amount of parenchyma in the primary xylem may well vary
within the individual plant. On the other hand, several species within a genus or
genera within a family may have so similar an anatomy that, if they were
fossilized and some of the detail lost, they would be difficult if not impossible
to distinguish. Thus, until organic connection can be proved, species such as
these, based on slight differences in anatomical organization will have to be
maintained.
Disposition ofDawson 's Dadoxylon hallii
The original description of Dadoxylon hallii (Dawson, 1862) was based on
isolated wood fragments found in the Devonian of New York State. For almost
a century, any wood fragment found in the Middle and lower Upper Devonian
of New York was usually placed in the single species of Dawson if it had
pycnoxylic secondary xylem with multiseriate round bordered pits on the
tracheids. However, six species, each having secondary xylem and belonging to
the Aneurophytales, have now been described from New York; Sphenoxylon
eupunctatum (Thomas) Read (Thomas, 1935; Matten & Banks, 1967),
Tetraxylopteris schmidtii Beck (Beck, 1957), Triloboxylon ashlandicum
(Matten & Banks, 1966; Scheckler & Banks, 1971a), Triloboxylon arnoldii
(Arnold, 1940; Scheckler & Banks, 1971a), Proteokalon petryi Scheckler &
Banks (Scheckler & Banks, 1971b) and Cairoa lamanekii (Matten, 1973).
Isolated wood fragments of any of these species would show pycnoxylic
secondary xylem with crowded, multiseriate bordered pitting on radial and
tangential walls, and the presence of narrow (normally uniseriate) tall rays. It
would be wise to place such fragments of secondary wood in a form taxon such
as Dadoxylon hallii Dawson while retaining the other genera and species for
only those specimens exhibiting critical features.
THE CAIRO FLORA FROM EASTERN NEW YORK
311
CLADOXYLON HUEBERI
Taxonomic considerations
Cladoxylon is currently considered to be a form genus for petrified axes with
a particular type of polystelic anatomy. The occurrence of Cladoxylon-type
anatomy in Pseudosporochnus and Calainophyton only strengthens the case for
using Cludoxylon as a form genus. Species of Cladoxylon about which
extensive morphological information is known should be removed from the
genus and a new genus should be established for them. Such an example is the
Middle Devonian Cladoxylon scoparium Krausel & Weyland (Krausel &
Weyland, 1926).
Specimens of Cladoxylon have been found in the Upper Devonian beds of
New York (Read, 1935; Banks, Hueber & Wilcox, 1959; Hueber, 1960).
Hueber (1960) described the most complete specimen of Cladoxylon found, to
date, in the Devonian of New York. Leclercq (1970) also described Hueber’s
material as “Cladoxylon sp. Hueber” and created of Hueber’s specific
e p i t h e t 4 dichotomum, a nomen nudum. Such a well-defined species deserves
a proper specific epithet and one is given that honours the man who first
described it.
Nomenclature
Cladoxylon hueberi sp. nov.
Cladoxylon sp. sensu Hueber (1960); Leclercq (1970).Leclercq, S., 1970. Classe
des Cladoxylopsida Pichi-Sermolli, 1959. In: Andrews, H. N., Arnold, C. A.,
Boureau, Ed., Doubinger, J. and Leclercq, S., Trait6 de Palkobotanique, IV
(I), Filicophyta. Paris: Masson, 138-9, figs 118, 119.
C. dichotomum Hueber ex Leclercq (1970), nom. nud. in adnot.
Diagnosis-As in Leclercq (1970: 138-9).
Illustrations of species-As in Leclercq (1970, figs 118, 119).
Holotype-Specimen no. 1634-1 in the Paleobotanical Collections, Cornell
University, Ithaca, New York.
Type locality-Quarry on the west slope of Cave Mountain on the road to
Jewett going south from State Route 23 in East Ashland, Greene County,
New York (Ashland, N.Y. 7% min. Quadrangle, Sheet 6168 IV SE, AMS Series
V821, 1946).
Horizon-The quarry is in the Oneonta Formation (Fisher et al., 1962), which
is equivalent in age to the marine Ithaca Formation, Genesee Group, Finger
Lakes Stage, Senecan Series, Lower Frasnian (Rickard, 1964).
Description of Cairo specimens
Six specimens have been determined as belonging to the form genus
CZadoxylon and in particular to Cludoxylon hueberi (SIPC CQ 11, CQ 51,
CQ 78, CQ 89, CQ 90, CQ 110). A total of 38 transverse and 9 longitudinal
sections was examined. The largest axis (CQ 89) was 22 mm wide and 5 5 mm
long, with other specimens ranging in diameter from 5 to 6 mm. Each specimen
contains numerous strands of primary xylem but due to compressions of the
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L. C. MATTEN
axes it is sometimes difficult to discern individual strands (Plate 4A). A similar
state of preservation was described for petrified axes of Pseudosporochnus
nodosus Leclercq & Banks (Leclercq & Banks, 1962) and for the original
material of Cludoxylon hueberi (Hueber, 1960). Such distortion due to
compression creates problems with reconstruction of the original form of the
stele. However, in specimen CQ 110 there appear to be about 34 xylem strands
in cross section (Fig. 2B) and in CQ 51 about 15 strands (Fig. 2A). Some of the
xylem strands are peripheral and others are central in position. They vary in
shape from round to oval or elongate strands that are straight or in U, Y or W
configurations. Although it is believed that the elongate strands were normally
elongated in the radial dimension in the living state, many of the strands appear
t o be elongated in a tangential orientation in the compressed fossilized state
(Fig. 2).
Figure 2. Camera lucida drawings of steles of two specimens of Cladoxylon hueberi from the
Cairo flora: A, SIPC C Q 51-2d,x15; B, SIPC CQ llO-ld,~ 1 0 .
The protoxylem is mesarch (Plate 4B) and no peripheral loops are visible
(the term peripheral loop, as used here, refers to the presence of either areas of
parenchyma in the position of the protoxylem or protoxylem lacunae).
However, some of the protoxylem areas appear disrupted as if either a
protoxylem lacuna was being formed or thin-walled parenchyma was present
and had been crushed during preservation. Protoxylem is found not only near
the ends of the elongate strands but also as a central band along their length
(Plate4B). Those xylem strands which are oval to round in cross section
normally have the protoxylem in a centrarch position. The xylem is all primary
and seems to be composed exclusively of tracheids. In transverse section
(Plate 4B), the tracheids range in diameter from 8 to 45 pm, the smallest
elements being protoxylem. The lateral walls of the tracheids are poorly
preserved (attempts to photograph details failed), but one can occasionally see
sectional views of bordered pits (tangential section) and rarely an indication of
THE CAIRO FLORA FROM EASTERN NEW YORK
313
2- t o 4-seriate bordered pitting in face view (radial section). In addition, some
tracheids appear to have scalariform or reticulate lateral wall thickenings.
In general, the ground tissue is not preserved. In the few sections that show
cells, they appear to be parenchymatous. No packets or bundles of sclerenchymatous cells could be determined, but their absence may well be due to
poor preservation.
Evidence of traces and lateral appendages is found in several specimens. In
C Q 7 8 the xylem strands seem to elongate radially in clusters of three. In
specimen CQ 5 1 part of a lateral appendage can be seen. I t seems t o contain a
single elongate vascular strand. The most interesting specimen is CQ 90. At one
level a primary lateral with three vascular strands is visible (Plate 4C). At a
higher level, a secondary lateral (probably borne on the primary lateral) with a
single vascular strand is visible in a position that is adaxial to the primary lateral
(Plate 4D). Above this level the primary lateral has two vascular strands. The
secondary lateral is not preserved beyond this level, so that it could not be
determined whether the division of the vascular strand was an indication of a
subsequent division. The primary lateral has two vascular strands at and above
the level of appearance of the secondary lateral (Plate 4D,E).
There are two tissue zones surrounding the vascular tissue of the primary and
secondary laterals. There is an inner zone of loose, dark-filled cells and an outer
zone of more compact parenchymatous ceIls (Plate 4C,D,E). N o sclerenchyma
was observed.
Discussion
The Cairo Cladoxylon is similar to the type of Cladoxylon hueberi in that
the formation of traces seems t o be by contiguous steles (usually three), by the
possession of terete, oval and elongate steles, by the lack of secondary xylem,
and by the lack of a well-defined peripheral loop. The one difference seems to
be the absence of sclerenchymatous bundles in the cortex of the Cairo
Cladoxylon and its presence in Cladoxylon hueberi (Hueber, 1960). Leclercq
(1970) remarked that the Lower Carboniferous species of Cladoxylon show
only minor differences between them and that some of them appear to
represent the structure of the same axis sectioned at different levels. This is also
probably true of the Devonian specimens from Cairo and Cave Mountain. The
lack of sclerenchyma in the Cairo specimens is most likely due to preservation
or morphological variation within a single plant or species. Thus the Cairo
Cladoxylon is considered to be part of the same plant as Cladoxylon hueberi.
The Cairo Cladoxylon hueberi is the first species of this genus to be reported
from the Middle Devonian of North America. Calarnophyton (Bonamo &
Banks, 1966) has been described from compression material collected from the
Ashoken Formation (= Givetian) in eastern New York. Other North American
species of Cladoxylon are Cladoxylon dawsonii Read (Read, 1935) and
Cladoxylon sp. (Read, 1936). The former has been described from the Upper
Devonian of New York and the latter from the Upper Devonian New Albany
Shale near New Albany, Indiana.
Cladoxylon hueberi differs from Cladoxylon dawsonii in the possession of
terete steles and the absence of secondary xylem, a well-defined peripheral loop
and radial bands of thick-walled cells in the cortex. This plant also differs from
314
L. C. MATTEN
Cladoxylon sp. (Read, 1936) in that it lacks a well-developed central zone of
parenchyma with perhaps included sclerenchymatous strands, it is much
smaller, and it lacks the distinct separation of circular steles in the centre of the
axis and radiating elongate steles on the periphery. Leclercq (1970) indicates
that, on the basis of the brief description and the single line-drawing of Read,
Cladoxylon sp, from the New Albany Shale shows certain affinities with
Pietzschia schulleri Gothan.
Of the European species of Cladoxylon, C. taeniatum (Unger) SolmsLaubach (including C. dubium Unger (Solms-Laubach, 1896)), C. solmsii
P. Bertrand (Bertrand, 1914), C. kidstonii Solms-Laubach (Solms-Laubach,
1910) and C. mirabile Unger (Unger, 1856) differ from C. hueberi in their
possession of secondary xylem. Cladoxylon radiatum (Unger) P. Bertrand
(Bertrand, 1935) possesses thick-walled cells in the ground parenchyma (absent
in C. hueberi) and the production of traces involves four to six steles (1-3 in
C. hueberi) and results in the formation of about ten traces (1-3 in C. hueberi).
Cladoxylon waltonii (Long, 1967) differs from C. hueberi in that it possesses
well-defined peripheral loops and clepsydroid traces that are formed from a
single xylem arm, and the stele in some axes is a U-shaped ‘actinostele’.
Cladoxylon scoparium (Krausel & Weyland, 1926) from the Middle
Devonian of Germany is the only species within the genus to show extensive
external morphology. Krausel & Weyland’s reconstruction of the plant showed
a digitately branching main axis bearing non-webbed, dichotomizing units
(sterile pinnules) and fan-shaped structures bearing sporangia. The morphology
of the fertile units (as they are presently understood) clearly distinguishes this
plant from the other genera in the family for which external morphology is
known (i.e. Pseudosporochnus and Calamophyton). Anatomically, the presence
of small amounts of secondary xylem and possibly the presence of small groups
of sclerenchyma in the ground tissue differentiate C. scoparium from
C. hueberi. As was stated earlier, Cladoxylon scoparium should be removed
from the form genus Cladoxylon and placed in a newly created genus within
the Cladoxylaceae.
Pseudosporochnus nodosus (Leclercq & Banks, 1962) differs from the genus
Cladoxylon in its external morphology, but its internal anatomy is so similar to
Cladoxylon that isolated petrified fragments of Pseudosporochnus would
undoubtedly be classified as a species of Cladoxylon. The same statement could
be made for Calamophyton (see Leclercq & Schweitzer, 1965). The occurrence
of Pseudosporochnus and Calamophyton in the Devonian of New York
indicates the possibility that Cladoxylon hueberi may be the structurally
preserved condition of one of these two genera.
COMPARISON OF THE CAIRO FLORA WITH THE GILBOA FLORA
To date, seven plants have been described and two additional ones have been
recorded from the Cairo locality. Those that have been recorded but not
described belong to the genera Stenokoleos (Matten, 1968a) and Eospermatopteris (Banks, 1966). The number of different plants in the Cairo flora is as
large as that of any single Middle Devonian locality in North America. The
Gilboa flora comes closest in numbers of different plants (Table 1). I t is very
close geographically (about 17 miles (27 km) west of the Cairo locality) and
THE CAIRO FLORA FROM EASTERN NEW YORK
315
Table 1. Comparison of Gilboa and Cairo floras. Lists compiled from Banks
(1966), Skog & Banks (1973), and Matten (1968a, 1968b, 1973 and this paper)
Gilboa Flora
-
Cairo Flora
~~
Eospeimatopteris erianus (Dawson) Goldring
Aneurophyton erianum (Dawson) Krausel & Weyland
Caulopten's lockwoodii Dawson
Amphidoxodendron dichotomum Grierson & Banks
Archaeosigillaria vanuxemi (Goppert) Kidston
Protolepidodendron scharianum Krejri
P. gilboense Grierson & Banks
Sigillaria? gilboense Goldring
Prosseria grandis Read
Ibyka amphikoma Skog & Banks
- stems and branches
Triloboxylon arnoldii sp. nov.
Cairoa lamanekii Matten
Reimannia aldenense Arnold
Actinoxylon banksii Matten
Rhacophyton ceratangium Andrews & Phillips
Eospermatopteris
Stenokoleos
Cladoxylon hueberi sp. nov.
Terete axes
stratigraphically (the Gilboa flora being of Moscow age and the Cairo flora of
Skaneateles-Ludlowville age and all units belonging to the Hamilton Group).
In fact, the two floras are not similar! The only common plant, which is
classified as Eosperrnatopteris-Aneurophyton, occurs in great numbers at
Gilboa as casts and impressions. The Gilboa flora, dominated by lycopods and
sphenopsids, immediately brings to mind the later Carboniferous swamp
communities. The swollen bases of the Eosperrnatopteris stumps are indicative
of a swampy environment, particularly since the stumps were found as if they
were fossilized in situ (Goldring, 1924; Banks, 1965). The Cairo flora is
dominated by ferns and progymnosperms. The plants are found associated with
scattered fish scales and large numbers of brachiopods. I t is likely that the
plant-bearing lens represents a localized stream or pond deposition on the
low-lying lands bordering the epicontinental Devonian sea of eastern North
America (Banks, 1965). The fragmentary nature of the remains indicates that
the plants were probably transported t o the depositionary environment.
Due to the fluctuating shoreline during Middle and Upper Devonian time it is
difficult to determine whether the Cairo flora was more of an upland flora than
the Gilboa flora; however, one can use the differences in the compositions of
the two floras to support the idea that they inhabited different ecological
niches. Even if each flora consists of allochthonous accumulations of plants,
the areas supplying them apparently were different. Populations of lycopods
etc. either grew at the Gilboa site or were washed into it, and populations of
ferns and progymnosperms similarly supplied the Cairo site. Such a difference
in composition indicates that the Devonian landscape was heterogenous, with
populations of different plants growing in different places forming different
communities. Because of this, reconstructions of the Devonian landscape
should show particular associations, separating the various ecological niches.
Comparisons between floras may have to be made on an ecological level rather
than on an evolutionary one.
It may be visualized that, as the shoreline fluctuated, the lowland flora of
the Devonian was forced to migrate eastward or westward. Plants inhabiting
certain niches persisted either as long as similar niches were available or until
they were replaced by better-adapted species (Banks, 1965). I t seems most
logical to assume that the ancestors of the coal swamp plants were Devonian
22
316
L. C. MATTEN
swamp plants and, from this viewpoint, the more inland plants of the Devonian
were then probably ancestral to the more inland plants of the Carboniferous.
Much more information will be needed before the westward migration and
evolution of floras in the Paleozoic of eastern North America can be fully
documented.
ACKNOWLEDGEMENTS
This work was supported, in part, by the Office of Research and Projects,
Southern Illinois University (2-2-22, 2-2-3 5 ) . The manuscript was completed
while the writer was on sabbatical leave at the School of Plant Biology,
University College of North Wales, Bangor, and he wishes to acknowledge with
thanks the facilities that it provided. In particular, the writer wishes to
acknowledge the hours of encouragement and constructive criticism given by
Dr William S. Lacey and Dr Jeffrey Duckett.
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EXPLANATION O F PLATES
PLATE 1
Sections of Rhacophyton ceratangium from th e Cairo flora.
A. Cross section showing radially aligned secondary xylem tracheids surrounding a barshaped
primary xylem, SIPC (Southern Illinois Paleobotanical Collection) CQ 60.1-10 ( ~ 2 8 ) .
B. Cross section showing bipolar primary xylem, SIPC CQ 60.1-7d ( ~ 3 5 ) .
C. Tangential section of secondary xylem showing bordered pits on t h e radial walls of the
tracheids, SIPC CQ 60.1-1.3 ( ~ 1 5 7 ) .
D. Cross section showing terete trace and periphery of secondary xylem, SIPC CQ60.1-20d
(x110).
318
L. C. MATTEN
PLATE 2
Cross sections of Rhacophyron cerarungiium from the Cairo flora (A-C) and West Virginia (D,E)
showing the similarity between the Middle Devonian and Upper Devonian material.
A. Section showing three traces, SlPC CQ 60.1-22u ( ~ 5 0 ) .
B. Section of mesarch trace in C, SIPC CQ 60.1-20d ( ~ 1 1 0 ) .
C. Section showing three traces at a higher level than A, indicating little lateral movement of
the traces, SIPC CQ 60.1-20d 1x43).
D. Section of Upper Devonian specimen from West Virginia with two terete mesarch traces
(from Krausel & Weyland, 1941: Taf. IX, Fig. l ) , Senck. Mus. B 3081 (x30).
E. Section of Upper Devonian specimen from West Virginia with a bipolar primary xylem
surrounded by secondary xylem. There is n o obvious peripheral loop in either this specimen or
the one figured in D (from Krausel & Weyland, 1941: Textabb. 7 ) . Senck. Mus. B 3082/2
(~14).
PLATE 3
Transverse (A,B) and longitudinal (C-F) sections of Triloboxylon arnoldii from the Cairo flora.
A. Section showing three-lobed primary xylem with several (six) mesarch protoxylems and
secondary xylem, SIPC CQ 20-1 (~50).
B. Secondary xylem showing angular tracheids and narrow rays, SlPC CQ 20-1 (~100).
C. Tangential section showing uniseriate rays (arrows), SIPC CQ 20-2-2 ( ~ 1 2 0 ) .
D. Section through t h e protoxylem showing helical and reticulate-scalariform elements, SlPC
CQ 20-2.1 ( ~ 1 2 0 ) .
E. Radial section through secondary xylem showing multiseriate, circular-bordered pitting,
SIPC CQ 20-4.2 ( ~ 1 2 0 ) .
F. Section showing tracheid with forked tip, SIPC CQ 20-2.1 ( ~ 1 8 0 ) .
PLATE 4
Transverse sections of Cladoxylon hueberi from the Cairo flora.
A. Section showing degree of compression of the axis forcing the xylem strands against one
another and obliterating the ground tissue, SlPC CQ 51-2u ( ~ 3 3 ) .
B. View of compressed mesarch xylem strands, SIPC CQ 51-211 (x100)
C. Primary lateral appendage with three xylem strands (arrows) below level of appearance of
secondary lateral appendage, SIPC CQ 90-3d ( ~ 5 3 ) .
D. Section at level of first appearance of the secondary lateral appendage. The secondary lateral
appendage, on the left, has a single xylem strand (arrow) and the primary lateral appendage, on
the right, has two xylem strands (arrows), SlPC CQ 90-3u (~53).
E. Section at higher level (than in D). There are t w o xylem strands each (arrows) in the primary
and secondary lateral appendages, SIPC CQ 90-2d (x53).
Bat. J. Linn. Sac., 68 (1974)
L. C. MATTEN
Plate 1
(Facing p . 318)
Bat. -7. Linn. SOC.,
68 (1974)
L. C. MATTEN
Plate 2
B0t.Y. Linn. SOC.,68 (1974)
L. C. MATTEN
Plate 3
Bat. 3.Linn. Sac., 68 (1974)
L. C. MATTEN
Plate 4