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The fern Sphenopteris through the Alpine Permian

Authors:
  • Museum Dolomythos

Abstract

The fern Sphenopteris constitutes one of the most common but also most misunderstood Per-mian ferns. During that time it was widespread across the Northern hemisphere. This short abstract will give brief information about changing and evolving tendencies of this fern in the Alpine Permian. Difficulties arose either because some vague terms were introduced in the last century confusing Lower with Upper Permian species or holotypes were lost-probably destroyed in the Second World War-and therefore it was necessary to introduce neotypes. This is sufficient for Sphenopteris suessi, belonging to the Early Artinskian, described from the Italian Collio-Formation by Hanns Bruno GEINITZ. For Sphenopteris battistii n. sp. (Artin-skian/Kungurian) from the Tregiovo-Formation it was useful to introduce a new species name due to distinctive differences from other Sphenopteris-ferns. April 2015.
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Dolomythos, 2015
The fern Sphenopteris through the Alpine Permian
by
Michael Wachtler
P. P. Rainerstrasse 11, 39038 Innichen, Italy; E-mail: michael@wachtler.com
Abstract
The fern Sphenopteris constitutes one of the most common but also most misunderstood Per-
mian ferns. During that time it was widespread across the Northern hemisphere. This short
abstract will give brief information about changing and evolving tendencies of this fern in the
Alpine Permian. Difficulties arose either because some vague terms were introduced in the
last century confusing Lower with Upper Permian species or holotypes were lost - probably
destroyed in the Second World War - and therefore it was necessary to introduce neotypes.
This is sufficient for Sphenopteris suessi, belonging to the Early Artinskian, described from
the Italian Collio-Formation by Hanns Bruno GEINITZ. For Sphenopteris battistii n. sp. (Artin-
skian/Kungurian) from the Tregiovo-Formation it was useful to introduce a new species name
due to distinctive differences from other Sphenopteris-ferns.
April 2015.
Key words: Early Permian floras, Pterophyta, Sphenopteris, Northern Italy
Division: Pteridophyta
Family: Osmundaceae (BERCHTOLD &
PRESL, 1820)
Genus: Sphenopteris (STERNBERG, 1825)
Sphenopteris suessi (GEINITZ,
1869)
1869 Sphenopteris suessi, Sphenopteris tridactylites,
Sphenopteris oxydata, GEINITZ, fig. 2-8, table V
1999 Sphenopteris suessi VISSHER ET. AL. fig. 6,Table
1, Sphenopteris kukukiana fig. 12, Table 1
Neotype
Coll. Ragazzoni N. 1 (Brescia, Museum of
Natural Sciences)
Paratype
COL 34 (Coll. Wachtler) Museum Brescia
Material
Coll. Ragazzoni N. 2, Coll. Ragazzoni N.
127, Coll. Allegretti, N. 76, Coll. Wachtler
COL 87, COL 114. All Brescia, Museum of
Natural Sciences)
Locality
Collio - Val Trompia, Lombardy, Italy
Type horizon and age
Early Permian (Sakmarian? - Artinskian)
Collio Formation
Diagnosis
Wedge-shaped and repeatedly incised fern-
fronds, leaflets bifid and with hidden vein-
structure. Fertile fronds skeletal, with the
sporangia densely aggregated on the single
pinnae.
Description
Fronds: Low growing fern, fronds alternate
and tripinnate. Neotype COL Brescia Museum
1 represents an incomplete semi-destroyed
frond from the old Ragazzoni collection with
partially fertile and sterile pinnae.
Pinnules: Secondary pinnae opposite to
alternate, bifid and deeply incised, apices
are complete and acute (COL 34). Pinnae
only allusively evidencing the mid-vein or
other internal veins, mirror-like pinnules
only on the basal leaves, upper part
irregularly incised. The attachment to the
rachis is narrow. They are up to 2.0 cm
long, 1 cm wide, often looking withered and
skeletonised.
Fertile pinnules: Much more skeletonised,
exhibiting only a rudimentary leaf structure
(COL 114). Single leaflets 1 cm long. The
sporangia clung densely to the lower side of
the pinnules.
Discussion
Beginning from the Carboniferous/Permian
border, most of the hygrophilous ferns dis-
59
Michael Wachtler and Thomas Perner: Permian Fossil plants from Europe
The original-plate of Hanns Bruno GEINITZ, (1869, Tav. 5). Über Fossile Pflanzenreste aus der Dyas von Val Trompia
featuring Sphenopteris ferns from the Early Permian Collio Formation. 2. Sphenopteris tridactylites, 3-7. Sphenopt-
eris suessi, 8. Sphenopteris oxydata. Courtesy Senckenberg Naturhistorische Sammlungen Dresden.
appeared so that only one genus of true
ferns became widespread in the Permi-
an: Sphenopteris. This pteridophyta is till
now the only certain representative of the
ferns, although - strangely enough - other
plants like conifers, cycads, but also horse-
tails were diffuse and variegated (PERNER &
WACHTLER, 2015).
In 1869, Hanns Bruno GEINITZ described
several floral elements from the Collio-For-
mation in the Lombardian Prealps passed
to him by the Austrian geologist Eduard
SUESS. They had spent several days in the
Val Trompia together with the indefatiga-
ble local botanist Giuseppe RAGAZZONI. In
1868 they had made a field trip to the Mon-
te Colombine (SUESS, 1869). SUESS ex-
pressed his great gratitude to Don Giovan-
ni BRUNI and his personal mountain-guide
RONCHINI as well as other collectors of the
fossil specimens (CASSINIS & SANTI, 2001).
Although all the SUESS specimens sent to
GEINITZ and stored in the Natural History
Museum of Dresden were lost in the
troubles of the Second World War, today we
nevertheless have a good overview of the
collected fossils due to the specimen from
the RAGAZZONI and Don BRUNI collection in
the Natural History Museum of Brescia.
Hanns Bruno GEINITZ published the fossil
slabs handed him on only one plate. Apart
from some doubtful Sphenopteris tridac-
tylites (BRONGNIART) and Sphenopteris
oxydata (GOEPPERT) classifications - previ-
ously known from the European Rotliegend
- all figures (from 3 to 7, Table V) feature
Sphenopteris suessi, a new species name
honouring Eduard SUESS’ activities in the
Val Trompia. Albeit the old plant slabs were
impregnated with varnish, probably by the
original collectors, and therefore sometimes
darkened, in the Brescia Natural-Museum
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Dolomythos, 2015
1) COL. Ragazzoni Brescia N. 1. Sphenopteris suessi. Designated neotype found by Giuseppe Ragazzoni in 1867. Fer-
tile and sterile fronds
2) COL. Allegretti Brescia N. 127. Sphenopteris suessi. Strange counterplate of N. 1, but found by Corrado Allegretti
(1894-1969), probably around 1950
3) COL. Allegretti Brescia N. 127. Sphenopteris suessi. Label on the specimen
4) COL. Ragazzoni Brescia N. 2. Sphenopteris suessi. Fertile and sterile pinnules
5) COL. Brescia N. 76. Sphenopteris suessi. Frond
1
2
3
45
1cm
2cm
1cm
2cm
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Michael Wachtler and Thomas Perner: Permian Fossil plants from Europe
6) COL 34 (Coll. Wachtler) Sphenopteris suessi. Paratype. Detail of frond
7-8) COL 34 (Coll. Wachtler) Sphenopteris suessi. Isolated pinnules
9) COL 87 (Coll. Wachtler) Sphenopteris suessi. Detail of a sterile frond
10) COL 114 (Coll. Wachtler) Sphenopteris suessi. Detail of a sterile frond. All Monte Dasdana, Wachtler Collection
69
7 8 10
5mm
5mm
5mm
5mm
5mm
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Dolomythos, 2015
we found sufficient opportunities to identify
the story of these ferns.
In the following decades, especially
in the 20th century, additional names
for Sphenopteris ferns from the Collio-
Formation, such as Sphenopteris kukukiana,
S. dichotoma, S. hoeninghausi or S. patens
were given, mostly because of distant
resemblances to species from German
Upper Permian localities. In the end this
culminated in the absurd situation - also
including the material in the Natural History
Museum in Brescia - that for all suggested
Sphenopteris ferns an equal number of
Sphenopteris species names were assigned.
By collecting plant slabs on Monte Dasdana
as well studying material in the museums it
became obvious that we were not dealing
with so many species. Therefore - because
the original material seemed definitely to
have been destroyed in the Second World
War - a neotype was selected from the old
RAGAZZONI-material stored in the Brescia
Museum with the suggestive number 1 and
collected in 1867; however, it was darkened
by lacquer. Amazingly, in examining the rest
of the collections, another specimen, N.
127, was noted identified on its backside by
a label that it was collected by ALLEGRETTI
but was the perfect counterplate to neotype
N. 1.
It can therefore be suggested, due to the
relatively fresh character of the slab, that
Corrado ALLEGRETTI - a famous Italian spe-
leologist - found this counterplate about
hundred years later in the heap of material
left by RAGAZZONI on Monte Colombine.
It can be stated that Sphenopteris suessi
was one of the dominant low-growing plants
of the Early Permian Collio-Formation,
whereas all the other Sphenopteris species
names have to be excluded because they
are synonymous, being taken from the Ger-
man Lower Permian Rotliegend or from more
younger formations like the Upper Permian
Zechstein.
It can even be asserted that the Sphenop-
teris species from the slightly younger Tre-
giovo-Formation have so many differences
in frond and leave organisation that we have
to choose a new name: Sphenopteris battis-
tii. It therefore constitutes an intermediate
fern element between the Lower Permian
Sphenopteris suessi and the Upper Permian
Sphenopteris dichotoma.
Sphenopteris battistii sp. n.
(WACHTLER, 2015)
1999 Sphenopteris suessi VISSHER ET. AL. fig. 6,Table
1, Sphenopteris kukukiana fig. 12, Table 1
2012 Sphenopteris dichotoma WACHTLER, p. 35
2013 Sphenopteris suessi WACHTLER p. 127, p 129
Holotype
TRE 541
Paratype
TRE 448 (fertile frond)
Material
TRE 37, 130, 166, 201, 301, 329
Repository
MUSE, Museum of Natural Science, Trento,
Coll Valentini, Coll. Wachtler
Etymology
Named after the Italian patriot and politician
of Austrian citizenship Cesare Battisti
(1875–1916), who became a prominent
Irrendentist and was sentenced to death.
Type locality
Tregiovo-Le Fraine, Trentino, Italy
Type horizon and age
Early Permian (Artinskian-Kungurian)
Tregiovo-Formation
Diagnosis
Tripinnate fronds, single pinnules alternate
and with narrow attachment. Distal pinnules
forming a slightly bifid tip. Fertile pinnules
skeletonised, with densely arranged sporan-
gia on the lower side of the leaflets.
Description
Fronds: Low growing fern with tripinnate
fronds. Holotype TRE 541 evidences an
overlapping 9 cm long frond, dividing on the
apical part into two equal branchlets. This
is a characteristic of almost all Sphenopteris
fronds, as seen in TRE 301 and 329.
Pinnules: Secondary pinnae opposite to
alternate, only slightly bifid and incised
(Holotype TRE 541). Single pinnules
are up to 2 cm long and 0.5 cm wide,
characteristically segmented into 3 mirror-
like leaves with a concluding only slightly
incised pinnula.
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Michael Wachtler and Thomas Perner: Permian Fossil plants from Europe
Fertile pinnules: Skeletonised, sporangia
densely covering the lower part, mainly
encasing the whole leave. Paratypes TRE
448, TRE 201 and 329 represent fertile
specimens. The leaflets are from 1 to 2 cm
long and 0.5 cm wide.
Discussion
Among the Collio-Flora, Sphenopteris
ferns were observed by the old authors
(GEINITZ, 1869) and described with
partially derivative names from different
epochs like Sphenopteris kukukiana S.
patens, S.tridactylites S. oxydata. Due
to the consistent differences from other
Sphenopteris ferns from Germany and
France, only Sphenopteris suessi can
therefore be regarded as corresponding to
the endemic terrain. The same is valid for
the slightly younger Tregiovo-beds located
nearby. Due to the poor preservation
observed for most species recovered there
by older researchers, given names like
Sphenopteris dichotoma (WACHTLER, 2012),
Sphenopteris kukukiana (VISSHER ET. AL.,
1999) or Sphenopteris suessi (VISSHER ET.
AL., 1999, WACHTLER, 2013) have to be
regarded with caution, because they may
have been taken from other localities in
different time periods. Profound studies and
researches established that the Sphenopteris
fern from Tregiovo is so different from the
Collio fern Sphenopteris suessi as to require
a new species name, Sphenopteris battistii.
Whereas in the Collio-Formation seven
striking disconnected leaves rounded on
the apex can be counted in the single last
pinnula, in Sphenopteris battistii we can
recognise in the single leaf an only slightly
bifid to sometimes dentate form.
The Upper Permian Sphenopteris dichotoma,
known mainly from German Zechstein,
with some doubtful specimens recovered
from the coeval Gröden-Formation from
Bletterbach, Recoaro (ULBE 89) and the
Valli del Pasubio (Ariche PAS 201), hold,
in contrast, smaller leaflets. Sphenopteris
leaves, especially in the Collio- but also in
the Tregiovo-Formation, can sometimes be
confused with Valentinia conifers, but they
hold longer and needle-like leaves that are
never incised.
Sphenopteris constitutes one of the most
dominant flora elements and sometimes
the only certain fern of the European Middle
to Late Permian. From the Upper Permian
German Zechstein, many Sphenopteris-
species have been described (SCHWEITZER,
1960) (S. dichotoma, S. geinitzii, S.
bipinnata, S. gillitzeri), often based only
on the existence of a single specimen,
disregarding the fact that we are dealing
with a fern with extremely changeable
leaflets (HAUBOLD & SCHAUMBERG, 1985).
Several findings from the Tregiovo-
Formation have established that the fern
Detail of a. Sphenopteris suessi from the slightly older Collio-Formation and b. Sphenopteris battistii from the Tregio-
vo-Formation. The incisions are more pronounced and skeletonised.
ab
64
Dolomythos, 2015
1) TRE 541. Sphenopteris battistii. Designed holotype. Part of a frond
2) TRE 541. Sphenopteris battistii. Designed holotype. Detail of the pinnules
3) TRE 541. Sphenopteris battistii. Designed holotype. Detail of a pinnula
4) TRE 448. Sphenopteris battistii. Paratype. Detail of a fertile frond
1
2
34
2cm
5mm
1cm
1cm
65
Michael Wachtler and Thomas Perner: Permian Fossil plants from Europe
Sphenopteris battistii n. sp. Reconstruction. a. Part of a frond {TRE 541), b. Single pinnula, c. Fertile frond (TRE
329), d. Part of a fertile frond, e. Entire plant.
a
b
c
d
e
66
Dolomythos, 2015
5
1 cm
4
1 cm
5) TRE 301. Sphenopteris battistii. Sterile frond
6) TRE 329. Sphenopteris battistii. Fertile frond
7) TRE 201 Sphenopteris battistii. Fertile frond
8) TRE 37B. Sphenopteris battistii. Part of a fertile frond
9) TRE 201. Sphenopteris battistii. Detail of the sporangia
91 cm
7
6
8
1 cm
1 cm
67
Michael Wachtler and Thomas Perner: Permian Fossil plants from Europe
Sphenopteris constitutes the most frequent
fern there. It could be stated that the
desiccated looking Sphenopteris is not
only the most common fern in the Permian
Europe but also forms the characteristic
fern of the Alpine Permian, being present
from the Lower to Upper Permian. Often
it represents the only fern traced, a
phenomenon that is hard to believe when
we think about the extraordinary richness
of ferns beginning from the Early Triassic
(Anomopteris, Ladinopteris, Gordonopteris,
Anotopteris, Wachtleria etc.) and also the
Carboniferous. Why the ferns declined so
much in the Permian after a peak in the
Carboniferous and further arose Triassic
must be studied, because other water-
loving plants like cycads and horsetails
were not affected during this decline. Their
insertion in extant fern-families is difficult,
but with caution they can be regarded as
Osmundaceae.
References
CASSINIS, G.,SANTI G., 2001: Hanns Bruno Geinitz:
A pioneer of the Permian stratigraphy of Eastern Lom-
bardy (Southern Alps, Italy). In: “Intern. Hanns Bruno
Geinitz Symp.”, Proceed. Symp. in Honour of Hanns
Bruno Geinitz, Dresden, Saxony, January 28-30, 2000,
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HAUBOLD, H., SCHAUMBERG, G. 1985. Die Fossilien
des Kupferschiefers. Pflanzen- und Tierwelt zu Beginn
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904127
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del Pasubio. Coll. Wachtler
ULBE 89. Sphenopteris sp. Upper Permian Ulbe Re-
coaro. Coll. Wachtler
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SUESS E. 1869. Über das Rothliegende im Val Trompia. 107-119, Sitzungsberichte vom 21. Jänner 1869, Wien SCHWEITZER, H.-J., 1960. Die Sphenopteriden des Zechsteins. Senckenbergiana Lethaea, 41, pp. 37-57, Frankfurt.
Hanns Bruno Geinitz: A pioneer of the Permian stratigraphy of Eastern Lombardy (Southern Alps, Italy)
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The latest Artinskian-Kungurian (Early Permian) Flora from Tregiovo -Le Fraine in the Val di Non (Trentino -Northern Italy) -Preliminary researches
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WACHTLER, M. 2012. The latest Artinskian-Kungurian (Early Permian) Flora from Tregiovo -Le Fraine in the Val di Non (Trentino -Northern Italy) -Preliminary researches, Dolomythos, 3-56 Innichen. ISBN 978-88-904127
In Perner & Wachtler: Permian fossil plants in Europe and their evolution, Dolomythos and Oregon Institute of Geological Research, Portland 1 cm PAS 201. Sphenopteris sp. Upper Permian Ariche, Valli del Pasubio. Coll. Wachtler ULBE 89
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WACHTLER M. 2013. The latest Artinskian/Kungurian (Early Permian) Flora from Tregiovo-Le Fraine in the Val di Non (Trentino, Northern Italy) -Additional and revised edition. In Perner & Wachtler: Permian fossil plants in Europe and their evolution, Dolomythos and Oregon Institute of Geological Research, Portland 1 cm PAS 201. Sphenopteris sp. Upper Permian Ariche, Valli del Pasubio. Coll. Wachtler ULBE 89. Sphenopteris sp. Upper Permian Ulbe Recoaro. Coll. Wachtler