IAWA Journal, Vol. 26 (4), 2005: 489–505
Transcript of IAWA Journal, Vol. 26 (4), 2005: 489–505
IAWA Journal, Vol. 26 (4), 2005: 489–505
CONIFER WOODS OF THE MIDDLE JURASSIC HOJEDK FORMATION (KERMAN BASIN) CENTRAL IRAN
Imogen Poole1* & Majid Mirzaie Ataabadi2, 3
SUMMARY
This paper documents the first record of permineralised wood from Mid-dle Jurassic coal bearing deposits to the north of Kerman, Iran, deposited c. 170 million years before present. The coniferous woods have character combinations resembling, and thus have been assigned to, the genera Xen-oxylon Gothan and Agathoxylon Hartig. Since Xenoxylon is essentially Laurasian and Agathoxylon has been recorded from the Northern Hemi-sphere during the Mesozoic, these woods help confirm conclusions from recent geological studies that place the Kerman Basin of Iran in southern Laurasia during the Jurassic.
Key words: Iran, Jurassic, conifer, Gondwana, Laurasia, fossil wood.
INTRODUCTION
Late Triassic–Middle Jurassic sediments extending from Central Iran northwards and east into Afghanistan have yielded excellently preserved floras. These Mesozoic plant-bearing deposits are important because they form a continuous, uninterrupted series from the Norian through to the Middle Jurassic (Schweitzer et al. 1997). The fossils orig-inate from the mining areas in Northern Iran (Alborz), Central Iran (Kerman Basin) and Northeast Afghanistan (Hindukusch) (Schweitzer et al. 2000). In western and northern Alborz the deposits are completely terrestrial whereas in central and eastern Alborz and to the south in the Kerman Basin they have occasional marine intercalations (Schweitzer et al. 1997). During the last few decades plant microfossils (e.g. Kimiyai 1968; Arjang 1975; Ashraf 1977; Achilles et al. 1984) and macrofossils from this area have been com-prehensively studied. From the leaf floras alone c. 60 genera of plant macrofos-sils have been described, ranging from bryophytes, lycophytes, equisetales and ferns (Schweitzer 1978; Schweitzer et al. 1997) to caytoniales, cycadophytes, bennettites, ginkgophytes, czekanowskiales and coniferales (Poliansky & Safronov 1973; Barnard & Miller 1976 and references therein; Sadovnikov 1976; Vassiliev 1985; Schweitzer & Kirchner 1995, 1996, 1998, 2003; Schweitzer et al. 2000; Mirzaie Ataabadi 2002). Based upon these publications the Rhaeto-Jurassic sediments of the Alborz region
1) Wood Anatomy Section, National Herbarium of the Netherlands, University of Utrecht branch, P.O. Box 80102, 3585 CS Utrecht, The Netherlands & Palaeontological Museum, Univer-sity of Oslo, P.O. Box 1172 Blindern, N-0318 Oslo, Norway [E-mail: [email protected]]. *Author of correspondence.
2) University of Esfahan, P.O. Box 81745/188, Esfahan, Iran.3) Present address: Division of Geology and Palaeontology, Department of Geology, University
of Helsinki, P.O. Box 64, FIN-00014, Helsinki, Finland [E-mail: [email protected]].
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IAWA Journal, Vol. 26 (4), 2005490 491Poole & Mirzaie Ataabadi — Jurassic conifer woods
appear to be the most fossiliferous locality in Iran to date (cf. Schweitzer & Kirchner 1995, 2003) but more recent surveys by Mirzaie Ataabadi (2002) and Vaez Javadi and Mirzaie Ataabadi (2006) indicate that the Kerman area is also rich in fossils. Fossil wood from this geographical region is rare when compared with leaf floras. Seward (1912) reports the presence of Cupressinoxylon in the Saighan Series of North Afghanistan, and Sitholey (1940) and Jacob and Shukla (1955) document Mesembri-oxylon sp. from the same locality. Schweitzer and Kirchner (1996), however, synony-mise the latter with Xenoxylon barberi (Seward) Kräusel which they describe from the same series in Northeast Afghanistan. From Middle Jurassic deposits of the Ferizi area, northeastern Iran, Fakhr and Marguerier (1977) in Fakhr (1977) assigned woods to Prototaxoxylon. More recently Nadjafi (1982) described Jurassic woods of Alborz Mountains (Fig. 1) and assigned them to five genera, Xenoxylon, Prototaxoxylon, Meta-taxodioxylonʼ, ʻProtosciadopitys and ʻProtopinoxylonʼ. The last three genera were newly erected in Nadjafiʼs (1982) unpublished thesis and are thus invalid according to the rules laid down by the I.C.B.N. (Greuter et al. 2000). The fossil woods, which form the subject of this study, originate further to the south in Middle Jurassic deposits in the Kerman Basin of Central Iran (Fig. 1; Lapparent & Davoudzadeh 1972; Schweitzer & Kirchner 1995) known for the coal bearing deposits that cover an area of 700 to 1050 km2. Even though palaeobotanical investigations in this Basin have been undertaken, workers have concentrated on the leaf and fructifica-tion flora (see references listed above) and to our knowledge no wood specimens have been recorded to date.
GEOLOGICAL SETTING
The area around northern Kerman (Fig. 1) had, until the Triassic, formed part of the Ola-cogen Basin with a connection to Palaeotethys. Significant palaeogeographical changes subsequently occurred as a result of the Early Cimmerian tectonic movements of the Late Middle–Late Triassic (Carnian–Norian) and thus played a major role in the geo-logical history of Jurassic deposits in this area. After these orogenic activities, faulting to the north and south of this area created a new basin in between the faults. The high amount of subsidence associated with these events caused the deposition of a thick sequence of terrigenous sediments that lasted until the Middle Cimmerian (Bajo-cian–Bathonian) (Berberian & King 1981). The detrital sediments in this basin vary in thickness from a few metres up to more than 3000 m. These sediments are known as the Shemshak group (formerly the Shemshak Formation) which includes the Nayband, Abhaji, Badamu and Hojedk Formations (Fig. 2) of Central Iran (Aghanabati 1977; Stöcklin & Setudenia 1991). The Hojedk Formation (Stöcklin & Setudenia 1991) comprises more than 1000 m succession of shale, sandstone, conglomerate and coal horizons (Fig. 2). The coal ho-rizons are fossiliferous yielding beautifully preserved leaf floras. The woody material studied herein was found in the “coal bearing series, coal horizon D” in the Kerman Basin equivalent to the “Dansirit Series” of the Alborz region (Table 1 of Schweitzer et al. 2000). The ammonite fauna from the underlying marine limestones of the Badamu
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IAWA Journal, Vol. 26 (4), 2005490 491Poole & Mirzaie Ataabadi — Jurassic conifer woods
Fig. 1. – a: Map showing the position of the study area in relation to Kerman, in south central Iran, with the area north of Kerman enlarged indicating the fossil locality in relation to other major towns in the area. – b: Geological map of the immediate area showing the Jurassic coal-bearing deposits in which the fossil flora was found. The fossil locality is indicated.
56° 24'
31° 31'
56° 35'
31° 31'
31° 15'
56° 35'56° 24'
31° 15'
a
b
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IAWA Journal, Vol. 26 (4), 2005492 493Poole & Mirzaie Ataabadi — Jurassic conifer woods
Thick to medium bedded limestone
Red conglomerate and sandstone
Alternation of greenish grey shale andsiltstones with black to medium bedded sandstones,locally containing abundant plant macrofossils
100 meters
Alternation of thick and medium beddedsandstones with traces of tree trunks
Alternation of grey shales, siltstones andsandstones with rare plant macrofossils
Thick bedded sandstones
Alternation of greenish gray shales, siltstonesand medium bedded gray sandstones with rareplant macrofossils
Thick bedded sandstones with fossil fragments
Alternation of sandstones, dark siltstones, coaly shales and coal seams (coal seams D9-D11) with abundant plant macrofossils
Thick bedded gray sandstones
Alternation of dark siltstones, coaly shales and coal seams (coal seams D2, D4 and D6) with abundant plant macrofossils
Medium to thick bedded greenish gray sandstones withinvertebrate fossils and sedimentary structures
Pale yellow shales
Medium to thick bedded yellowish oolithic limestones withabundant brachiopods, pelecypods, gastropods, ammonites
STA
GE
FO
RM
AT
ION
LIT
HO
LOG
Y
SU
ITE
(ME
MB
ER
)
B
a
t
h
o
n
i
a
n
D
a
s
h
t k
h
a
k
H
o
j
e
d
k
G
u
m
r
u d
B
a
j
o
c
i
a
n
Bab
nizu
Badamu
Cal. Bidu AsadAbad
DESCRIPTION
Fig. 2. Stratigraphical column of the section through the Hojedk Formation, North of Kerman, with the strata in which the fossils were found indicated by the wood symbol.
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IAWA Journal, Vol. 26 (4), 2005492 493Poole & Mirzaie Ataabadi — Jurassic conifer woods
Formation is dated as Toarcian–Lower Bajocian (Seyed Emami 1971) and Aghanabati (1998) concluded that the Hojedk Formation, covering an area from Southeast Central to East Central Iran, must have been deposited during the Middle Bajocian–Lower Batho-nian. Therefore, the fossil wood, found in the lower horizons of the Hojedk Formation, is probably Middle to Late Bajocian in age. This paper describes two gymnospermous taxa as yet undescribed from these deposits and thus increases the diversity of the flora preserved within the Hojedk Formation. This paper forms one of a series of publications focusing on the palaeobotany (pre-dominantly leaf floras) of this Formation.
MATERIAL AND METHODS
Two specimens of fossil wood, one pyritized (EUGM–PC1574P) and one substituted with iron oxide (EUGM–PC1565P), deposited in the Palaeobotanical Collection at the Esfahan University Geology Museum, were collected from the Pabedana coalmine to the north of Kerman (Fig. 1). They originate from coal horizons D2–D4 (Fig. 2) of the Hojedk Formation which comprise the famous coal seams of this area. The wood speci-mens were thin sectioned along three planes, transverse section (TS), radial longitudinal section (RLS) and tangential longitudinal section (TLS) and studied using transmitted light microscopy. Descriptions follow the terminology of the IAWA Committee (2004) wherever possible. The material described has been referred, rather than assigned, to fossil species at this stage because not only are fossil conifer wood species ill-defined but both quantitative and qualitative anatomical characters can vary greatly throughout one tree (cf. Chapman 1994; Wheeler & Baas 1998; Falcon-Lang & Cantrill 2000) and furthermore each xylotype described below is only represented by one specimen.
RESULTS
XENOXYLON Gothan 1905Holotype: Xenoxylon latiporosum (Cramer) Gothan 1905Xylotype: Xenoxylon cf. latiporosum (Cramer) Gothan 1905
Material — This taxon is described from a single specimen, EUGM–PC 1565P. Description — An isolated piece of homoxylous secondary xylem (ʻtracheidoxylʼ, Creber 1972), with a total diameter of 4 cm and length of 10 cm, with 25 growth rings, composed predominantly of tracheids (Fig. 3). Parenchyma present and diffuse (Fig. 3). Growth increments predominantly narrow (c. 0.5 mm) although wider towards the outer edge (2.75 mm) and centre (5.5 mm) of the specimen. Ring boundaries distinct but de-marcated by only 1–2 layers of latewood tracheids, rectangular in cross section, thick- or thin-walled (double wall thickness 7–30 μm) when not compressed (Fig. 3). Latewood radial and tangential mean diameters measuring 18.5 μm (range: 12.5–25 μm) and 49 μm (range: 37.5–65 μm) respectively. Tracheids in the earlywood are thin-walled (double radial wall thickness 6–17 μm), regularly arranged in radial rows, square-rectangular in cross section with radial and tangential mean diameters measuring 51 μm (range:
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IAWA Journal, Vol. 26 (4), 2005494 495Poole & Mirzaie Ataabadi — Jurassic conifer woods
Fig. 3–9. Photomicrographs of Xenoxylon cf. latiporosum (EUGM – PC1565P). – 3: TS showing nar-row growth increments with subtle ring boundaries. – 4: TLS showing relatively low rays. – 5: RLS showing a typical ray. – 6: TLS. – 7: Window-like cross-field pits in RLS. – 8: Intertracheary pitting in LS. – 9: Window-like cross-field pits in RLS. — Scale bar = 200 μm in Fig. 3; 100 μm in Fig. 4; 50 μm in Fig. 5; 25 μm in Fig. 6; 12.5 μm in Fig. 7; 10 μm in Fig. 8; 15 μm in Fig. 9.
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IAWA Journal, Vol. 26 (4), 2005494 495Poole & Mirzaie Ataabadi — Jurassic conifer woods
37.5– 67.5 μm) and 57 μm (range: 42.5–72.5 μm) respectively. No spiral thickenings present. Intertracheary pits (Fig. 8) on radial walls uniseriate, bordered, vertically con-tiguous, elliptical 15 –17.5 μm in diameter, with small round-oval apertures, of the abietinoid type (cf. Philippe 1995) usually separated by a distance of at least a quarter of the diameter of the radius if not adpressed, flattened; extremely rare subopposite biseriate pits. Pits on the tangential walls are rare but small, circular, bordered and up to 10 μm in diameter. Cross-field pits (Fig. 7 & 9) are predominantly solitary, occasion-ally 2, possibly even 3, seemingly of the large, simple, window-like fenestriform type (feature 90, IAWA Committee 2004) (cf. the normal or circopore oopore of Philippe 1995) throughout the rays. Rays (Fig. 4 & 5) are homocellular, uniseriate, (1) 2–8 cells high, with no differentiation in shape of the marginal cells. Pith region absent. This speci-men is from the inner portion of a branch/ trunk as deduced from the growth ring cur-vature. One branchlet / twig/ leaf trace can be seen running through at least 21 growth rings. Systematic affinities — Specimen EUGM–PC1565P is characterised by seemingly large, simple, circular cross-field pitting and contiguous uniseriate flattened intertrache-ary pitting on the radial walls. This pitting is characteristic of Xenoxylon, a Mesozoic northern hemisphere xylotype with uncertain systematic position (cf. Philippe & Théve-nard 1996). When this material is compared with other woods from this geographical region (Table 1), there are similarities (e.g. window-like cross-field pits, uniseriate inter-tracheary pits that are flattened when in contact) to Xenoxylon latiporosum from Tazareh (Nadjafi 1982) and X. barberi from Ferizi (Schweitzer & Kirchner 1996). However, when the comparison is widened to include other material from other regions, this wood is more similar to X. latiporosum described by Suzuki and Terada (1992) from Japan (see Table 2). Thus, this specimen is referred to Xenoxylon latiporosum.
AGATHOXYLON HartigHolotype: Agathoxylon cordaianum Hartig 1848Xylotype: Agathoxylon sp.
Material — This taxon is described from a single specimen, EUGM–PC 1574P. Description — An isolated piece of homoxylous secondary xylem (ʻtracheidoxylʼ, Creber 1972) (Fig. 10), measuring 15 cm in diameter and 20 cm in length. Growth rings present. At least three, wide (4–5 mm) growth-ring increments can be seen with distinct growth ring boundaries and a gradual transition from earlywood to latewood within the growth ring (Fig. 13). Parenchyma present as rare, diffuse cells. Latewood relatively wide, tracheids more or less rectangular in cross section, generally thick-walled (double radial wall thickness 12.5–25 μm) with respect to the tracheid diameter. Latewood radial and tangential mean diameters measuring 20 μm (range: 12.5–25 μm) and 31.5 μm (range: 22.5–40 μm) respectively. Tracheids in the earlywood, thin-walled (12.5–20 μm), regularly arranged in radial rows, rectangular in cross section (Fig. 13) with mean tangential and radial diameter measuring 37 μm (range: 27.5–45 μm) and 60 μm (range: 47.5–75 μm) respectively. Tracheids occasionally with features remi-
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IAWA Journal, Vol. 26 (4), 2005496 497Poole & Mirzaie Ataabadi — Jurassic conifer woodsTa
ble
1. S
umm
ary
of th
e an
atom
ical
cha
ract
ers
of th
e Ju
rass
ic f
ossi
l woo
ds p
revi
ousl
y de
scrib
ed f
rom
the
Iran
-Afg
hani
stan
geo
grap
hica
l re
gion
. – E
W =
ear
lyw
ood;
LW
= la
tew
ood;
— =
not
pro
vide
d.
Cupr
essin
oxyl
on o
rient
ale
Mes
embr
ioxy
lon
sp.
Xeno
xylo
n ba
rber
i Pr
otot
axox
ion
feriz
iens
e
loc
ality
Is
hpus
hta
(Afg
hani
stan
) Is
hpus
hta
(Afg
hani
stan
) Is
hpus
hta
(Afg
hani
stan
) Fe
rizi (
Iran
)
age
Sa
igha
n se
ries,
Dog
ger
Saig
han
serie
s, D
ogge
r Sa
igha
n se
ries,
Dog
ger
Shem
shak
Fm
, Dog
ger
ref
eren
ce
Sew
ard
1912
Ja
cob
& S
hukl
a 19
55
Schw
eitz
er &
Kirc
hner
199
6 Fa
khr &
Mar
guer
ier 1
977
in
Fakh
r 197
7
pre
serv
atio
n si
licifi
ed
silic
ified
—
si
licifi
ed
gro
wth
ring
s na
rrow
late
woo
d na
rrow
0.
25–2
.1 m
m w
ide;
103
in
indi
stin
ct, n
umer
ous,
0.6–
3 m
m
5.5
cm
wid
e
tra
chei
ds
—
—
EW (2
3) 4
8–57
μm
; 22
–50
μm
× 1
5–38
μm
LW
48
μm
res
in b
odie
s —
pr
esen
t ab
sent
ab
sent
ray
s un
iser
iate
, rar
ely
part
unis
eria
te, s
omet
imes
par
t un
iser
iate
, hom
ocel
lula
r; un
iser
iate
, rar
ely
part
bise
riate
;
bise
riate
; 1–2
0 (2
5–30
) bi
seria
te; 3
–25
(38–
40)
2–12
cel
ls h
igh
(40–
270
μm
) 1–
13 c
ells
hig
h; ta
ll m
argi
nal
ce
lls h
igh
cells
hig
h
cells
ind
entu
res
—
—
—
abse
nt
cro
ss-fi
eld
pits
po
ortly
pre
serv
ed; 1
–4,
poor
ly p
rese
rved
; 1–2
and
us
ually
1, r
arel
y 2,
ova
l us
ually
1, r
arel
y 2–
3 in
mar
gina
l
?bor
dere
d, ?
oopo
res
fairl
y la
rge
oopo
res,
bord
ered
ce
lls; o
culip
ores
, wea
kly
bord
ered
tra
chei
d pi
ts
RLS
bis
eria
te, o
ppos
ite,fl
at-
unis
eria
te, b
orde
red,
irre
gu-
RLS
bor
dere
d, u
nise
riate
, con
- R
LS b
orde
red,
mai
nly
unis
eria
te
tene
d w
hen
in c
onta
ct; s
ome-
la
rly d
istri
bute
d on
radi
al
tiguo
us, fl
atte
ned
or sp
aced
, (c
ontig
uous
or s
pace
d), r
arel
y
times
uni
seria
te, o
ccas
iona
l w
alls
, inf
requ
ently
bis
eria
te
roun
d; p
roto
pino
id
bise
riate
(alte
rnat
e or
opp
osite
)
grou
ps o
f 3 T
LS u
nise
riate
, an
d op
posi
te; p
odoc
arpo
id
or
mix
ed
cont
iguo
us in
LW
par
ench
yma
pres
ent
pres
ent
pres
ent
pres
ent
tyl
oses
—
—
—
—
bar
-like
orn
amen
tatio
n —
?b
ars o
f San
io
—
—
spi
ral t
hick
enin
g —
—
se
emin
gly
pres
ent b
ut p
rese
r- tru
e th
icke
ning
s pre
sent
, som
e-
vatio
n fe
atur
e tim
es d
oubl
e
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IAWA Journal, Vol. 26 (4), 2005496 497Poole & Mirzaie Ataabadi — Jurassic conifer woods T
able
1 c
ontin
ued
Xeno
xylo
n “M
etat
axod
ioxy
lon
Prot
otax
oxyl
on
“Pro
tosc
iado
pity
oxyl
on
“Pro
topi
noxy
lon
latip
oros
um
irani
cum
” “p
ersic
um”
bour
eaui
” fa
khrii
”
loc
ality
Ta
zare
h (I
ran)
Ta
zare
h (I
ran)
Sa
ngro
ud (I
ran)
Ta
ch (I
ran)
Fe
rizi (
Iran
) a
ge
Lias
Ju
rass
ic
Jura
ssic
Ju
rass
ic
Jura
ssic
ref
eren
ce
Nad
jafi
1982
N
adja
fi 19
82
Nad
jafi
1982
N
adja
fi 19
82
Nad
jafi
1982
pre
serv
atio
n si
licifi
ed
silic
ified
—
si
licifi
ed
silic
ified
gro
wth
ring
s nu
mer
ous,
dist
inct
, 1–2
.5
num
erou
s, di
stin
ct, 2
–5
indi
stin
ct, c
. 4 m
m
dist
inct
, 2 m
m, l
atew
ood
num
erou
s, di
stin
ct, 4
–6 m
m,
m
m, l
atew
ood
1–3
cells
m
m, l
atew
ood
1–2
cells
1–2
cells
la
tew
ood
1–3
cells
tra
chei
ds
LW 6
–30
(42)
× (9
) 18–
LW
5–2
3 ×
23–4
8 μ
m
LW 7
–22
× 26
–33
(74)
μm
LW
4–1
8(–3
0) ×
(11)
18–
LW
5–1
4 ×
23–3
9 μ
m
48 (5
4) μ
m
EW 3
2–46
× 3
4–46
μm
EW
26–
48 ×
18–
52 μ
m
44 (5
5) μ
m
EW 2
8–62
× 2
3–51
μm
EW
30–
66 ×
12–
54 μ
m
EW 2
6–67
× 2
6–66
μm
res
in b
odie
s ab
sent
ab
sent
ab
sent
ab
sent
pr
esen
t as v
ertic
al c
anal
s
in L
W r
ays
unis
eria
te, 1
–10
cells
un
iser
iate
, par
t bis
eria
te
unis
eria
te, r
arel
y bi
seria
te
unis
eria
te, p
art b
iser
iate
un
iser
iate
, 1–2
2 ce
lls h
igh
(2%
), 1–
24 c
ells
hig
h (1
%),
1–22
cel
ls h
igh
(2%
), 1–
18 c
ells
hig
h i
nden
ture
s —
—
—
—
pr
esen
t c
ross
-fiel
d pi
ts
1, o
ccas
iona
lly 2
fene
s-
1–2
(3–4
) tax
odio
id
1 (2
–3),
taxo
dioi
d, o
culi-
1–
3 (4
), ta
xodi
oid,
ocu
li-
1–4,
taxo
dioi
d
trifo
rm p
its o
ccup
ying
pore
s; ?
oopo
re
pore
s; o
opor
es
the
who
le c
ross
-fiel
d ar
ea
tra
chei
d pi
ts
RLS
uni
seria
te, c
ontig
u-
RLS
uni
seria
te (7
9%),
RLS
uni
seria
te, fl
atte
ned
RLS
uni
seria
te, b
iser
iate
R
LS u
nise
riate
(94%
),
ous,
flatte
ned,
rare
ly b
i- fla
ttene
d; b
iser
iate
(21%
) (5
5%);
circ
ular
, con
tigu-
(7
%),
arau
caro
id
flatte
ned;
bis
eria
te (6
%)
se
riate
, alte
rnat
e al
tern
ate,
ara
ucar
oid
ous (
20%
); bi
seria
te a
lter-
al
tern
ate,
ara
ucar
oid
TL
S lo
cally
pre
sent
nate
(20%
); ab
ieto
id (4
%)
par
ench
yma
unce
rtain
due
to th
e no
t obs
erve
d pr
esen
t ab
sent
ab
sent
pr
esen
ce o
f tyl
oses
and
sept
ae t
ylos
es
—
—
—
—
pres
ent
bar
-like
orn
amen
- —
—
fin
e ho
rizon
tal ʻ
calli
troid
—
—
ta
tion
bars
spi
ral t
hick
enin
g —
—
pr
esen
t, do
uble
fa
lse
thic
keni
ngs
—
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IAWA Journal, Vol. 26 (4), 2005498 499Poole & Mirzaie Ataabadi — Jurassic conifer woodsTa
ble
2. S
umm
ary
of th
e an
atom
ical
cha
ract
ers
of th
e w
oods
des
crib
ed h
erei
n an
d th
e sp
ecie
s of
Xen
oxyl
on w
ith w
hich
spe
cim
en E
UG
M-
PC15
65P
show
s gre
ates
t ana
tom
ical
sim
ilarit
y. –
EW
= e
arly
woo
d; L
W =
late
woo
d; —
= n
ot p
rovi
ded.
EUG
M-P
C15
74P
EUG
M-P
C15
65P
Xeno
xylo
n la
tipor
osum
loc
ality
K
erm
an Ir
an
Ker
man
Iran
Ja
pan
age
H
ojed
k Fm
, Dog
ger,
Jura
ssic
H
ojed
k Fm
, Dog
ger,
Jura
ssic
Ea
rly C
reta
ceou
s
ref
eren
ce
here
in
here
in
Suzu
ki &
Ter
ada
1992
pre
serv
atio
n py
ritiz
ed
iron
oxid
e —
gro
wth
ring
s di
stin
ct w
ith a
brup
t LW
-EW
bou
ndar
y,
dist
inct
, 0.5
–5.5
mm
late
woo
d na
rrow
di
stin
ct, 0
.25–
1.35
mm
late
woo
d na
rrow
,
4–5
mm
late
woo
d w
ide
with
gra
dual
1–
2 ce
lls
2–se
vera
l cel
ls
EW-L
W tr
ansi
tion
tra
chei
ds
EW: 2
7.5–
45 μ
m ×
47.
5–75
μm
EW
: 37.
5–67
.5 ×
42.
5–72
.5 μ
m
EW: 6
0–80
× 4
0–80
μm
LW
: 12.
5–25
μm
× 2
2.5–
40 μ
m
LW: 1
2.5–
25 ×
37.
5–65
μm
LW
: 20–
30 ×
40–
80 μ
m
res
in b
odie
s ab
sent
ab
sent
da
rk o
cclu
sion
s in
ray
cells
ray
s un
iser
iate
, hom
ocel
lula
r, 2–
32 c
ells
un
iser
iate
, hom
ocel
lula
r, (1
) 2–8
cel
ls
unis
eria
te, 1
–10
cells
hig
h
high
, rar
ely
part
bise
riate
hi
gh
ind
entu
res
pres
ent
abse
nt
abse
nt
cro
ss-fi
eld
pits
up
to 6
, bor
dere
d, c
upre
ssoi
d 1(
–2),
win
dow
-like
(oop
ore)
1(
–2),
larg
e, w
indo
w-li
ke (o
opor
e)
tra
chei
d pi
ts
RLS
: rar
ely
unis
eria
te a
nd tr
iser
iate
, R
LS: u
nise
riate
, elli
ptic
al b
orde
red
RLS
: bor
dere
d, e
llipt
ical
, rou
nd a
pertu
res,
m
ainl
y bi
seria
te, a
ltern
ate,
occ
asio
n-
TLS:
ver
y oc
casi
onal
smal
l, ci
rcul
ar,
cont
iguo
us to
smal
l, ci
rcul
ar, s
catte
red
al
ly su
bopp
osite
bo
rder
ed
TLS:
smal
l, ci
rcul
ar, b
orde
red
TL
S: ra
re u
ni–b
iser
iate
par
ench
yma
rare
pr
esen
t ab
sent
tyl
oses
ab
sent
pr
esen
t pr
esen
t
bar
-like
orn
amen
tatio
n ab
sent
ab
sent
ab
sent
spi
ral t
hick
enin
g —
ab
sent
—
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IAWA Journal, Vol. 26 (4), 2005498 499Poole & Mirzaie Ataabadi — Jurassic conifer woods
Fig. 10–17. Photomicrographs of Agathoxylon sp. (EUGM – PC 1574 P). – 10: TS showing distinct rays. – 11: TS showing the remains of the pith. – 12: TLS showing rays. – 13: TS of growth ring boundary. – 14: RLS of cupressoid cross-field pits. – 15: RLS showing alternate intertracheary pitting. – 16: RLS showing pseudo-thickenings of the tangential wall of the ray cells. – 17: LS tracheid pitting and features caused by fungal decay of the cell wall that could be confused with spiral thickenings. — Scale bar = 50 μm in Fig. 10; 100 μm in Fig. 11–13; 15 μm in Fig. 14; 25 μm in Fig. 15 & 17; 10 μm in Fig. 16.
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IAWA Journal, Vol. 26 (4), 2005500 501Poole & Mirzaie Ataabadi — Jurassic conifer woods
niscent of spiral thickenings, although these are more likely to be features caused by fungal decay in the S2 layer of the cell wall (Fig. 17; cf. Kräusel & Jain 1964) or cell wall cavities associated with compression wood. Intertracheary radial wall pits (Fig. 15) are bordered, rarely uniseriate (< 10%), predominantly biseriate (c. 85%), rarely triseriate (< 10%), predominantly polygonal and alternate when multiseriate, biseriate pitting is occasionally subopposite, 10.5–17.5 μm in diameter, with small round-oval apertures; uniseriate pitting is confined to the narrow latewood tracheids with pits usually contigu-ous, occasionally scattered. Multiseriate pitting occurs in the wide earlywood tracheids. Tracheid tangential wall pitting is rare but uni- to biseriate when present. Cross-field pits (Fig. 14) are bordered and of the ʻcupressoid type (feature 93, IAWA Committee 2004) (cf. cupressoid oculipore of Philippe 1995) with up to six pits per cross field in both the marginal and central part of the rays. Cross-field pits poorly preserved but gen-erally more or less circular when not touching; when adpressed, pits appear to be alter-nate. Rays are uniseriate, 2–32 cells high, rarely locally biseriate, with biseriate regions up to two cells in length with no difference between the cells of the margins and body (Fig. 12). Tangential walls of some ray cells appear to have thickenings (Fig. 16) but could represent pseudo-thickenings formed by iron sulphide bacterial activity or, alter-natively, checking associated with compression wood. Pith is composed of thin-walled parenchyma cells (Fig. 11). Systematic affinities — Alternate biseriate tracheidal pitting, which is characteristic of specimen EUGM–PC1574P, is also found in the specimens of Prototaxoxylon and the invalid genera ʻMetataxodioxylonʼ, ʻProtosciadopityoxylon and ʻProtopinoxylon from the Alborz region of Iran (Nadjafi 1982; see Table 1). However, similar cross-field pits in combination with alternate tracheidal pitting occur only in Prototaxoxylon specimens from Alborz (Fakhr & Marguerier 1977 in Fakhr 1977; Nadjafi 1982). The differences between the specimens previously assigned to Prototaxoxylon and the specimen described herein include the abundance of biseriate and presence of triseri-ate tracheidal pitting, up to four cupressoid pits in the cross fields, and the absence of true spiral thickenings in EUGM–PC1574P; therefore this material cannot be assigned to this xylotype.
Alternate and subopposite radial intertracheary pitting and cupressoid crossfield pitting is also characteristic of araucariaceous fossil wood and wood of Brachyoxylon Hollick et Jeffrey (1909). However, the intertracheary pits are predominantly polygonal and alter-nate when multiseriate and the cross-field pits appear alternate when adpressed (although better preserved material will enable the determination of the precise pit characters of the cross fields) such that this fossil bears greater similarity to araucariaceous fossil woods than to those assigned to Brachyoxylon. Therefore this specimen has been as-signed to the xylotype Agathoxylon erected for woods with araucarian radial wall pit-ting and araucarioid cross-field pits. No attempt has been made to place this specimen within an existing species since there are no other woods of this type yet found from Iran and the morphogenera for araucariaceous fossil wood are in need of revision (Bamford & Philippe 2001).
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IAWA Journal, Vol. 26 (4), 2005500 501Poole & Mirzaie Ataabadi — Jurassic conifer woods
DISCUSSION
During the Middle Jurassic the habitats of Iran and Afghanistan were located on the southern coastal edge of the Middle Asian province of the Euro-Sinian region (Vakh-rameev 1991). Fakhr (1977) suggested that the Middle Jurassic basins of North Central Iran and North Afghanistan, once formed part of a basin-island complex along the coast of Laurasia and that the terrestrial deposits were laid down in the Caucausus–Turkistan geosyncline which branches into the eastern part of Central Iran. Studies by Repin (1985) concluded that the Kerman area was one such island situated in close proxim-ity to other land masses, which were periodically united into a large basin. Middle Jurassic vegetative remains were deposited in this basin and subsequently became the coal deposits of Central Iran. A study by Seyed Emami (1971) also considers the Kerman area to have been sited on the southern margin of Laurasia due to similarities between the terrestrial Rhäto–Jurassic deposits in North and Central Iran and those of Afghanistan, Turkmenistan and Armenia which are known to have been situated on the southern margin of Laurasia. However, Vozenin-Serra and Taugourdeau (1985) con-sider the Kerman area to have been connected to the northern part of Gondwana with migration from northern Gondwana prevented by the large expanse of water between India and this island complex (e.g. Delle 1967). The composition of the Jurassic floras within the Middle Asian province (extending south from the Karagandar-Turgay region in the north to the coast of the Jurassic sea in the south; Vakhrameev 1991) changed markedly along the north-south transect (Sykstel 1954). Floras of the Middle Jurassic are relatively species-rich, with a diversity of lycopods, horsetails, ferns and gymnosperms (Gomolitzky & Khudayberdyev 1976). Palaeobotanical evidence derived from the coal deposits of the Georgia-Transcaucasus region to the north indicates a typical rich and diverse Middle Jurassic flora similar to that found in the European floristic province (Barale et al. 199; Vakhrameev 1991). However, components characteristic of coeval floras growing in the Central Asian and Indian provinces were also present (Delle 1960; Junusov 1975). Understorey ferns (e.g. Coniopteris, Cladophlebis), ginkgos and conifers (e.g. Araucarioxylon, Podocarpoxy-lon and Xenoxylon) (Delle 1960; Barale et al. 1991 and references therein) occupied higher elevations whereas the cycadophytes (especially Nilssonia) and woody ferns grew at lower elevations, giving way to horsetails and neocalamites in moist swampy zones. A similar floral composition can also be found in Turkmenistan to the east (Delle 1960). To the south, the floras of northern and south-eastern (Kerman) Iran covering low-land areas adjacent to the marine basin can be referred to the Transcaspian subprovince (Vakhrameev 1991). The gymnosperm component of the vegetation from the Lower Jurassic comprised members of the Voltziaceae/Taxodiaceae (including Xenoxylon), Cheirolepidiaceae and Palissyaceae (e.g. Schweitzer & Kirchner 1996) with a decreasing Czekanowskiales (Delle 1967; Vakhrameev 1991; Barale et al. 1991). Palaeobotanical studies (Mirzaie Ataabadi 2002) of the Middle Jurassic macrofloras from the Kerman area of Iran show up to 50% similarity with those of Laurasian floras, the Transcauca-sus (Georgia; Delle 1967), East Urals (Russia; Genkina 1963) and the Saighan Series
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IAWA Journal, Vol. 26 (4), 2005502 503Poole & Mirzaie Ataabadi — Jurassic conifer woods
of North Afghanistan (Jacob & Shukla 1955) which also form part of the Euro-Sinian floral region (Vakhrameev 1991). By the Middle Jurassic the Czekanowskia and the Ginkgoales had become scarce in terms of number and diversity and the ancient Pinaceae were almost completely absent (Delle 1967; Vakhrameev 1991; Barale et al. 1991). The Lower-Early Middle Jurassic arboreal vegetation, as determined from the wood record, included Prototaxoxylon (Fakhr & Marguerier 1977 in Fakhr 1977; Nadjafi 1982), Xenoxylon (Nadjafi 1982, herein), Agathoxylon, and three other coniferous taxa assigned to the (unpublished and thus invalid-) genera ʻProtosciadopityoxylonʼ, ʻProtopinoxylon and Metataxodioxylonʼ (Nadjafi 1982). Xenoxylon survived into the Middle Jurassic and together with Agathoxylon and characteristic understorey ferns (such as Klukia, Eboracia and Coniopteris) and cycads, e.g. Nilssonia (Vakhrameev 1991) contributed to a more diverse and species-rich vegetation relative to the preced-ing Lower Jurassic flora (Gomolitzky & Khudayberdyev 1976). The presence of Xenoxylon, an essentially northern hemisphere (Laurasian) fossil wood taxon from the Middle Jurassic of Central Iran, may help substantiate the hy-pothesis that Central Iran was situated in the Middle Asian province of the Euro-Sinian floral region during the Middle Jurassic. However, the presence of a wood type in the Kerman Basin similar to Agathoxylon, which is distributed in both the northern and southern hemispheres, may indicate a connection enabling floral exchange between the Iran–Afghanistan region (forming the Iranian–Afghani Plate) and the northern edge of Gondwana at this time. Detailed studies of this and other palaeofloras will help further our understanding of the palaeogeography and vegetation dynamics across this region during the Mesozoic. The extensive survey of the fossil record of Xenoxylon by Philippe and Thévenard (1996) across the northern hemisphere shows this genus to favour cool, seasonally wet areas. The authors conclude that Xenoxylon favoured, and was later confined, to climates with a mean annual temperature of 5–15 °C during the Early Cretaceous. The presence of Xenoxylon wood in Pabedana along with other moisture loving plants such as bryophytes, equisetales and ferns (Schweitzer 1978; Schweitzer et al. 1997) and the relative abundances of Bennettites, Coniopteris and Nilssonia suggest that the palaeoenvironment was cool and at least locally moist – an observation which would agree with the Kerman area being an island at this time (Repin 1985) and evidenced by the vast coal deposits. The presence of growth rings in both specimens described here may suggest some seasonality. However, in the Xenoxylon specimen the rings are relatively indistinct rather than well-defined. Well-defined rings in Xenoxylon led some authors to believe that this taxon was deciduous (Philippe & Thévenard 1996 and references therein). Further growth ring analysis was not undertaken to explore this climatic aspect since there is debate as to the reliability of using such palaeoden-drochronological methods for fossil material (e.g. Brison et al. 2001).
CONCLUSION
Two coniferous taxa, Xenoxylon and Agathoxylon, are recorded from Middle Jurassic deposits of the Kerman Basin, Iran. This area was a lowland coastal zone located on the southern margin of Laurasia in the Middle Asian province of the Euro-Sinian floral
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IAWA Journal, Vol. 26 (4), 2005502 503Poole & Mirzaie Ataabadi — Jurassic conifer woods
region at this time. These specimens represent the first record of permineralised wood from these Middle Jurassic coal bearing deposits and provide information pertaining to the arboreal component of the vegetation. Additional studies focusing on fossil woods from sites across Iran and Afghanistan, the most southerly of all Laurasian localities, in combination with syntheses of the floras already described will further our understanding of the nature, affinities and palaeogeographical distribution of the Jurassic vegetation that grew along the edge of the Laurasian landmass. Moreover, palaeobotanic studies of fossil floras will enhance understanding of the spatial and temporal dynamics of the plate(s) on which they rode.
ACKNOWLEDGMENTS
We thank Dr Mohammad Sadegh Fakhr (Laboratory of Paleobotany, University of Tehran, Iran), Professor Han van Konijnenburg-van Cittert (Utrecht University), Professor Elisabeth Wheeler and especially Dr Marc Philippe (Université Claude Bernard, Lyon) for providing relevant literature, helpful discussions and comments that have improved this manuscript. Otto Stieckma (Utrecht University) is acknowledged for sectioning the fossil material. We are also grateful to Vachik Hairapetian (Islamic Azad University, Khorasgan branch, Esfahan, Iran) for his help in drawing the maps and the stratigraphic column used in this paper, and Iwona Rejniewicz for translating the Russian literature.
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