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A short history about the evolution of gymnosperms

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 A short history about the evolution of gymnosperms
Michael Wachtler
P. P. Rainerstrasse 11, 39038 Innichen, Italy; E-mail: michael@wachtler.com

The slow evolution of life is an indisputable scientific principle. However, it seems that all of
today’s widespread plant tribes were already present at the Carboniferous–Permian boundary
and that their common ancestors can be largely dated back to the Devonian. This means that
the origins of gymnosperms and angiosperms should be found there. In the earliest Permian
(and probably before) all the features of the gymnosperms were highly developed; hence,
in the following 300 million years they changed only slightly. The straight evolution line to
modern-day Araucariaceae was clearly defined in the Permian. The one-seeded genus Ortiseia
had the same structural appearance as modern-day Araucarias, while wing-seeded Majonica
represented the firs. Also the Pinus ancestor Valentinia was widespread in the Early Permian,
as well as the typical three-, four- (or more) lobed seed scales of the Voltziaceae, which were
potential progenitors of today’s Cupressaceae. The typical blueprint of the Ginkgoales was in
the same way already defined in the Paleozoic (Baiera), and even fully evolved Cycadales like
Bjuvia, Taeniopteris or Nilssonia could be encountered. The cycad-like Wachtleropteris or ru-
dimentary conifer Perneria can be regarded as real living fossils in the Permian. The double Y-
lobed leaves and seed scales of Perneria are the last memory of a common plant origin in the
Devonian, where all tribes hold this feature. This blueprint can be observed in its hidden form
in all modern-day floras.
Online: December 2016
Key words: Gymnosperms, Coniferales, Ginkgoales, Cycadophyta

An Early Permian landscape that evidences the different and fully evolved gymnosperm families at that time: Left
Ortiseia male and female cone (Araucaria-progenitor), in the middle Wachtleropteris with fertile organ (Cycadalean-
ancestor) and Baiera (Ginkophyta), right Valentinia (Pinus-progenitor), above right Majonica male and female (Abi-
etaceae-ancestor) and above in the middle Seymourina (Voltziaceae), a thought Cupressaceae-progenitor.

Michael Wachtler: The evolution of gymnosperms in the Triassic

3
 The most important feature used to the Carboniferous–Permian, two-, three- or
distinguish the distinct conifer genera is their more-lobed seed scales with seeds attached
female fructification. There we encounter on the upper side of fertile leaves. Usually
a problem that originates in the taxonomy the scales were surrounded by dwarfish
of modern conifers. Usually today’s Pinales sterile, tapered micro-leaves and especially
are grouped in the Pinaceae, Araucariaceae, in the Permian with one the seed scale
Po d o c a r p a c e a e , Sciadopityaceae, largely projecting sterile bract (Wachtlerina,
Cupressaceae, Cephalotaxaceae and Majonica, Seymourina, Pseudovoltzia). They
Taxaceae families. However, what do the were accompanied by male organs consisting
big five Pinales groups (namely, Pinaceae of cones with small tapered leaves and
– with its four sub-families; the Pinoideae pollen-clusters on the lower side. However,
comprising the genus Pinus – with more if all these plants, holding common features,
than 110 species; the Piceoideae – with were largely separated on the Carboniferous–
Picea; the Laricoideae – with Cathaya, Larix Permian border, where should we look for
and Pseudotsuga; and the Abietoideae – their common grandfather?
with Pseudolarix, Abies, Cedrus, Keteleeria,
Nothotsuga and Tsuga) have in common? The origin of all gymnosperms
In the Early Permian the genus Pinus
(Valentinia) and the genus Abies/Picea The ancestor of all gymnosperms must be
(Wachtlerina/Majonica) were already searched for close to the Devonian–Car-
separated and passed through the following boniferous border, which marks the dawn of
300 million years in a markedly different the most primitive plants. In fact, the first
way (Wachtler, 2013, 2015). The same is Psilophytes were a complex of single Y to
valid for all other conifer groups, which were two or four Y-dichotomising fertile and ster-
jumbled together in artificial systems without ile leaves, sometimes also equipped with
consideration of their evolutionary record. minute leaves on the basal parts of the
Around the world we can find, starting from plants, called “emergences”. Therefore, it

The phylogenetic tree of gymnosperms

Taxus Cryptomeria Larix Abies Pinus Zamia Cycas
Picea Araucaria Ginkgo
Cephalotaxus Juniper Pseudolarix 1-2-3-5-8 needles Encephalartos
Torreya Sequoiadendron Stangeria
Cunninghamia

Larix-line
Wachtlerolarix
Abies-line Picea-line
Taxus-Torreya
Farjonia Estellencsia
Kandutschia

Araucaria-line
Cupressus-line Abies-line Ortiseia Ginkgo-line Zamia-Cycads Cycas-Cycads
Voltzia Majonica Northern extinct Baiera Nilssonia Taeniopteris
Taxus-line Pseudoctenis Bjuvia
Ullmannia Southern globe
Metasequoia
line
Pinus-line Pinus-line
Valentinia Valentinia
1-2-3 needles 5 needles

Pinus-line
First Cupressus-line Abies-line Araucaria-line Valentinia Ginkgo-line Zamia-Cycads Cycas-Cycads
Perneria Seymourina Wachtlerina Wachtleropteris Taeniopteris
Ortiseia Baiera Nilssonia
Last Devonian Voltzia Majonica Pseudoctenis Last-cycad Bjuvia
survivor Devonian-
survivor

Main Devonian ancestor

From the fossil-record can be deduced, that the most important tribes were separated just in the Early Permian.
Red: Time-scale of the first appearance.

Dolomythos, 2016
4
 2 cm
1 cm

2 3
1 cm

1 cm
10 cm

1 4 5
Calamophyton primaevum from the Schiffarth Quarry, Lindlar (Middle Devonian) 1. Complete plant (known formerly
as Duisbergia mirabilis); 2–3. Branchlets bearing primitive cones with two sporangia; 4–5. Detail of the archaic spo-
rangia (All Coll. Pohl). Plants like these should be investigated to determine the origin of the gymnosperms.

Michael Wachtler: The evolution of gymnosperms in the Triassic

5
 can be assumed that the megasporophylls
originated from forking telomes, whereas
the microsporophylls derive from modified
emergences. The evolutionary notion of all
gymnosperms with a forefather in the De-
vonian and with rapid initial mutation to all
extant gymnosperm groups, including Cy-
cadales, Ginkgoales and Coniferophyta, can
be understood only from this point of view.
The building concept developed by Rudolf
Florin is no longer an issue. This means that
Permian conifer seed scales evolved as in-
florescences and formed the basis for the
development of all conifers existing today,
through the continuous merging of sterile
bracts and fertile seed scales (Florin, 1938–
1945). Unfortunately, this Walchia theory
was based merely on a few isolated and of-
ten poorly preserved seed scales collected
in the Thuringian Forest; to make matters
worse, the mixed group of specimens was
also taken from different strata and periods. 1cm

The theory of the Cordaitales as the

ancestors of all conifers is not more tenable.
Cordaites was a tree-like plant with long,
tongue-shaped leaves that characterised
the landscapes from the Early Carboniferous
to the Early Permian. Their cone-shaped
reproductive organs gave the impression
that they were somehow related to the
conifers but they do not fit our image of
conifers today. The female organs formed
similar loose, cone-like structures and a high
number of sterile protecting leaves grew
out of the axils of each bract. It was likely
that they acted only as a dead-ending side
branch, as many other plants in the history
of evolution.
The conifer enigma
Unfortunately, we encounter several conifer
lines on the Carboniferous–Permian border
that are connected by some similarities but
also display net differences. Primarily, there
is an unambiguous distinction between
one-seeded, two-winged-seeded and more-
seeded genera. Furthermore, their pollen
cones were remarkably different. Only in
a general context can it be established
that the Voltziaceae, Ortiseiaceae and
Wachtlerinaceae (Majonica) form a relative
allied group of gymnosperms in a system in
the enigmatic Walchiaceae (Wachtler 2013,
2014, 2015). Otherwise, the Pinus conifers
(Valentiniaceae) have a completely different
Devonian Buthotrepis mosellae (Kräusel & Weyland)
Schweitzer from Waxweiler, Germany with its furcating
axis (Coll. Stapf, Nierstein)
origin and are more connected with the
Ginkgoales (Baiera) and the two-seeded
Cycadophyta (Nilssonia, Pseudoctenis).
In order to determine in which ways
the conifers, cycads and ginkgos differ
or are related, we have to follow more
steps. The first step is to establish which
gymnosperms in the Paleozoic can be
accepted as the most primitive. Perneria
thomsonii (Wachtler, 2013), which was
recorded on the Carboniferous–Permian
boundary, has until now been regarded as
the most rudimental conifer. The seed scales
were two- to four-lobed and, interestingly,
their branchlets also hold primitive four-
lobed leaves, a feature never recorded later
but common in Devonian times. However,
Perneria cannot be accepted as the ancestor
of all conifers but could perhaps be just
the last representative of a very primitive
gymnosperm-conifer.
The cycad gymnosperms
Wachtleropteris valentinii represents a ru-
dimental cycad gymnosperm from the Early
Permian. This shrubby plant was equipped
with leaves that extended upwards on a

Dolomythos, 2016
6
stem and branched twice, which is not typi-
pohli can be regarded, because of its
cal for cycads, and each of the leaves split
inchoate and irregular branching system,
into two independent branches. The leaves
as the most primitive ginkgo recorded until
were tongue-shaped and have a pronounced
now, with affinities to some progymnosperm
midrib. The arrangement of the cones on the
plants from the Devonian. Yet it has all of
end of a pinnate leaf identifies them as gym-
the features of real ginkgos: a collar-like
nosperms but it is not possible to draw clear
ring from which the leaves emanate and two
parallels with the Cycadales. Their classifica-
ovules/seeds aggregated together on the
tion is therefore questionable, even though
apical part of modified leaf segments.
they formed a bridge as the “last repre-
Isolated needles can be easily confused
sentatives of a very old and primitive spe-
with the leaves of the most primitive
cies having cycadalean affinities” (Wachtler,
Pinus conifer, Valentinia. Like almost every
2015) from the Devonian until the Permian.
other family of conifers, the Pinoideae
From the Permian and all through the Tri-
had already experienced an explosive
assic we then encounter isolated wedge-
development by the time of the transition
shaped appendices holding two ovules/
from the Carboniferous to the Permian,
seeds on the lower shield surface, known as
which brought them close to the modern-
Olangocarpus. They belong to cycadalean
day pines in a very short time. Strangely
progenitors, like Nilssonia, Apoldia or Pseu-
enough, the multilobed and laciniate
doctenis, and have an amazing resemblance
individual leaves of its Devonian ancestors
to today’s Zamia species.
again played a role in this: they became
Additionally, from the Permian and especial-
longer, forming the slender needles that we
ly in the Triassic, feathered cycad megaspo-
know today. The cones reached their current
rophylls with multiple seeds/ovules attached
appearance during the Early Permian. An
on the lower side of an often closed or semi-
interesting feature of their needles is a
open fruit-blade are found. They have the
striking similarity to the ginkgo leaves of
organ name Dioonites and pertain to the
that period.
other big group of modern-day cycads: the
To date, Valentina wachtleri from the
Cycas family. However, not even these can
Early Permian (Artinskian) is the oldest
be connected in a reasonable time in the
known ancestor of the genus Pinus. The
Carboniferous–Permian with the two-seeded
irregularly forking needles were variegated
cycads. That means that their line-splitting
and grouped in bundles of two to eight.
must also have happened in the Devonian,
The pollen cones were dwarfish, remaining
and, surprisingly, has to be connected with
below 1 cm in size. The seed cones grew
the other main group of today’s plants: the
on a minute leaf/stem, were no more than
angiosperms. It can be supposed that the
2 to 3 cm high and were composed of a
more-seeded Cycas cycads are, in reality,
few megasporophylls arranged in a helix,
primitive angiosperms.
each bearing two short-winged seeds on
Entire male cones, independent of
its surface. The apophysis was keeled with
cycad genus, are described within the
a short umbo, like most of today’s pines.
morphogenus Androstrobus, whereas
In this regard, Valentinia wachtleri can be
isolated microsporophylls of cycadalean
considered as a conifer close to the origins
affinities with pollen sacs on the lower part
of Pinoideae. Astonishingly, in a short time
or inside of a covering roof are classified as
of about 7 million years, the Pinus ancestors
Pizperesia. They represent parts of decaying
diverged. Kungurian Valentinia angelellii
pollen cones and could belong to all possible
still exhibited leaves, irregularly bundled in
cycad ancestors.
groups of one to three. An evolution in the
direction towards extant diploxyl pines is
The amalgation of Gingko and Pinus
suggested. A second pine, coeval Valentinia
Another enigmatic gymnosperm group is cassinisi, grouped its long needles in bundles
the Ginkgoales. Modern-day ginkgo leaves of five, evidencing basal sheaths and being,
are fan-shaped with veins radiating out into in that respect, a potential ancestor of
the leaf blade. But at the Carboniferous– today’s haploxyl pines. The fertile organs
Permian border, their leaves were irregularly of the two Valentinia species are similar,
lobed and needle-like. Early Permian Baiera having rounded hard woody cones with

Michael Wachtler: The evolution of gymnosperms in the Triassic

7
 Milestones in the evolution of the gymnosperms
Carboniferous-Permian (Gzehlian):
Preconiferophyta
Perneria thomsonii: Oldest known conifer.
Leaves on upper part double-bifurcating, basal
needle-like (thought to emerge from the Devo-
nian). Microsporophylls on single leaves (emer-
gences). Megasporophylls upper part, double-
bifurcating, holding one seed on each lobe.

Carboniferous-Early Permian:
Walchiaceae

Ortiseia-line (Pre-Araucarian):
Single leaves, male cones moderately large in
size, seed-scales surrounded by dwarfish leaves
(emergences), one fused wingless seed on two
pseudo-separated scales.
Wachtlerina-line (Pre-Abietaceae): Single
leaves, small male cones, seed-scales sur-
rounded by dwarfish leaves (emergences), two
winged seeds on two pseudo-separated scales.
Voltzia-line (Pre-Cupressaceae): Single
leaves, medium-sized male cones, seed-scales
divided in three or more lobes. From each upper
side, hanging dorsiventrally, is one small-sized
wingless seed.

Early Permian:
Valentiniaceae

Valentinia-line (Pinus-conifers):
Leaves in the Early Permian (Artinskian) grouped
in bundles from 1–8 (Valentinia wachtleri), in
Kungurian one line generating from 1–3 needles
(Valentinia angellelii), others have 5 needles
(Valentinia cassinisi). Pollen cones are dwarfish,
megasporophylls have two short-winged seeds.

Early Permian:
Pre-Cycadaceae

Wachtleropteris valentinii: Oldest known pu-

tative cycad. Leaves double-bifurcating. Micro-
sporophylls are a compound of many microleaves
(Pizperesia). Megasporophylls, called Olangocar-
pus hold two seeds covered by a shield. All ar-
ranged in cones.

Early Permian:
Pre-Ginkgoales

Baiera-line (Ginkgo-ancestors):
Leaves double-bifurcating. Seeds either single or
paired on an elongated leaf (Baiera pohli).

Dolomythos, 2016
8
3 1cm

Evolving trends in
primitive conifers
Pinus-like- Valen-
tinia
1. TRE 483 Valentinia
cassinisi. Holotype
with exemplary fos-
silised fascicle (7
cm long) with five
attached needles en-
cased by a sheath
2. TRE 601. Valentin-
ia cassinisi. Branchlet
2 cm

1cm
3. TRE 548 Valentinia
angelellii. Paratype.
Cone with two seeds
on each scale. 2 1

The approaching of Cycadales, Coniferales and Ginkgoales in Early Permian

Evolving trends in
the most primitive
Ginkgoales.
Note the two- to four-
times dichotomously
branched leaflets that
could reach up to 20
cm in length.
1. TRE 45. Baiera
pohli. Holotype. Leaf
with two seeds on the
upper right side and
one on the left side.
2. TRE 86. Baiera
pohli. Mature leaf
with a basal spur
shoot.

1 1cm 2 1cm

Michael Wachtler: The evolution of gymnosperms in the Triassic

9
 Voltziaceae, Ortiseiaceae, Wachtlerinaceae: Evolving trends of the Walchiales

Wachtlerinaceae. Earliest fossil record on the Carboniferous-Permian border

Wachtlerina bracteata. (Kasimovian and Gzhelian,
Carboniferous-Permian): Niederhausen, Germany (Coll.
Perner). Male cone, female cone, seed scale with long
bract, winged seed.
Remarks: Female cones elongated with one projecting
bract and seed-scale fused together densely in the
middle suggesting only one winged seed
Majonica suessi. (Artinskian, Early Permian): Collio,
Northern Italy (Coll. Valentini). Male cone, female cone
with long bracts, seed scale and two winged seed.
Remarks: Female cones elongated. Seed scales split in
two, two winged seeds for each scale, with one project-
ing sterile bract, additionally to sterile micro-leaves on
the lower part.

Ortiseiaceae. Earliest fossil record on the Carboniferous-Permian border

Ortiseia uhli. Ortiseiaceae (Kasimovian and Gzhelian,
Carboniferous-Permian): Niederhausen, Germany (Coll.
Perner). Male cone, female cone, seed-scale with sur-
rounding sterile leaves, one seed inside.
Remarks: No projecting bract, but seed scales sur-
rounded by sterile microleaves. Only one seed, but split
scale in the middle
Ortiseia daberi. Ortiseiaceae (Kungurian, Early Perm-
ian): Tregiovo, Northern Italy (Coll. Valentini). Male
cone, female cone, seed-scale with surrounding sterile
leaves and one seed inside.
Remarks: Female cones round-bodied, no projecting
bract, but seed scales surrounded by sterile microle-
aves. Only one seed but split scale in the middle

Dolomythos, 2016
10
 Voltziaceae, Ortiseiaceae, Wachtlerinaceae: Evolving trends to the Walchiales

Wachtlerinaceae. Conducting to extant Abietaceae

Majonica alpina (Wuchiapinginan Late Permian) Do-
lomites, Italy (Coll. Wachtler). Male cone, female cone,
seed scale divided with elongated bract-leaves, two
winged seeds.
Remarks: Female cones elongated with one projecting
bract and additional sterile leaves; seed scales split in
two. Two winged seeds for each scale
Picea alba Living. Male cone, female cone, seed scale
holding two winged seeds, winged seed
Remarks: Female cones elongated, sterile bracts
sometimes evidenced (Picea) sometimes clearly visible
(Abies), seed scales fused together. Two winged seeds
for each scale

Ortiseiaceae. Conducting on the Southern hemisphere to the Auracariaceae

Ortiseia leonardii. Ortiseiaceae (Wuchiapinginan Late
Permian) Dolomites, Italy (Coll. Wachtler). Male cone,
female cone, seed scale with surrounding sterile leaves,
seed inside with one seed.
Remarks: Female cones round-bodied, no projecting
bract, but seed scales surrounded by sterile microle-
aves. Only one seed but split scale in the middle.
Araucaria araucana, Living. Scales: Araucaria bid-
willii. Araucariaceae (Recent) Male cone, female cone,
each seed scale holding one seed
Remarks: Female cones round-bodied, seed scales
embedded. Only one seed.

Michael Wachtler: The evolution of gymnosperms in the Triassic

11
 Valentinia wachtleri
Collio
Artinskian

Valentinia cassinisi
Valentinia angelellii Tregiovo
Tregiovo Kungurian
Kungurian

Pinus monophylla Pinus mugo Pinus ponderosa Pinus sylvestris Pinus edulis Pinus cembra

Evolutionary trends in the genus Pinus

The biggest changes occurred in a relatively short time (about 7 million years) in the Early Permian from Artinskian
to Kungurian. In the Artinskian the major 1-2-3-5 needled Pinus-ancestors are just recognizable.

Dolomythos, 2016
12
deep grooves and furrows on the underside
and bearing an umbo at the end. The two-
seeded scales were only a few millimetres in
size and were slightly to heavily winged.
The birth of the Araucariaceae
The one-seeded Ortiseia conifers, which can
be regarded as potential extant Araucaria
ancestors, were already widespread in the
Paleozoic. In the northern hemisphere, a
mainly clear evolutionary line is observed
from earliest Permian (Kasimovian/Gzhelian)
Ortiseia uhli (Wachtler, 2015) to Early
Permian (Kungurian) Ortiseia triumpilina
(Wachtler, 2015), slightly younger Early
Permian (Artinskian), Ortiseia daberi
(Wachtler, 2013) and Late Permian Ortiseia
leonardii (Florin, 1964), Ortiseia vissheri,
Ortiseia jonkeri (Clement-Westerhof, 1984)
and Ortiseia zanettii (Wachtler, 2015). Then,
in the Early Triassic, there was an apparent
and strange disappearance of the Ortiseiaceae
in the northern hemisphere, possibly with
some doubtful last representatives (Ortiseia
1 represent another important gymnosperm
2 aliform seed (Clement-Westerhof, 1987;
3 line, but lying nevertheless on this Abies
The coating of some Permian conifers with sterile mi-
croleaves. 1. Permian Majonica (Abietaceae). The ori-
gin of the aliform seeds originating from microleaves.
2. Permian Ortiseia (Auracariaceae), adaxial-abaxial
and topside-view. Minute leaves surrounding the seed.
3. Permian Ullmannia (Seedberry-conifers) In this
process is to search the building of the fleshly aril of
today’s Taxaceae
Michael Wachtler: The evolution of gymnosperms in the Triassic
collii) on the Balearic islands (Juárez &
Wachtler, 2015) and a continuation in the
southern hemisphere until all Araucaria
species of today. The main distinctive
features of pre-Araucarian conifers are the
round-bodied female Ortiseia cones, which
rarely exceeded 10 cm. Their ovulate scales
were composed of an agglomeration of many
minute sterile leaves completely surrounding
the once-segmented seed blade and the
only one submerged seed. The evolutionary
steps of the Araucaria family can be easily
understood based on this concept. The pollen
cones were slender and about 10 cm long,
although there are some difficulties observing
the typical blueprint of all extant Araucaria
microsporophylls in the Permian (the free-
standing pollen sacs hanging from the apical
lower side dorsiventrally to the main axis).
The microsporophylls of Ortiseia leonardii,
O. vissheri and O. jonkeri from the Dolomites
were probably so densely compressed
together that single pollen sacs are difficult
to observe in fossilised specimens. The Late
Permian Ortiseia zanettii can be accepted
as the first undoubted fossil specimen with
distinctive hanging pollen sacs (Wachtler,
2015).
The triumphal march of firs (Abies)
The winged-seed conifers of the Abies family
group. Carboniferous–Permian Wachtlerina
bracteata (Kasimovian), followed in the Early
Permian by Majonica suessi (Kungurian)
and Cassinisia ambrosii (Artinskian) to Late
Permian (Lopingian) Majonica alpina reveal
a clear evolutionary line characterised by
conifers with dwarfish pollen cones and
two-lobed seed scales – each holding one
Wachtler 2013, 2014, 2015). Almost all
species, like the extant conifer genus Abies,
shed their seed scales entirely and were
further characterised by a projecting bract.
An evolutionary line to today’s widespread
Abietaceae is suggested. A slightly different
group, is supposed for the Early–Middle
Triassic Farjonia campeiae or Middle Triassic
Farjonia presegliei. It seems that, since the
start of the Permian, the genus Ortiseia was
one-seeded on a two-lobed scale, while the
Wachtlerinaceae indeed hold two winged
seeds on an also two-lobed deeply incised
13
 scale. Ortiseia and Majonica have therefore
more parental affinities than previously
thought.
The birth of the spruces (Picea)
From the outside, Abies and Picea conifers
seem to be closely related; however, from
the fossil record this does not seem to be
the case. A distinct feature of all Abies
progenitors from the Permian (Wachtlerina,
Majonica and Cassinisia) and the Triassic
(Farjonia) is the loosening of their single
ovulate scales; also, in addition, in most
cases, a projecting sterile bract. Conifers
corresponding to the present-day Picea or
Keteleeria were recovered with Estellencsia
saezii for the first time from Early–Middle
Triassic (Anisian) layers on the Balearic
Islands in Spain (Juárez & Wachtler, 2015).
These fossils had spreading branches and
elliptical juvenile leaves with a rounded
apex, while adult foliage and leaves on the
main stems were elongated and needle-like.
Female cones were elongated with striate
scales and no visible bract, while male cones
were small in size (Juárez & Wachtler, 2015).
Estellencsia conifers are also recorded in the
Dolomites. It seems that the Picea conifers
separated from the Abies line at the dawn
of the Permian or before. After that period,
both coexisted in the fossil record.
The larches (Larix)
The evolution of the important conifer fam-
ily Larix was in the past shrouded by ques-
tions. It can be now stated that they were
already present from the Early–Middle Trias-
sic. Wachtlerolarix weissii, in particular, has
all the features of today’s larches, such as
small-sized seed cones, needles clustered on
short shoots and small male strobili borne
solitarily from the apex of short shoots. The
origin of the larches’ typical false whorls lies
in a reduction and compression of the nee-
dles of a twig to a bundle. In contrast to the
evolution of the genus Pinus (Valentinia)
with its 2–3–5 regularly assembled needles,
which occurred in the Early Permian, the
larch family has a different development his-
tory also demonstrated by their irregularly
bundled leaves. The larches have their ori-
gin in a reduction and compression from a
shoot to a bundled short shoot. For this rea-
son, the number of leaves in each tuft varies
14
considerably. It seems that over time a re-
duction from normal shoots to short shoots
must have occurred independently several
times, as seen in the extant Cedrus, Larix,
Pseudolarix or Sciadopitys. The reduction
to this bundled short shoot never occurred
in Metasequoia glyptostroboides or in Taxo-
dium distichum although progenitors were
also already present in the Permian.
The evolution of the berry-seeded conifers
Some modern Cupressoideae, such as
Torreya, Cephalotaxus and Calocedrus,
are normally characterised by thick,
fleshy, berry-like fruits. For a long time
no satisfying answer could be given about
their development. Amazingly, these
conifers evolved at the same time as the
other conifers began to form and in the
same way as the first Araucarias (Ortiseia):
sterile micro-leaves began to coat the seed,
forming a fleshy aril. There is no other way
to explain the berry-like cone structure of
the conifer Ullmannia in the Upper Permian.
Early–Middle Triassic Kandutschia kuehnii
has the same blueprint. If the berry held
only one seed inside, which is often difficult
to determine, then these can be considered
as an ancestral line to the present-day
Torreya, Cephalotaxus and Taxus.
The development of the Cupressus-Voltziales
Some living conifers cannot be classified
in any of the evolutionary lines described
above, because they hold three or more
seeds on each scale. Today, they play only
a minor role in the conifer kingdom. At the
Carboniferous–Permian border, three- and
more-seeded conifers were widespread, and
often they were categorised as Voltziales.
Although this important group is often re-
garded as leading to all modern-day conifers,
particularly in the Mesozoic, it is not so.
As first certain Voltzia conifer can be
regarded Seymourina niederhauseni (Perner
& Wachtler, 2013), originating just between
the Kasimovian and Gzhelian (Regional stage
Western Europe: Stephan B-C). Female
cones were extraordinarily tall, sometimes
25 cm in length, while male cones reached
a length of about 10 cm. A fair number of
dwarfish, sterile tapered leaves surrounded
the scales. The seed scales were divided
into the three or four (and sometimes more,
Dolomythos, 2016
 3

2 5
Twigs and needles can be variegated in the same genus, but their fructifications build the connection link between
the species: 1. Araucaria cunninghamii; 2. Araucaria laubenfelsii; 3. Araucaria araucana; 4. Araucaria angustifolia; 5.
Araucaria heterophylla

then called Thuringiostrobus) fertile rounded the type-conifer Pseudovoltzia liebeana is

lobes and additionally were equipped on
the outside with a projecting sterile bract.
Ovules/seeds were collocated and hung
inverted from the upper side of each lobe.
From the Early Permian until the Late
Permian similar Voltzialean conifers, mostly
the three-lobed varieties, are found. In the
Alps, these were classified as Seymourina
viallii with its seed scales Dolomitia nonensi
(Kungurian) and Lopingian Dolomitia
cittertiae (Clement-Westerhof, 1984;
Wachtler, 2012). In the coeval Middle
European Upper Permian Zechstein basin,
encountered. Most Voltzia cones disintegrate
at maturity on the tree. In the Triassic, a
Voltzialean conifer explosion occurred in
Europe, with many arborescent to shrubby
subgenera and species that usually held
three lobes and were three-seeded (many
Voltzia species) but could be as much as
five-lobed and five-seeded (Aethophyllum,
Swedenborgia).
The vegetative branches were not uniform
in the same way as today’s Araucarias
or Cupressaceae. Some (Seymourina
niederhauseni, Seymourina viallii), held

Michael Wachtler: The evolution of gymnosperms in the Triassic

15
 slender, geometrically regular branchlets
with falcate needles, such as the extant
Norfolk pine (Araucaria heterophylla).
Others were characterised by their rope-
like protruding twigs, irregularly and
sparsely branched and armoured by leathery
triangular leaves (Voltzia unescoensis,
Voltzia rietscheli, Voltzia dolomitica), much
like today’s Araucaria araucana or Araucaria
bidwillii. Additional types were linguiform,
such as the enigmatic Pelourdea vogesiaca,
with foliage that was up to 15 cm long and
resembled the extant Agathis conifers.
Elongated, slender needles about 20 cm long
pertain to Aethophyllum stipulare but some
were also thorny (Agordia). In the Alpine
Voltziales, a consistent line can be followed
from the Early–Middle Triassic (Anisian)
Voltzia unescoensis, Voltzia agordica and
Voltzia rietscheli, all of which had rope-
like branchlets and hard (but not leathery)
leaves. The Ladinian Voltzia dolomitica
(Wachtler & Van Konijnenburg-Van Cittert,
2000) bore thick-walled armoured leaves
on massive shoots, as did the Late Triassic
(Carnian) Voltzia carinthica.
It may be that these widespread and
influential Voltziales disappeared completely
at the Triassic–Jurassic border, or that they
survived in some extant form. Some genera
in the fairly inhomogeneous family of the
Cupressaceae can otherwise be accepted as
possible Voltzia descendants. The usually
three- or more-seeded Cryptomeria,
Cunninghamia or Sequoiadendron, but
also the Juniper tree, can be regarded as
the most likely potential relatives of the
Voltziales.
The beginning of the Cupressaceae
The large and globally ranging group of the
Cupressaceae (or cypress) family unfortu-
nately represents an arbitrarily put together
system of conifers, such as Cunningham-
hioideae, Taiwanioideae, Athrotaxidoideae,
Sequoioideae, Taxodioideae, Callitroideae
and the super-group of Cupressoideae with
the genera Juniper and Cupressus.
Many of the forerunners of widespread co-
nifer families seem to be now already oc-
cupied. This is true of the Auraucariaceae
(occupied with Ortiseia as the most likely
potential ancestor), the Abietaceae (Ma-
jonica) and Piceoideae (Estellencsia), the
Pinoideae Pinus (with Early Permian Valen-
tinia), the Cephalotaxaceae and Taxus (Ul-
16
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lmannia, Kandutschia), and the Laricoideae
(Wachtlerolarix). Other genera from the
Cupressaceae remain, such as Taiwania or
Glyptostrobus. Early–Middle Triassic Alpia
anisica or the Ladinian Alpia ladinica, as
well as the Late Triassic Pusteria maribelae,
with its needle- or awl-like foliage and small
cones, have some similarities with them and
can be seen as descendants. The origin of
other Cupressaceae, which have foliage in
flat sprays, such as the Chamaecyparis or
Cupressus, or leaves arranged in scale-like
decussate pairs, like Thujopsis, were un-
known until now.
References
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