Occurrence of fossil bamboo pollen and a fungal conidium of Tetraploa cf.

aristata in
Upper Miocene deposits of Józefina (Poland)
Elżbieta Worobiec
, Grzegorz Worobiec
⁎, Przemysław Gedl
Władysław Szafer Institute of Botany, Polish Academy of Sciences, Lubicz 46, PL-31-512 Kraków, Poland
Institute of Geological Sciences, Cracow Research Centre, Polish Academy of Sciences, Senacka 1, PL-31-002, Kraków, Poland
a b s t r a c t a r t i c l e i n f o
Article history:
Received 26 November 2008
Received in revised form 24 April 2009
Accepted 5 May 2009
Available online 7 May 2009
pollen grains
fossil bamboos
fossil fungi
wetland vegetation
Upper Miocene
The paper presents results of pollen analysis of the Upper Miocene deposits from a borehole at Józefina
(Kraków-Silesia Upland, central Poland), including data concerning bamboo type pollen grains, as well other
sporomorphs (pollen, spores and freshwater phytoplankton). Fossil pollen grains of bamboos (Graminidites
bambusoides Stuchlik) were found in two samples, and in one sample they were accompanied by conidium of
Tetraploa-like fossil fungus, resembling the recent Tetraploa aristata Berkeley & Broome. This fungus could
grow on leaves of the bamboos or other grasses, but also on variety of other plants. Comparison of
palynological data, studies of leaf floras and the nearest living relatives, confirmed the connection of the fossil
bamboos with wetland, reed and riparian vegetation. Results of these analyses indicate a moderately wet and
warm temperate palaeoclimate.
© 2009 Elsevier B.V. All rights reserved.
1. Introduction
Pollen grains of bamboos are rarely reported from the Neogene
palynofloras of Europe. Till now in Poland only two localities were
known for the fossil bamboo taxon Graminidites bambusoides Stuchlik
(Stuchlik, 1964; Worobiec, 2000; Worobiec and Worobiec, 2005). A
borehole through Upper Miocene sediments of the Kraków-Silesia
Upland in central Poland provides a very good state of preservation of
pollen grains, spores, phytoplankton and fungi, as well as high
taxonomical diversity of sporomorphs in samples with G. bambusoides,
enabling detailed examination of the palynoflora accompanying the
bamboo type pollen grains. Additionally, together with the bamboo
pollen, a fossil fungal conidium, similar to the recent Tetraploa, was
found. Fungi of this genus were extremely rarely reported from
European Tertiary. Tetraploa may have grown on leaves of bamboos or
on other grasses.
2. Materials and methods
The material was collected from a borehole drilled at Józefina
(northern edge of the Kraków-Silesia Upland, central Poland; Fig. 1).
During the Miocene this area was situated at the Meta-Carpathian
Swell, which separated the marine basin of the Carpathian Foredeep
in the south fromthe lowland areas of central Poland (in the north). In
the latter area, continental and brackish sedimentation took place,
leaving brown coal seams (Piwocki et al., 2004).
The Miocene succession at Józefina starts at ca. 57 m depth, with
conglomerates composed of limestone clasts derived fromthe Jurassic
substratum. Above these conglomerates, at 53.5–51.0 m depth, pale-
coloured sandy–loamy complex occurs. Higher, at 51–45 m depth,
dark-coloured (dark-greenish and black) non-calcareous clays occur.
The uppermost part of the Miocene succession (depth 45–25 m)
consists of monotonous pale-coloured (pale-grey–greenish to willow-
green) non-calcareous clays. In our study ten samples from a depth of
25.2–56.4 m were examined.
The samples were prepared using hydrochloric acid (HCl), hydro-
fluoric acid (HF), heavy liquid (ZnCl
+HCl; density 2.0 g/cm
separation, ultrasound for 10–15 s, and sieving with a 15 µm nylon
mesh. The microscope slides were made using glycerine jelly as a
mounting medium. The quantity of rock processed varied from 50 to
100 g. The rock samples, palynological residues and slides are stored in
the collection of the Institute of Geological Sciences, Cracow Research
Centre, Polish Academy of Sciences, Kraków.
The taxa have been classified to an appropriate palaeofloristical
element, mainly onthe basis of the checklist of selectedpollenandspore
taxa from the Neogene deposits (Ziembińska-Tworzydło et al., 1994a).
Review of Palaeobotany and Palynology 157 (2009) 211–217
⁎ Corresponding author.
E-mail addresses: e.worobiec@botany.pl (E. Worobiec), g.worobiec@botany.pl
(G. Worobiec), ndgedl@cyf-kr.edu.pl (P. Gedl).
0034-6667/$ – see front matter © 2009 Elsevier B.V. All rights reserved.
Contents lists available at ScienceDirect
Review of Palaeobotany and Palynology
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Microphotographs were taken using a NIKON Eclipse E400 microscope
fitted with Canon A640 digital camera.
3. Results of pollen analysis
Ten samples from Józefina were examined, but only three of them
(taken from dark-greenish and black non-calcareous clays, depth
45.7–47.5 m) yielded well-preserved sporomorphs suitable for
detailed pollen analysis. About 600–800 pollen grains and spores, as
well about 60–260 specimens of freshwater phytoplankton were
encountered in each of these samples. The studied spectra were
morphologically differentiated, and about 40–50 taxa of pollen, spores
and phytoplankton were identified in each sample. An abridged list of
the taxa, and their botanical affinity are given in Table 1.
Results of pollen analysis show a significant role of wetland and
riparian vegetation. Occurrence of Nymphaeaceae pollen, Salvinia-
ceae/Azollaceae microremains, and abundant freshwater phytoplank-
ton: Sigmopollis as well as zygospores of Zygnemataceae (Ovoidites,
Tetraporina and Stigmozygodites), desmids (Closteritetrapidites) and
others, point at the presence of a water basin at the time of sedi-
mentation. This freshwater basin (meandering river, ox-bow lake or
small lake) was surrounded by swamp-aquatic vegetation, composed
of herbs (including grasses, Chenopodiaceae and Asteraceae), and
riparian forests dominated by Pterocarya, Carya and Liquidambar,
accompanied by Ulmus, Juglans and others. Drier terrains were
vegetated by mixed mesophytic forests dominated by Pinus, Fagus
and Carpinus, accompanied by Tsuga, Sciadopitys, Abies, Picea, and
Results of pollen analysis indicate that the climate was warm
temperate and moderately wet. The composition of pollen spectra and
mutual ratio of palaeotropical and arctotertiary elements point at a
Late Miocene age of the deposit. The studied assemblage shows
similarity with the XII climatic phase belonging to the Carpinipites–
Juglandaceae spore-pollen zone, deposited in the Pontian (spore-
pollen zones according to Piwocki and Ziembińska-Tworzydło, 1997).
4. Fossil pollen grains of bamboos from Józefina
Graminidites Cookson 1947 ex Potonié 1960
Graminidites bambusoides Stuchlik in Ziembińska-Tworzydło et al.,
1994b (Pl. I, 1–3)
1964 Gramineae “Bambusa” type; Stuchlik, p. 76, pl. 24, Figs. 1–2.
1970 Graminidites sp. A (Bambusa-typus); Krutzsch, p. 51, pl. 1,
Fig. 1.
1994b Graminidites bambusoides Stuchlik sp. nov.; Ziembińska-
Tworzydło et al., p. 14, pl. 8, Figs. 4–5.
2005 Graminidites bambusoides Stuchlik; Worobiec and Worobiec,
pp. 42, 44, pl. 3, Figs. 1–3.
2007 Poaceae (Bambusoideae); Ivanov et al., pl. 2, Figs. 26–27.
Description: Pollen grains monoporate, oval to circular, 50–60 µm
in diameter. Pore 4–5 µm in diameter, with distinct annulus 4.0–
4.5 µm broad. Exine composed of two layers, very thin (about 1.0 µm
thick), with secondary folds, surface very finely granulate. Pollen
grains often deformed.
Remarks: Pollen grains of the fossil taxon Graminidites bambu-
soides Stuchlik were found in two samples from Józefina (depth
47.5 m and 46.2 m), and were accompanied by other (smaller) pollen
grains of grasses (Plate I, 4). In one sample 28 pollen grains of G.
bambusoides (about 4.5% of spore-pollen sum) were encountered.
Occurrence of attached pollen grains (Plate I, 1) points at in situ
presence of this pollen.
Graminidites bambusoides Stuchlik differs from the nearest G.
pseudogramineus Krutzsch by greater dimensions, considerable
broader annulus and thinner exine.
Fossil pollen grains of bamboo type morphology were described
from the Miocene deposit from Rypin (central Poland) as Gramineae
“Bambusa” type (Stuchlik, 1964), and later named Graminidites
bambusoides Stuchlik (Ziembińska-Tworzydło et al., 1994b). Accord-
ing to Stuchlik (1964) they resemble pollen of the recent genera
Bambusa Schreber, Oryza L. and Dendrocalamus C.G. Nees. Pollen
grains of G. bambusoides were also found in the Middle Miocene
deposit from Legnica in southwestern Poland (Worobiec, 2000;
Worobiec and Worobiec, 2005). A comparison of this fossil taxon
with pollen of recent cereals and bamboos shows that G. bambusoides
is morphologically close to the bamboos, e.g. representatives of the
genus Arundinaria Michx.
5. Fossil conidium of Tetraploa from Józefina
Tetraploa Berkeley & Broome 1850
Tetraploa cf. aristata Berkeley & Broome 1850 (Pl. II, 1a–1d)
1850 Tetraploa aristata Berkeley & Broome; p. 459.
1971 Tetraploa aristata Berkeley & Broome; Ellis, pp. 51–52, Fig. 22.
1978 Type 89: Tetraploa aristata Berkeley & Broome; Van Geel,
p. 96, pl. 17, Fig. 89.
Description: Conidiumbrown, withbodyand4 appendages. Body of
conidium32 μmlong and 20 μmwide, coarsely granulate to verrucate;
ornamentationis more densely towards basal part, base rounded. Body
composed of most probably four columns (but only up to two columns
are visible on Plate II), 8.5–10.0 μm wide. The number of cells in a
Fig. 1. Simplified map of the Upper Miocene deposits in Poland (after Piwocki, 1998;
changed), and location of the Józefina borehole.
212 E. Worobiec et al. / Review of Palaeobotany and Palynology 157 (2009) 211–217
column is 3 or 4, cells are 7.5–12.5 μmlong. Each column gives rise to a
septate setose appendage of approximately equal length (47–50 μm),
at the base slightly verrucose and smooth in the upper part, ca. 5 μm
thick at the base and 4 μm thick at the apex with 3 septa preserved.
Remarks: The above-described fungal microfossil was found in a
sample with numerous Graminidites bambusoides pollen grains (depth
47.5 m, Table 1). Morphologically it exactly corresponds to conidia of the
extant Tetraploa Berkeley & Broome. However, a similar microfossil has
been also described as an acritarch under the genus name Frasnacrite-
trus Taugourdeau from Devonian (Frasnian) deposits of France
(Taugourdeau, 1968). This author mentioned that erection of the new
genus was done on the basis of one specimen and he cannot eliminate
the possibilityof contamination by recent material. Taugourdeau(1968)
consideredthe possibilities of contaminationandfinally he rejectedthat
notion (Kalgutkar and Jansonius, 2000). Subsequently the genus name
Frasnacritetrus has been used for Tetraploa-like microfossils (Saxena
and Sarkar, 1986; Sarkar and Singh, 1988; Kumar and Takahashi, 1991;
Kumaran et al., 2001; Pedrão Ferreira et al., 2005; Antoine et al., 2006;
Kumaran et al., 2008) but more frequently the name Tetraploa was used
Table 1
Results of palynological analysis (number of palynomorphs) of samples from borehole
Józefina. S—spores, G—Gymnospermae, An—Angiospermae, V—varia, P—phytoplankton,
Taxon Botanical affinity Depth
45.7 m 46.2 m 47.5 m
S Baculatisporites sp. sp. Osmundaceae 1 11 2
S Corrusporis sp. Bryales 1 1
S Echinatisporis sp. Selaginella 1 2
S Foveotriletes sp. sp. Unknown
botanical affinity
2 1
S Laevigatosporites sp. sp. Polypodiaceae/
9 4 10
S Leiotriletes maxoides/maximus
2 1 1
S Leiotriletes sp. Sp. Schizaeaceae/
7 3 2
S Retitriletes frankfurtensis Krutzsch Lycopodium 1 1
S Rudolphisporis sp. Anthocerotaceae 1
S Rugulatisporites quintus Pflug &
Osmunda 1
S Stereisporites sp. Sphagnum 1
S Verrucatosporites sp. sp. Dennstaedtiaceae 2 2
G Abiespollenites sp. sp. Abies 31 28 12
G Cathayapollis sp. sp.
+Pinuspollenites sp. (Pinus
haploxylon type)
Pinaceae, Cathaya,
94 102 76
G Cedripites sp. sp. Cedrus 1 6
G Inaperturopollenites sp. Taxodiaceae/
G Keteleeriapollenites sp. Keteleeria 4 3 4
G Piceapollis sp. sp. Picea 24 18 9
G Pinuspollenites labdacus (Potonié)
Raatz ex Potonié
Pinaceae, Pinus 316 300 212
G Sciadopityspollenites sp. sp. Sciadopitys 28 12 28
G Sequoiapollenites sp. Sequoia/
G Zonalapollenites sp. sp. Tsuga 58 31 40
An Aceripollenites sp. Acer 1
An Alnipollenites verus Potonié ex
Alnus 1 2 2
An Araliaceoipollenites edmundi
(Potonié) Potonié ex Potonié
Araliaceae 1 1
An Arecipites pseudoconvexus
Arecaceae 1
An Carpinipites carpinoides (Pflug)
Carpinus 15 5 28
An Caryapollenites simplex (Potonié)
Raatz ex Potonié
Carya 38 7 15
An Celtipollenites sp. Celtis 3 4
An Cercidiphyllites minimireticulatus
(Trevisan) Ziembińska-Tworzydło
Cercidiphyllum 1
An Chenopodipollis multiplex
(Weyland & Pflug) Krutzsch
Chenopodiaceae 1 7
An Cichoreacidites gracilis (Nagy)
An Corsinipollenites ludwigioides
Oenotheraceae 1
An Cyperaceaepollis neogenicus
Cyperaceae 2
An Diervillapollenites sp. Diervilla/Weigela 1
An Ericipites callidus (Potonié)
Calluna 1 1
An Ericipites ericius (Potonié) Potonié Erica 3 1 1
An Ericipites roboreus (Potonié)
Arbutus unedo
1 2 2
An Faguspollenites verus Raatz ex
Fagus 136 21 47
An Graminidites bambusoides Stuchlik Poaceae,
1 28
An Graminidites sp. Poaceae, Pooideae 4 3 11
An Intratriporopollenites
cordataeformis (Wolff) Mai
Tilia 1 1 4
An Intratriporopollenites instructus
(Potonié) Thomson & Pflug
Tilioideae 1
An Juglanspollenites sp. Juglans 3 5
Table 1 (continued)
Taxon Botanical affinity Depth
45.7 m 46.2 m 47.5 m
An Lonicerapollis gallwitzi Krutzsch Caprifoliaceae,
6 2
An Nympheapollenites sp. Nympheaceae 5 2
An Nyssapollenites sp. Nyssa 4
An Periporopollenites stigmosus
(Potonié) Thomson & Pflug
Liquidambar 30 3 5
An Persicarioipollis sp. Polygonum 1
An Polyatriopollenites stellatus
(Potonié) Pflug
Pterocarya 31 17 28
An Quercoidites sp. Quercus 4 1
An Ranunculacidites sp. Ranunculaceae 2
An Salixipollenites sp. Salix 1
An Tricolporopollenites
pseudocingulum (Potonié)
Thomson & Pflug
An Tricolporopollenites staresedloensis
Krutzsch & Pacltová
Hamamelidaceae 1
An Trivestibulopollenites betuloides
Betula 4
An Tubulifloridites granulosus Nagy Asteraceae,
An Ulmipollenites undulosus Wolff Ulmus 2 1 8
An Vaclavipollis pacltovae Krutzsch Caryophyllaceae 1
An Vitispollenites sp. Vitaceae 1
V Varia (pollen and spores indet.) ? 3 6
V Reworked palynomorphs ? 3 1
P Botryococcus sp. Botryococcus sp. 2
P Closteritetrapidites sp. Desmidiales,
1 2
P Diagonalites diagonalis Krutzsch &
1 5 3
P Dinoflagellate cysts Dinoflagellate 3 7
P Ovoidites sp. +Cycloovoidites
cyclus Krutzsch & Pacltová
3 17 4
P Pseudoschizaea rubina (Rossignol)
P Sigmopollis sp. ?Cyanophyta 75 23 225
P Spintetrapidites sp. ?Desmidiales,
2 3
P Stigmozygodites sp. Zygnemataceae,
1 2 4
P Tetraporina sp. Zygnemataceae,
1 4 7
P Zygodites sp. ?Desmidiales,
5 9 4
F Microthyriaceae indiff. Microthyriaceae 2
F Tetraploa cf. aristata Berkeley &
Tetraploa aristata
Berkeley &
Sum 960 664 898
Question marks in the table signify uncertain botanical affinity.
213 E. Worobiec et al. / Review of Palaeobotany and Palynology 157 (2009) 211–217
(Gupta, 1970; Van Geel and Van der Hammen, 1973; Sharma, 1976; Van
Geel, 1978; Kumar, 1990; Head, 1993; Hoorn, 1994; Davis et al., 2002;
Anzótegui and Garralla, 2004; Mudie et al., 2004; Marinova and
Atanassova, 2006; Singh and Chauhan, 2006; Connor et al., 2007;
Gómez et al., 2007; Bera et al., 2008; González et al., 2008). Saxena and
Sarkar (1986) emended the diagnosis of Frasnacritetrus allowing
inclusion of Tetraploa-like microfossils in this genus. According to
Head(1993) the microfossil describedas Frasnacritetrus by Taugourdeau
is almost certainly a modern contaminant of Tetraploa. Thus Frasnacri-
tetrus should be considered as junior synonym of Tetraploa and should
not be used as a form genus for Tetraploa-like fossils (Head, 1993).
Kalgutkar and Jansonius (2000) are of the opinion that microfossils
described as Tetraploa may have to be reassigned to the genus Frasna-
critetrus, but if it can be proven that the type of Frasnacritetrus is a recent
contaminant, it should become a junior synonym of Tetraploa.
Additionally, Jansonius and Hills (1987) noted that it was unlikely that
a close relative of Tetraploa already existed during the Devonian. The
present authors support the opinion that the microfossil which was the
basis for the erection of the genus Frasnacritetrus represents a
contamination by recent Tetraploa as there is a huge gap between the
first Devonian record of Frasnacritetrus and the next continuous record
of Cenozoic Tetraploa-like fossils beginning during Palaeocene (as Fras-
nacritetrus, Pedrão Ferreira et al., 2005). A gap in the fossil record of
almost 300 Ma is very unlikely.
In respect of shape and size the Tetraploa-like conidium from
Józefina is similar to Frasnacritetrus sp. 3 from the Tertiary of Himahal
Pradesh, India (Saxena and Sarkar, 1986) and especially to microfossil
type 89: Tetraploa aristata Berkeley & Broome from Holocene peat
deposits in the Netherlands and Germany (Van Geel, 1978).
Considering that extant conidia of Tetraploa aristata show strong
morphological variability (see description of T. aristata and Fig. 22 in
Ellis, 1971, also Tseng and Chen, 1982, and Becerra Hernández et al.,
2007), Tetraploa from Józefina should be regarded as belonging to T.
aristata. However, due to sparseness of material (one specimen) and
consequent lack of data on variability, it was named Tetraploa cf.
Plate I.
1. Two attached pollen grains (arrows) of Graminidites bambusoides Stuchlik, borehole Józefina, 47.5 m.
2–3. Graminidites bambusoides Stuchlik, pollen, borehole Józefina, 47.5 m.
4. Graminidites sp., pollen, borehole Józefina, 47.5 m.
214 E. Worobiec et al. / Review of Palaeobotany and Palynology 157 (2009) 211–217
Conidia of Tetraploa differ from the rather similar fossil stauros-
porae fungus Eoglobella longipes Bradley because of their muriform
multicellular body (Kalgutkar and Jansonius, 2000).
The reliable fossil record of Tetraploa (also as Frasnacritetrus) starts
fromthe Palaeocene (Pedrão Ferreira et al., 2005). Beginning fromthe
Miocene it is frequently recorded, mainly outside of Europe, e.g. in
India and South America (Gupta, 1970; Saxena and Sarkar, 1986;
Sarkar and Singh, 1988; Kumar and Takahashi, 1991; Hoorn, 1994;
Kumaran et al., 2001; Pedrão Ferreira et al., 2005; Antoine et al., 2006;
Kumaran et al., 2008). For the European Neogene Tetraploa is known
from the uppermost Pliocene St. Erth Beds locality in Cornwall,
England (Head, 1993). From Quaternary deposits (mainly Holocene)
Tetraploa is reported mostly outside Europe (Van Geel and Van der
Hammen, 1973; Sharma, 1976; Kumar, 1990; Davis et al., 2002;
Anzótegui and Garralla, 2004; Singh and Chauhan, 2006; Gómez et al.,
2007; Bera et al., 2008; González et al., 2008). In the European
Quaternary Tetraploa was found in the Holocene deposits of areas
with warm and mild climate, e.g. the Netherlands and Germany (Van
Geel, 1978), Bulgaria (Marinova and Atanassova, 2006) and Marmara
Sea in the Mediterranean area (Mudie et al., 2004). Close to Europe it
was reported from the Holocene deposits of the Colchis Lowland of
Western Georgia, Transcaucasia (Connor et al., 2007).
6. Discussion
6.1. Taxonomy, geographical distribution and ecology of extant
representatives of Tetraploa
Tetraploa is the anamorphic (asexual) stage of pleomorphic fungus
classified as a member of the dematiaceous staurospore hyphomy-
cetes (Ellis, 1971). A teleomorph (sexual stage) of Tetraploa aristata
was described by Scheuer (1991) as Massarina tetraploa Scheuer
Plate II.
1a. General view of conidium of Tetraploa cf. aristata Berkeley & Broome, borehole Józefina, 47.5 m.
1b–d. Detail of external and internal structure of conidium of Tetraploa cf. aristata Berkeley & Broome, borehole Józefina, 47.5 m.
215 E. Worobiec et al. / Review of Palaeobotany and Palynology 157 (2009) 211–217
(Ascomycetes, Pleosporales). The current name for the holomorph
(species in all its morphs: T. aristata and M. tetraploa) is Lophiostoma
tetraploa (Scheuer) Aptroot & K.D. Hyde (Hyde et al., 2002; Index
Fungorum, 2008).
Till now there have been about 11 described extant species of the
genus Tetraploa (Index Fungorum, 2008), Tetraploa aristata is most
frequently found. Today T. aristata is distributed mainly in subtropical
and tropical regions (Ellis, 1971; Farr et al., 2008). In areas with
temperate climate in Europe it was found almost exclusively in
countries with mild climate: Belgium (de Meulder, 2002), United
Kingdom, Denmark, Germany, Italy (Farr et al., 2008), and as Tetraploa
sp. in Spain (Diaz et al., 1998) and France (Proctor and Lambert, 1961).
So far records of the geographical distribution of another species, Te-
traploa ellisii Cooke are restricted to the subtropics and tropics (Ellis,
1971; Farr et al., 2008). The only Tetraploa species recorded exclusively
from European countries of temperate climate with continental
influence (Hungary and Lithuania) is T. setifera Révay (Révay, 1993;
Markovskaya, 2007).
Tetraploa aristata and Tetraploa ellisii commonly occur as sapro-
phytes on grasses and other monocotyledons on leaf bases and stems
just above soil (Ellis, 1971; Farr et al., 2008) although T. aristata was
also recorded on dicotyledons e.g. Citrus, Eucalyptus, Heracleum, Li-
quidambar, Quercus and also on Pinus (Farr et al., 2008) and rarely on
soil and decaying wood and leaf litter (Vittal, 1976; Révay, 1993).
Sometimes Tetraploa is found as aquatic fungus together with aquatic
hyphomycetes (e.g. Schoenlein-Crusius and Piccolo Grandi, 2003).
Conidia of Tetraploa often occur as accessory element among
airborne fungal spores in subtropical and tropical regions, e.g. India,
USA, Taiwan, Canary Island, Spain, Australia, and Martinique (Jothish
and Nayar, 2004; Codina et al., 2008; Tseng and Chen, 1982; Diaz et al.,
1998; Mitakakis and Guest, 2001; La-Serna et al., 2002; Desbois et al.,
2006) but sporadically they are aerial transported as far as Antarctica
(Marshall, 1996).
Interestingly, Tetraploa aristata occasionally could be a human
pathogen causing keratomycosis or phaeohyphomycotic cysts (Mark-
ham et al., 1990).
6.2. Palaeoecology of fossil bamboos
In the studied material pollen grains of bamboos were accom-
panied by abundant freshwater phytoplankton, as well as pollen and
spores of representatives of swamp-aquatic vegetation and riparian
forests. So, it would seem that in the Late Miocene the bamboos grew
in plant communities surrounding the water basin. They could be
components of riparian forests or grow at the edges of both riparian
and mesophytic forests as well. They also could occur in reed
association at the water's edges. Similar deductions could be inferred
on the basis of the pollen analytical results of the Miocene deposits
from Rypin, central Poland (Stuchlik, 1964) and the Middle Miocene
deposits from Legnica, southwestern Poland (Worobiec, 2000;
Worobiec and Worobiec, 2005), as well as investigations on fossil
leaves of bamboos “Bambusa” lugdunensis Saporta from the Upper
Miocene of the close locality Bełchatów Lignite Mine, central Poland
(Worobiec, 2003). Ivanov et al. (2007) recorded bamboo pollen grains
in Upper Miocene deposits from Bulgaria, and also regarded them as
components of riparian forests. Also the recent Arundinaria Michx.
bamboos growin swamp forests, along river and streambanks, in pine
forests or at the edge of flatwoods (Kearney, 1901; Johnson and
Hoagland, 1999; Lu and Liu, 2003; Wunderlin and Hansen, 2003).
6.3. Palaeoecology of fossil Tetraploa
Sharma (1976) and Saxena and Sarkar (1986) noted that fossil
conidia of Tetraploa are often associated with graminaceous pollen
grains. In the Józefina locality a conidium of Tetraploa cf. aristata is
also accompanied by numerous pollen grains of Poaceae, including
pollen grains of Graminidites bambusoides Stuchlik, which presumably
has a bamboo origin. Considering this it is probable that the discussed
Tetraploa occurred on grasses. Taking into account that recently three
new species of Tetraploa were described from bamboos from Japan
(Hatakeyama et al., 2005) and based on the data in literature (Ellis,
1971) as well as database record (Farr et al., 2008), G. bambusoides
bamboo could be a host plant of Tetraploa at the Józefina locality.
The occurrence of Tetraploa cf. aristata in Józefina points to a mild,
at least warm temperate climate of this period and confirms results of
pollen analysis suggesting a moderately wet and warm temperate
7. Conclusions
To conclude it would seem that bamboos were more common in
Europe in the Neogene time than we previously thought. Therefore, it
is necessary to pay more attention to pollen grains and leaves of the
bamboo type in fossil palyno- and macrofloras. The same attention
should be paid during palynological investigations on fossil Tetraploa
remains in pre-Quaternary deposits, especially when it is found in
association with monocotyledonous pollen grains. Tetraploa should be
considered as palaeoclimatical indicator of warm temperate climate.
We would like to thank Prof. Dr. hab. Leon Stuchlik and Prof. Dr.
hab. Ewa Zastawniak (Władysław Szafer Institute of Botany, Polish
Academy of Sciences, Kraków) for critically reading the manuscript,
as well as Prof. Dr. hab. Jerzy Głazek, Dr. Rafał Ratajczak and
Dr. Przemysław Dąbrowski (Adam Mickiewicz University, Poznań)
for supplying cored material from the Józefina borehole. Prof. Dr.
Steven R. Manchester (Florida Museum of Natural History and
Department of Botany, University of Florida) is gratefully acknowl-
edged for linguistic advice on the manuscript.
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