Saxon amber (Eocene, 3757 million years B.P.) consisting of 55
nologBaltic and Saxon amber is a valuable source for
fossils from many groups of organisms. It originated
in oakpine forests in Fennoscandia at the north
shore of a branch of an ocean during Eocene (57
37 Ma). The amber is attributed to the resin produc-
quaternary deposits from NE Germany to the Baltic
Sea. It is not exactly known how the amber accumu-
lated in the Oligocene and Miocene in Eastern
Prussia and Saxony (Schlee, 1990; Weitschat and
Wichard, 1998). There are two hypotheses for the
origin of the Baltic amber. According to the first1. Introduction grad, also in Saxony (Bitterfeld) and scattered inspecimens is reported. The specimens belong to extant species such as Trachycystis flagellaris (Sullivant and Lesquereux)
Lindberg, Trachycystis microphylla (Dozy and Molkenboer) Lindberg, Hypnodontopsis conferta (Goeppert and Berendt) J.-P.
Frahm, Atrichum cf. rhystophyllum (C. Muller) Paris, Haplocladium angustifolium (Hampe and C. Muller) Brotherus,
Tristichella glabrescens Iwatsuki, Ctenidium capillifolium (Mitten) Brotherus, Campylium cf. squarrosulum (Bescherelle and
Cardot) Kanda, not specified species of extant genera such as Aptychella, Campylopus, Campylopodiella, Barbella, Brotherella,
Brachythecium, Bartramia, Drepanocladus, Echinodium, Rhizogonium, and Symphyodon, but also to species described in form
genera such as Muscites pilifer J.-P. Frahm, Muscites serratus Goeppert and Berendt and Dicranites subflagellare Caspary and
Klebs. Several fossil mosses could not be identified but are described here as new as Hypnodonopsis pilifer J.-P. Frahm,
Eurohypnum revolutum J.-P. Frahm, Hypnum palaeocircinale J.-P. Frahm, Hypnites lanceolatus J.-P. Frahm and Hypnites
complanatus J.-P. The extant species Hypnodontopsis mexicana (Theriot) Robinson has proved to be synonymous with the
fossil H. conferta (Goeppert and Berendt) J.-P. Frahm, which has priority. T. glabrescens Iwatsuki and C. squarrosulum
(Bescherelle and Cardot) Kanda from Japan are reported for the first time from the Baltic amber.
D 2004 Published by Elsevier B.V.
Keywords: Mosses; Bryophytes; Baltic amber; Saxon amberSo far, the largest collection of mosses from the Baltic andReceived 3 May 2002; accepted 28 November 2003
AbstractA new contribution to the moss
Botanisches Institut der Rheinischen, Friedrich-Wilhelm-U
Review of Palaeobotany and Palytion of Pinus succinifera (Conwentz) Schubert. Large
masses of amber are found in marine deposits from
the upper Eocene or lower Oligocene near Kalinin-
0034-6667/$ - see front matter D 2004 Published by Elsevier B.V.
* Fax: +49-228-733-120.
E-mail address: email@example.com (J.-P. Frahm).ora of Baltic and Saxon amber
sitat, Meckenheimer Allee 170, D 53115 Bonn, Germany
y 129 (2004) 81101hypothesis, the resin was transported by streams into
the ocean; however, it is not exactly known how the
resin got into the streams in such masses. According
to another hypothesis, the amber forests were
inundated by a transgression (Schubert, 1961) during
which the trees were drowned, the resin was washed
out by tides, transported by ocean current and depos-
Amain problem with the identification of mosses in
amber is that relevant characters are only visible in
single leaves but not in whole plants, where leaf bases
with alar cells are not visible or laminal cells cannot be
studied in transmitting light. Therefore, a large number
of fossils could not be determined or only tentatively
assigned. Many of these fossils are, however, found
repeatedly. Therefore, these specimens are described
here in part as fossil species in extant genera, not
regarding that the discovery of sporophytes or better-
preserved specimens could reveal the identity with
and consists of 13 leaves that are somewhat com-
planate. The laminal cells are rounded. All these
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 8110182ited in clay sediments. Saxon amber is found in
deposits from the upper Oligocene or lower Miocene.
It is controversely discussed whether the Saxon
amber is just a reworked Baltic amber or of indepen-
dent origin. An argument for an independent origin is
that the amber was, according to IR-spectographic
studies, not produced by P. succinifera but by another
conifer, Cupressospermum saxonicum Mai.
Besides the amber produced by Pinus succinifera
(succinite), an amber with a different chemical com-
position (glesinite, redamite) is also known which
shows, that other plant families except for Pinaceae
and Taxodiaceae were also involved in the produc-
tion of resin.
The fossil moss flora of the Baltic amber was only
poorly known for more than 100 years. There were only
five publications (Goeppert and Berendt, 1845; Goep-
pert, 1853; Caspary, 1907; Dixon, 1922; Magdefrau,
1957) dealing with seven species, which all were
described in form genera. Since 1994, the author, a
moss taxonomist, started to study mosses from amber
(Frahm, 1994, 1996a,b, 1999a,b, 2000a,b, 2001a,b)
and increased our knowledge of the moss flora of the
Eocene considerably. In total, 150 species of mosses
are known from the Tertiary of Europe (Jovet-Ast,
1967; Miller, 1984; Frahm, 2000b) as compared to
about 1200 species at present. However, most records
are from the Pliocene (95 species) and Miocene (49
species). There are only four species recorded from the
Oligocene and none from the Palaeocene. All records
except for one from the Eocene are from the Baltic and
Saxon amber. They concern 47 species (Frahm, 2001a),
of which 27 species are extant species and the remain-
ing were described as fossil species or in form genera.
The authors studies were predominantly based on
specimens provided by amateur collectors, who con-
tributed much to the knowledge of the moss flora of the
amber forest. By courtesy of Christel and Hans-Werner
Hoffeins (Hamburg), Carsten Grohn (Glinde), Heinrich
Grabenhorst (Celle),ManfredKutscher (Sassnitz), Jens
von Holt (Hamburg), Franziska Witsch (Koln) and
Prof. Dr. Wichard (Bonn) I received again numerous
mosses from the Baltic and Saxon amber, which are
described and illustrated here. The Grabenhorst and
Kutscher collections consist of the Saxon amber (Bit-
terfeld), all others from the Baltic amber. The speci-
mens are kept in the private collections of thecollectors.characters refer to the genus Mnium; however, the
laminal cells are mamillose, which undoubtedly
places this specimen in the genus Trachycystis.
In total, six species (three fossil and three extant)
are known from the genus Trachycystis. Of the three
Fig. 1. Trachycystis flagellaris (Grohn 2023), plant.
Fig. 2. Trachycystis flagellaris (Grohn 2023), leaves.
Fig. 3. Trachycystis microphylla (Wichard 14).
Fig. 4. Trachycystis flagellaris and T. microphylla (Wichard 14).
Fig. 5. Hypnodontopsis conferta (Grohn 2031).extant species. In addition, several specimens are listed
and described, which cannot be determined and not
even attributed to a genus. They are nevertheless
included to show the richness and diversity of the
Eocene moss flora.
2.1. Trachycystis flagellaris (Sullivant and Lesquer-
eux) Lindberg (Grohn 2023, Grabenhorst La 3,
Wichard 14 p.pte.)
Grohn 2023 (Figs. 1 and 2) consists of a 7-mm-
long part of an acrocarpous moss with 14 distant
leaves. The leaves have a percurrent costa, which is
serrate at the dorsal side. The leaf margin is bordered
and shows paired teeth. Grabenhorst La 3 is similarFig. 6. Hypnodontopsis conferta (Grohn 2042).
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101 83
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 8110184fossil species, two have been reported from the
Baltic amber (Frahm, 2000a). A key for all extant
and fossil species of the genus is given by Frahm
(2001a). The combination of characters of bordered
leaves with paired teeth refers to the species Trachy-
cystis flagellaris (Fig. 4), which is still found in
Eastern Asia. The flagellate branches typical for this
species have, however, never been found in fossil
specimens. The fossil Trachycystis microphylla is
similar with regard to the leaf shape but has no
Trachycystis szaferi Szafran, described as fossil
species from the Miocene of Poland (Szafran, 1958),
shall be differentiated by the other species of the genus
by a costa toothed at the dorsal side. This character also
concerns, however, Trachycystis flagellaris (Noguchi,
19871994), and also matches the specimen La 3.
Therefore, the fossil T. szaferi is probably synonymous
with the extant T. flagellaris.
Trachycystis flagellaris was so far recorded once
from the Baltic amber (Caspary, 1907 as Muscites
hauchecornei; Frahm, 1994) and exists stillas all
extant species of this genusin East Asia (Koponen,
2.2. Trachycystis microphylla (Ddozy and Molken-
boer) Lindberg (Hoffeins 1161.3, Wichard 14 p.pte.)
Hoffeins 1161.3 consists of 14 intact plants as
well as several other less well-preserved specimens
of 57 mm length. The leaves of most plants are
somehow flattened (perhaps by embedding), but
there is one single plant with normally spirally
arranged leaves. The leaves are oblong ovate, acu-
minate, with percurrent costa and serrate margins.
The preservation of the leaves is so perfect that the
rounded small laminal cells are visible which are
distinctly mamillose. The specimen can be attributed
to the genus Trachycystis because of the Mnium-like
appearance, however, with mamillose laminal cells.
Because of a lacking border and unpaired teeth, the
specimen can be attributed to Trachycystis micro-
phylla (Frahm, 2001a). It has already been reported
from the Baltic amber and exists still in East Asia
By a unique chance, two plants of different species
of Trachycystis were imbedded in Wichard 14. Thefirst plant is about 5 mm long and has 20 distantleaves, which are not bordered and have single teeth
along the leaf margins (Fig. 3).
The second plant (Fig. 4) has about the same size,
but has 18 somewhat narrower leaves with double
teeth along the leaf border, characters, which refer to
Trachycystis flagellaris. This remarkable specimen
reveals that both species were sympatric and existed
at the same time in the same place.
2.3. Hypnodontopsis conferta (Goeppert and Berendt)
J.-P. Frahm (Grohn 616, Hoffeins 1417/1, 2, Gra-
benhorst La 23, 24, Von Holt 2)
These specimens belong to the most common moss
in the Baltic amber. It was described already in 1845 by
Goeppert and Berendt (1845) as Muscites confertus.
Part of the syntypes of Dicranites casparyi also belong
here (Frahm, 2001a), which was described by Klebs in
Until recently, only sterile specimens were known.
The first fertile specimen was published by Frahm
(1999b) as Dicranites grollei. At that time, the iden-
tity of the fossil with the extant genus Hypnodontopsis
was not known, mainly because this genus is repre-
sented by only two species: Hypnodontopsis apiculata
Iwatsuki and Noguchi from Japan and Hypnodontop-
sis mexicana (Theriot) Robinson described from Mex-
ico and known there only from the type collection in
Michoacan and recently reported for Uganda. Another
fertile specimen was reported by Frahm (2001b),
which was then attributed to the recent genus Hyp-
nodontopsis (Rhachitheciaceae). An identity with H.
apiculata could be excluded because of a different
leaf shape, but H. mexicana substantially resembles
the fossil specimens. A synonymy could, however,
not be confirmed because H. mexicana has pluripa-
pillose laminal cells, which could not be observed in
the fossil material. Therefore, Muscites confertus was
combined to the extant genus Hypnodontopsis as
Hypnodontopsis confertus (Frahm, 2001b). These
plants are characterized by narrowly lanceolate leaves
with percurrent costa prominent at the back and
conspicuously only 56 rows of isodiametric laminal
cells at each side of the costa.
Grohn 2031 (Fig. 5) has by chance large decol-
ourized transparent parts of the lamina, which make
distinct papillae visible characteristic for Hypnodon-topsis mexicana. The latter has also the conspicuous
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101 85few rows of laminal cells (cf. Fig. 447 in Sharp et al.,
1994). Therefore, the extant species H. mexicana is
most likely identical with the fossil Hypnodontopsis
conferta, a name, which antedates H. mexicana.
According to Art. 11.7 of the ICBN (see also exam-
ple 27), the names of fossil plants have priority over
the names of extant species. Therefore, the name H.
conferta has to be used for H. mexicana:
Hypnodontopsis conferta (Goeppert and Berendt)
J.-P. Frahm, Trop. Bryol. 20: 80, 2001 (confertus)
Muscites confertus (Goeppert and Berendt) Bernst.
Org. Reste Vorwelt 1: 112, 1845)
Hypnodontopsis mexicana (Ther.) Robins., Bryol-
ogist 67: 449, 1964, syn. nov.
Dicranites grollei (J.-P. Frahm) Haussknechtia
Beih. 9: 129, 1999, syn. nov.
Oreas mexicana (Ther.), Smiths. Misc. Coll. 81: 2,
Since the Greek name opsis is feminine and the
gender is kept in the latinized genus name, the species
name must be conferta and not confertus as in Frahm
Besides this species, another species, Hypnodon-
topsis fossilis J.-P. Frahmwas described from the Baltic
amber (Frahm, 2000a), which differs by shorter setae
(0.5 mm instead of 1.52 mm long), longer, narrower
leaves and long cylindric shape of the capsule.
Except for the first fertile specimen of Muscites
confertus, from which the generic position of this
moss could be concluded, several more fertile speci-
mens showed up within a short period of time:
Grohn 2042 (Fig. 6) is a spectacular specimen with
one sterile and one fertile plant with well-developed
sporophyte. It shows a twisted seta characteristic for
this genus, a short cylindric capsule with eight striae
and peristome teeths united to eight pairs. Similar to
Grohn 2042, parts of the lamina are decolourized and
make the papillae visible. The seta is 1.5 mm long,
thus resembling H. conferta.
In Hoffeins 1417/1, the capsule is longer as in
Grohn 2031. Grabenhorst La 23 (Fig. 7) is a conspic-
uous specimen with four fertile plants and numerous
sterile plants. The setae are 1.5 mm long, twisted,
while the capsules (Fig. 8) are 0.5 mm long and ovoid
to almost globose, with eight distinct striae. Theperistomes are no longer preserved. La 24 includes adozen single plants, one of which has a sporophyte in
the same shape as La 23.
Hypnodontopsis conferta is the most common
moss found in the Baltic and Saxon amber. It was
already described and illustrated by Goeppert and
Berendt (1845, Taf. VI, Figs. 2931). Caspary
(1907, figs. 4445) included it in the syntypes of
Dicranites casparyi, a reason, that this species had
later to be lectotypified (Frahm, 2001a). Probably
because it was an epiphyte, it produced terminal leaf
rosettes which were easily spread. It has previously
been reported from numerous collections by Frahm
(1999a, 2000a, 2001b) as acrocarpous moss or
In addition, there is also a number of sterile speci-
mens of Hypnodontopsis (Hoffeins 1343/2,4,5,6,
Grohn 2040, Grabenhorst 6, 8, 9, 16, von Holt 2),
which show the characteristic longly lanceolate leaves
with percurrent costa and few rows of round cells at
both sides of the costa. Grabenhorst 9 includes several
sterile plants. Grohn 2040 consists of an apical rosette
of leaves, which apparently serve for vegetative
propagation. Hoffeins 1343/2, 4, 5, 6 as well as
Grabenhorst 6 and 16 represent bud-like rosettes of
leaves, too. Most fossil specimens of Hypnodontopsis
consist of such rosettes.
2.4. Hypnodontopsis fossilis J.-P. Frahm (Hoffeins
1417/2, Grabenhorst La 7, 14)
This species has the typical vegetative andgenerative
characters of the genus Hypnodontopsis as discussed
under Section 2.3; however, much longer and narrower
capsules (four times longer than broad) as compared
with Hypnodontopsis conferta, which are gradually
merging into the seta and not abruptedly as in H.
conferta (cf. Fig. 9). Furthermore, the leaves are longer
and narrower. Such a plant has been described as H.
fossilis J.-P. Frahm(2000a).Grabenhorst La14 (Fig. 10)
consists of a single plant with perfectly preserved
peristome teeth, which are erect, spreading and united
into eight pairs.
Vegetatively, the plants substantially resemble the
sterile specimens of Dicranites casparyi (cf. Caspary,
1907, fig. 43, lectotype of the latter, not figs. 4445,
which belong to Hypnodontopsis conferta) as well as
Dicranites obtusifolius (Caspary, 1907, fig. 46). How-ever, the few longitudinal rows typical for the fossil
Diagnosis: Plantae steriles, foliis dense appressis,
anguste linearibus, costatis, margine anguste revolu-
tis. Costa in pilum longum excurrens. Cellulae lam-
inae subquadratae, mamillosae, seriatae in 56
The new species is easily distinguished from all
other species of the genus by its leaves ending in a
The genus Hypnodontopsis now comprises four
species: Hypnodontopsis apiculata confined to Ja-
pan, Hypnodontopsis conferta known from Mexico
and Uganda and also from the Baltic and Saxon
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 8110186specimens of Hypnodontopsis cannot always be rec-
ognised, and the type of D. casparyi and D. obtusi-
folius are sterile that the sporophytes typical for the
genus Hypnodontopsis cannot be studied. Further-
more, there are plants similar to those of H. fossilis
(Kutscher 3, 7, Frahm, 1999a as Dicranaceae; Hof-
feins 874, Frahm, 2000a as D. casparyi) which have,
however, an elongate seta and can therefore not be
conspecific with H. fossilis. Therefore, D. casparyi
and/or D. obtusifolius may also belong to this (un-
Hypnodontopsis fossilis was so far only known
from the type specimen (Frahm, 2000a).
2.5. Hypnodontopsis pilifer J.-P. Frahm sp. nov.
The amber includes a relatively large piece of bark
of 3.5 2 cm, which is fully covered with a crusta-ceous lichen with lecanoroid apothecia. Two plants
of an epiphytic hepatic (Frullania sp.) are creeping on
the lichen. Attached to the lichen is a small fragment
of an apparently young moss plant with several small
(12 mm long) branchlets and narrow linear longly
apiculate leaves (Fig. 11). A higher magnification
(Fig. 12) reveals that the leaves have a costa which
is excurrent as long straight hair, that the leaf margins
are narrowly recurved and that the lamina is formed
by few rows of round, mamillose cells arranged in
A similar fossil moss with such narrow linear
leaves and long hairpoint was described as Muscites
pilifer (Frahm, 1999b). In the type specimen of the
latter species, the shape of the laminal cells is not
visible; however, a similar plant described here shows
elongate laminal cells, that this specimen seems not to
Round mamillose laminal cells arranged in few
longitudinal rows are highly characteristic for the
extant species Hypnodontopsis conferta (syn. H.
mexicana), see Section 2.3, and also for the fossil
species H. fossilis, see Section 2.4. It could be argued
that this is a characteristic of at least some species of
the same genus, and therefore this specimen is de-
scribed here as a new (fossil) species of the genus
Holotype: coll. Witsch #1. Etymology: The speciesis named after its costa ending in a long hairpoint.amber, as well as Hypnodontopsis fossilis and
Hypnodonopsis pilifer only known from the Baltic
and Saxon amber. Fossil records of H. conferta are
much more numerous than the two extant records.
In addition, fossil species are more numerous than
extant species that it can be argued that the genus
Hypnodontopsis has reached its peak of speciation
in the Tertiary.
2.6. Campylopodiella sp. (Grohn 2028, Grabenhorst
La 13, 15, 19, 21, 22)
Grohn 2028 (Figs. 13 and 14) consists of a bundle
of several plants 3 mm high, as well as a single plant
with some separate leaves. The leaves are erect,
concave-tubulose and give the impression of Leuco-
bryum leaves. Some leaves show a dark stripe in the
middle of the leaf, which is typical for the genus
Campylopodiella and caused by a group of stereids in
transverse section of the costa.
Grabenhorst La 22 consists of a dense mat of
several sterile plants with the same characteristics.
La 13, 15 and 21 contain only few plants, but several
Fig. 7. Hypnodontopsis conferta (Grabenhorst La 23), several fertile
Fig. 8. Hypnodonopsis conferta (Grabenhorst La 23), capsule.
Fig. 9. Hypnodontopsis fossilis (Hoffeins 1417/2).
Fig. 10. Hypnodontopsis fossilis (Grabenhorst La 14).
Fig. 11. Hypnodontopsis pilifer (Witsch 1), plant.Fig. 12. Hypnodontopsis pilifer (Witsch 1), branch.
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101 87
genus is discussed. However, the fossil specimens
differ from the extant species in that they have
considerably smaller leaf length. Extant species are
distinguished by the position of the perichaetia, a
character which is not expressed in the fossil material.
Species of Rhizogonium substantially resemble those
of Trachycystis, which is shown by the fact that T.
microphylla (Doz. and Molk.) Lindb. was also de-
scribed as Rhizogonium sieboldii Hampe, but differ by
narrow lanceolate leaves, which are distantly arranged
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 8110188separate leaves, in which the broad costa is visible,
which occupies 3/4 of the width at leaf base and fills
the leaf apex. Similar specimens have been found
twice in the Saxon and Baltic amber (Frahm, 1996b,
1999a), one with sporophytes typical for the genus
with cylindric capsules and long, twisted setae. La 19
is attributed to this species with some reservation
because the plants are intermixed with dirt.
The genus Campylopodiella (Dicranaceae, Paral-
eucobryoideae) is today represented by two species
in Latin America, one in Papua New Guinea and
one in the eastern Himalaya. Of these species, the
fossil material resembles C. himalayana (Brotherus)
J.-P. Frahm occurring from Bhutan to Yunnan, as
well as C. stenocarpa (Wilson) P. Muller and J.-P.
Frahm occurring in Central America. Both are very
similar (Muller and Frahm, 1987). The latter differs
from C. himalayana by longer capsules, width of
the costa and broader lamina at leaf base, thus not
significantly. The leaf characters cannot be proved
in the fossil material. Capsules were found in fossil
material in Grohn 1022 (Frahm, 1999a) and Strie-
bich s.n. (Frahm, 1999b). The old empty capsules
are four times as long and as wide in both speci-
mens. Striebich s.n. shows in addition a young
capsule which is only 2.5 times longer than wide.
In contrast, the capsules of C. stenocarpa are five
times longer than wide and those of C. himalayana
23 times. Therefore, the fossil material cannot be
attributed with certainty to one of the extant spe-
cies. It can be assumed that both extant species are
very closely related and are perhaps sister species,
which may have been derived from the Eocene
ancestor described here, which has an intermediate
2.7. Rhizogonium sp. (Hoffeins 1416)
The amber includes two parts of plants of 5 viz.
7 mm length, which are inadequately visible. Be-
sides, there are several leaves or fragments of
leaves 1.4 mm long, which are perfectly preserved.
They are lanceolate with percurrent costa, bordered
leaf margin with paired teeth and round laminal
cells (Figs. 15 and 16).
The species is apparently identical with Kutscher
M24, which was attributed to the genus Rhizogonium(see also Frahm, 2001a), where the affiliation to thisalong the stem, longer teeth along the leaf margins
and smooth laminal cells.
The genus is mainly distributed in SE Asia and
2.8. Atrichum cf. rhystophyllum (C. Muller) Paris
The piece of amber includes several plants of 5
8 mm length, with 3.5- to 4-mm-long, broadly lance-
olate, patent leaves (Figs. 17 and 18). These are
mostly incrusted by dirt, but few leaves are clearly
visible. These show four relatively high lamellae on
the ventral side of the costa, a character, which is
characteristic for the genus Atrichum (Polytrichaceae).
The laminal cells are round, the leaf margin bordered
with paired teeth. Another specimen of Atrichum
(Grohn 2019) was already reported by Frahm
(2000b). They possess six costal lamellae. At present,
it cannot be determined whether or not both speci-
mens belong to the same species.
A fossil species of Atrichum, Atrichum subundu-
latum Goeppert, was already described in the 19th
century. The type specimen is lost and there exists
neither a description nor an illustration of this species
that the name A. subundulatum must be regarded as a
nomen dubium, and our specimen cannot be com-
Fig. 13. Campylopodiella sp. (Grohn 2028).
Fig. 14. Campylopodiella sp. (Grohn 2028).
Fig. 15. Rhizogonium sp. (Hoffeins 1416), several leaves.
Fig. 16. Rhizogonium sp. (Hoffeins 1416), single leaf.
Fig. 17. Atrichum cf. rhystophyllum (Grohn 2041).Fig. 18. Atrichum cf. rhystophyllum (Grohn 2041).
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101 89
and Saxon amber such as Hoffeins 1161/1, perhaps
also Kutscher M22 (Frahm, 2001a) and Grohn 869
(Frahm, 1999a) and named as Hypnum sp. Ker-
negger 1992/27 consists of a 3-mm-long stem with
leaves of identical shape. Grohn 2034 is a fragment of
a hypnaceous moss with one branch with similar
narrow lanceolate, hamate leaves. The leaves are
conspicuously ovate and concave at base, sheathing
the stem, and end in a very long acumen. Although
not as regularly hamate as Hoffeins 1417b, it seems to
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 8110190pared with that species. In contrast to the extant A.
undulatum, the leaves are much smaller and not
transversely undulate. The fossil plants resemble,
however, the extant A. rhystophyllum with regard to
plant and leaf size, leaf shape, position of leaves
(leaves of equal size and shape equally arranged along
the stem) and number of costal lamellae. A final
decision, whether the fossil specimen is identical with
the extant species, cannot be made because this would
require a transverse section of the costa to determine
the length of the costal lamellae. On the other hand,
there is no apparent characteristic that would distinctly
differentiate the fossil specimen from the extant spe-
cies. Therefore, the fossil material is tentatively re-
ferred to the extant A. rhystophyllum, a species
distributed today in East Asia and Central America
2.9. Haplocladium angustifolium (Hampe and C.
Muller) Brotherus (Grohn 2039, Damzen 120, Von
Grohn 2039 consists of a fragment of a pleuro-
carpous moss of 4 mm length with dense foliation.
Damzen 120 (Figs. 19 and 20) consists of a 4-cm-
long, regularly pinnate plant. The leaves are from
ovate base narrowly acuminate and have a percur-
rent costa which is prominent at the back. The leaf
margins are revolute, the laminal cells round. Von
Holt 3 is tentatively placed to this species. It
consists of a fragment 1 cm long, which is less
well preserved but shows the same leaf shape, a
percurrent costa and rounded laminal cells.
Such specimens were already reported several
times from the Baltic amber (Frahm, 1996a,b,
1999a,b as Leskeaceae) and identified as Hapocla-
dium angustifolium (Frahm, 2001a), a species found
mainly in southern Asia and Central America. In
Europe, it is found only in the Southern Alps,
where it is regarded as a relict from the Tertiary.
2.10. Hypnum palaeocircinale sp. nov. (Hoffeins
1417b, Kernegger 1992/27, Grohn 2034)
A 4-mm-long part of a stem of a densely foliate
pleurocarpous moss with strongly circinate (almost
180j) and homomallous leaves (Fig. 21). A specimen
of this species was already reported from the Balticbe identical with the latter. Similar plants were
reported by Frahm (1996a, 1999a).
Although the specimens resemble East Asiatic spe-
cies of this genus in appearance, they cannot be
attributed to one of these extant species with certain-
tythe reason being that anatomical characters such as
the basal laminal cells or the alar cells are not visible in
the specimens in amber. Because there are several
representatives of the genus Hypnum in the Baltic
and Saxon amber, these specimens are described here
as new as fossil species to avoid confusion with other
similar specimens named also Hypnum sp.
Holotype: Hoffeins 1161/1, Paratypes: Hoffeins
1417b, Kernegger 1992/27, Grohn 2034. Etymolo-
gy: The species is named after its resemblance to
the extant Hypnum circinale. Diagnosis: Plantae
specie generis Hypni persimilis, cum foliis valde
circinatis, homomallis, ecostatis, in apicibus longis
irregulariter serratis excurrentibus.
The species differs from all other representatives of
the genus in the Baltic or Saxon amber by its large
size, dense foliation with strongly circinate leaves.
2.11. Campylopus sp. (Grabenhorst La 7)
La 7 includes merely a single, but very clearly
embedded sporophyte with twisted seta and capsule,
Fig. 19. Haplocladium angustifolium (Damzen 120).
Fig. 20. Haplocladium angustifolium (Damzen 120).
Fig. 21. Hypnum paleocircinale sp. nov. (Hoffeins 1417/4).
Fig. 22. Campylopus sp. (Grabenhorst La 7), capsule.
Fig. 23. Eurohypnum revolutum sp. nov. (Grabenhorst La 10), plant.Fig. 24. Eurohypnum revolutum (Grabenhorst La 10), branch.
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101 91
2.13. Tristichella glabrescens Iwats., Sematophylla-
ceae (Grabenhorst La 12)
This specimen includes (amongst other mosses)
three 4-mm-long branches of a pleurocarpous moss
with small, distant, erect spreading lanceolate, con-
cave leaves (Fig. 25). The leaves are conspicuously
arranged in three rows.
Mosses with tristichous arrangement of leaves are
met only very rarely, and if, usually amongst the
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 8110192of which even the exothecial cells are visible (Fig.
22). The capsule is 1.5 mm long, and the seta 2.5
mm long. A similar capsule without preserved
peristome was named as Campylopus sp. Both
genera, Campylopus and Hypnodontopsis, have
twisted and cygneous setae and Hypnodontopsis
fossilis has capsules with similar shape; however,
Campylopus has a haplostomous, haplolepideous
peristome, Hypnodontopsis a diplostomous, diplole-
pideous one. Furthermore, the capsules of Hypno-
dontopsis from amber are 0.5 mm long, the setae
1.5 mm or less, whereas the present sporophyte is
much larger and can therefore be attributed to the
Campylopus is a genus with about 160 extant
species worldwide, and occurs mainly in tropical
montane and subantarctic regions. A determination
by sporophyte characters is not possible.
2.12. Eurohypnum revolutum sp. nov. (Grabenhorst
The preserved plant is 1 cm in length with creeping
stem and several erect, 34 mm long branches (Figs.
23 and 24). The branches are densely and very
imbricately foliate. The branch leaves are ovate and
narrow into a long, fine acumen. Leaf margins are
conspicuously narrowly revolute. The leaves have no
costa and elongate laminal cells.
Pleurocarpous mosses with ecostate leaves and
prosenchymatous laminal cells must be placed into
the Hypnaceae or Sematophyllaceae. The latter have
often papillose laminal cells and a different appear-
ance without julaceous stems, which are found in the
Hypnaceae. This specimen resembles certain species
of Hypnum with julaceous branches, but has straight,
not homomallous leaves. It resembles with its sub-
pinnately branched stems, terete and imbricate folia-
tion and leaf shape Eurohypnum leptothallum (C.
Mull.) Ando from Japan (Noguchi, 19871994), a
monotypic genus close to Homomallium, but differs in
terms of longer leaf apices and revolute leaf margins.
Holotype: collection Grabenhorst La 10. Etymolo-
gy: The species is named after its revolute leaf margins.
Diagnosis: Caulis 1 cm longa, ramosus, ramis rectis,
dense imbricatis. Folia ovata, in apicem angustum
contracta, marginibus revolutis, since costa. Cellulaelaminae elongatae.acrocarpous taxa. The only pleurocarpous genus with
tristichous leaves is the genus Trichistichella Dixon.
The present fossil matches perfectly the description of
T. glabrescens Iwats., a species from Japan and Taiwan
(Noguchi, 19871994), growing on branches of trees.
Kutscher M 25 (Frahm, 2001a: 275 without name)
belongs to the same species.
2.14. Sematophyllaceae (Grabenhorst La 12)
Associated with Tristichella glabrescens is another
4.5-mm-long piece of another moss, with broadly
lanceolate leaves that are complanate (Fig. 26). The
leaf base is sheathing the stem and decurrent along the
stem. The leaves are broadly lanceolate acuminate,
ecostate, with serrulate margins at the leaf tip and
elongate, smooth laminal cells.
All these characteristics place this moss into the
Sematophyllaceae, which has many similar represen-
tatives in different genera.
2.15. Ctenidium capillifolium (Mitt.) Broth.
(von Holt 4)
The specimen consists of a 6-mm-long fragment of
a pleurocarpous moss with one branch (Figs. 27 and
28). The plant is densely foliate with extremely
Fig. 25. Tristichella glabrescens (Grabenhorst La 12).
Fig. 26. Sematophyllaceae (Grabenhorst La 12).
Fig. 27. Ctenidium capillifolium (von Holt 4).
Fig. 28. Ctenidium capillifolium (von Holt 4).
Fig. 29. Campylium squarrosulum (von Holt 6).Fig. 30. Campylium squarrosulum (von Holt 6).
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101 93
long flagellae. This (in connection with the leaf shape,
shape of laminal cells and percurrent costa) is typical
for the genus Barbella (Meteoriaceae), a tropical
genus, which goes form north to Mexico and southern
Japan and typically grows pendant from branches.
Similar plants have been reported from the Baltic
amber by Frahm (1996a: 132 Taf. 2, figs. 78;
2.18. Brotherella sp. (Grohn 619)
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 8110194narrow lanceolate, longly acuminate leaves, which are
slightly incurved at the apex. Details of the leaf
anatomy are not visible.
The specimen resembles several other fossils
(Grohn 869, Wichard 326, Teuber 1565: Frahm,
1999a: 235, fig. 13; Teuber 1606: Frahm, 2000a: 34;
Wenzel s.n.: Frahm, 2000b: 126, fig. 6) with regard to
the appearance, leaf shape and incurved leaf apices,
which were referred, also because of prosenchymatose
laminal cells, serrate leaf margins and ecostate leaves,
to Ctenidium capillifolium. This species is distributed
in China, Taiwan, Korea and Japan.
2.16. Campylium cf. squarrosulum (Besch. and
Card.) Kanda (von Holt 6)
The amber includes several pieces of a pleuro-
carpous moss with length of up to 7 mm (Figs. 29
and 30). The plants are conspicuously foliate with
patent leaves. The leaves are distant, concave at
base and tubulose in the apex, from ovatecordate
base gradually contracted into a fine acumen. The
margins are finely serrate, the laminal cells short,
elongate oval. The costa is short and reaches to
midleaf. Plants with such squarrose distant leaves
with this typical shape are found in the genus
Campylium (Amblystegiaceae). Therefore, this spec-
imen can be referred to this genus. The fossil plants
match perfectly the description and illustration of
Campylium squarrosulum in Noguchi (19871994),
occurring in Japan except for details of the areola-
tion, which are not visible in the fossil. Therefore,
the specimen is tentatively referred to this species.
Most species of this genus grow in bogs and fens;
however, this species grow on rotten wood.
2.17. Barbella sp. (von Holt 1, 5)
In von Holt 1, a bunch of three pleurocarpous
mosses is preserved, which are connected at the base
(Figs. 31 and 32). The plants are 1.52 cm long and
are loosely and irregularly pinnate. The leaves are
from broader base gradually contracted into a long
fine point. Von Holt 5 is in a similarly bad condition.
Because of the relatively bad preservation, details
of the leaves are hardly visible. Laminal cells are
visible in only a few leaves and elongate; a costa isnot visible. However, some of the branches end inGrohn 619 consists of a 5-mm-long plant with some
branches. The leaves are distinctly complanate foliate,
ovate lanceolate and finely pointed into an acumen,
which is serrate and often twisted. Such plants were
described as Muscites tortifolius Caspary and Klebs
(Caspary, 1907) and later recorded as Brotherella sp.
from several specimens from the Baltic and Saxon
amber by Frahm (1996a, 2000a,b, 2001a). The reason
is that ecostate leaves with prosenchymatous laminal
cells, serrate leaf tips and complanate foliation are only
found amongst the Sematophyllaceae in the genera
Brotherella and Heterophyllium. Both genera can only
be distinguished by their alar cells, which are visible
only in detached leaves.
2.19. Brachythecium sp. (Grabenhorst La 18)
This specimen shows (amongst fragments of anoth-
er moss species) fragments of a pleurocarpous moss
with ovate lanceolate, finely tipped leaves, serrate
margins and a costa reaching 3/4 of leaf length. The
laminal cells are prosenchymatous. This combination
of characters is realized in the genus Brachythecium
(Hypnales, Brachytheciaceae), a species-rich genus
with mainly holarctic distribution. The plant resembles
Fig. 31. Barbella sp. (von Holt 1).
Fig. 32. Barbella sp. (von Holt 1).
Fig. 33. Muscites pilifer (Grabenhorst La 27).
Fig. 34. Drepanocladus sp. (Kernegger 1998/117).
Fig. 35. Hypnaceae (Hoffeins 1343/1).Fig. 36. Muscites serratus (Grabenhorst La 26).
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101 95
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 8110196much those of Brotherella sp. (cf. Section 2.11), which
has, however, ecostate leaves.
2.20. Bartramia sp. (Grabenhorst La 18)
In the same specimen with Brachythecium sp.,
there are upper parts of two plants and single leaves
of a moss with narrowly lanceolate, canaliculate
leaves, percurrent costa, round upper laminal cells
and sheathing leaf base. Such plants have been named
before as Bartramia sp. (Frahm, 2001a). Such names
are suppositions and give the nearest possible guess;
however, they help to mark identical fossil specimens.
2.21. Muscites pilifer J.-P. Frahm (Grabenhorst
The amber contains part of a stem or a branch of 3
mm length (Fig. 33). The leaves are very narrow and
elongate, have elongate laminal cells and are con-
tracted into a piliferous leaf tip of 1/41/3 leaf length.
The leaf tip is hyaline. The specimen resembles M.
pilifer described by Frahm (1999b). This species seems
not to match any extant species. The branching of the
type of M. pilifer as well as the elongate laminal cells,
which are visible in this specimen, seem to indicate a
pleurocarpous moss. However, the extant pleurocar-
pous species have not such hairpoints. It could be that
the leaf apices are decolourate and in fact, no hyaline
hairpoints but very longly acuminate leaf tips.
2.22. Hypnaceous moss (Grabenhorst La 20)
Here, a 2-cm tuft of a pleurocarpous moss is
embedded, which is densely branched with short
branches and densely foliate with very narrowly
lanceolate, very gradually narrowed acute leaves,
which are apparently ecostate and have elongate
laminal cells. It can be attributed to the families
Sematophyllaceae or Hypnaceae, in which numer-
ous species in several genera match this description.
2.23. Drepanocladus sp. (Kernegger 1998/117)
A terminal part of the branch or a stem of a pleuro-
carpousmoss of 1.5mm length (Fig. 34). The leaves are
narrowly lanceolate, hamate, and longly and finelyapiculate.Thecosta is robust andreaches into the leaf tip.Similar fossils with narrow, hamate leaves were
reported several times (Frahm, 1996a: 133; Frahm,
1999a: 229, fig. 13; Frahm, 2001a: 277, fig. 11);
however, they lack any costa and were therefore
attributed to the Hypnaceae. The present specimen
has, however, a very distinct costa. Hamate, costate
leaves are found in the genus Drepanocladus
(Amblystegiaceae). Most species of this genus are
growing in swamps, but Drepanocladus uncinatus
also epiphytic in damp conditions.
2.24. Hypnaceae (Hoffeins 1343/1)
A 1.5-mm-long apical fragment of a pleurocarpous
moss in the Saxonian amber with hamate, narrowly
lanceolate leaves and complanate foliation (Fig. 35).
The leaves have a prosenchymatous areolation and no
Hoffeins 962/2 as well as Teuber 1606 are very
similar (Frahm, 2000a: 34, figs. 3 and 4). They were
named as Hypnaceae.
2.25. Echinodium sp. (Kutscher 15, Witsch 6)
This specimen was already described in Frahm
(1999a) and also illustrated by Kutscher (1999,
Tafel 3 Bild 6) but was erroneously identified as
Haplocladium angustifolium. (Haplocladium is nev-
ertheless present in the Baltic and Saxon amber, see
Section 2.8). A revision revealed that it belongs to
the genus Echinodium (Echinodiaceae), which is
characterized by narrow, longly acuminate, very
densely foliate leaves with costa. Within this genus,
Echinodium savicziae A. Abramov and I. Abra-
mova is known from the Pliocene of Russia from
a subtropical to warm temperate palaeo-environ-
ment, although it is represented at present with
two species in New Zealand and four species in
Macaronesia (Churchill, 1986). Therefore, it can be
concluded that the species got extinct in Eurasia
perhaps at the beginning of the Quaternary but
survived in New Zealand and Macaronesia. It is
known that the Macaronesian Islands harbour a
larger number of flowering plants which were
present in Tertiary in Europe.
Another four specimens are included in Witsch 6,
two larger 10 viz. 12 mm long, and two smaller 3 viz.5 mm long. The leaves of the smaller specimens are
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101 97longitudinally split and give the impression of a
2.26. Plagiotheciaceae (Grohn 618, 2033)
Grohn 2033 consists of a 2-mm-long fragment of a
pleurocarpous moss with extremely complanate folia-
tion. The leaves are narrowly lanceolate and ending in a
long tip. They are ecostate, and have prosenchymatous
laminal cells and entire margins.
Grohn 618 consists of a 15-mm-long piece of a
moss. Although the leaves are substantially eroded,
they have apparently no costa and prosenchymatous
cells. Conspicuous is the complanate foliation.
Fragments of this species were already often found
in the Baltic amber (Kutscher 2b, Hoffeins 962/2,
Hoffeins 1161/2, Grohn 370, cf. Frahm, 1999a,
2000a,b). They were referred to the family Plagiothe-
ciaceae because of their complanate foliation with
ecostate leaves and elongate laminal cells, a combina-
tion of characters, which is found only in this family.
They resemble Brotherella sp. (cf.), which has, how-
ever, serrate leaf tips and twisted leaf apices.
2.27. Muscites serratus Goepp. and Berendt (Gra-
benhorst La 26, Grohn 615)
A 2-mm-long apical part of the stem or branch of a
pleurocarpous moss with narrow lanceolate, plane,
ecostate leaves, which have prosenchymatous laminal
cells (Figs. 36 and 37). The leaf margins are serrulate.
Grohn 615 is a pleurocarpous moss of 17 mm length
with 11 short, erect branches. The leaves are ecostate,
lanceolate and have elongate laminal cells. The plant is
covered with 11 claviform structures, which originate
from the stem as well as from the branches. They could
be sporangia of parasitic fungi.
Similar plants were reported more frequently from
the Baltic and Saxon amber as Hypnum sp. (Frahm,
1996a, 1999a). In addition, the Pleurokarpes Laub-
moos 5 in Frahm (1999a) may belong here as well as
KutscherM 25 (Frahm, 2001a, as pleurocarpousmoss).
Such a fragment was described by Goeppert and
Berendt (1845) as Muscites serratus. Because of the
bud-like shape, the authors compared it with the moss
genera Phascum and Gymnostomum, which is not
probable because these genera have isodiametric lam-inal cells.2.28. Dicranites subflagellare Caspary and Klebs
(Hoffeins 1343/3, Kernegger 1994/259, Grabenhorst
The specimens consist of short apical parts of
pleurocarpous mosses with narrow lanceolate leaves.
The leaves are erect patent to appressed in the tip, and
are narrowed from ovate base into a long fine tip.
They are ecostate and entire at margins. Such (appar-
ently easily detachable) apices of stems or branches
apparently serve for vegetative reproduction. A sim-
ilar or identical fragment was described as Dicrantes
subflagellare, although the illustration (Caspary,
1907, Fig. 47) indicates a hypnaceous rather than a
Flagellate branches (with shorter, appressed leaves)
are a common method of vegetative propagation in
mosses. The branches can be fully flagelliform or the
apices of normally developed branches end in flagellae.
At present, the specimens cannot be attributed to a
known species. The branches of Barbella sp. (see
Section 2.17) end in such long flagellae, and it could
be that these flagellae belong (at least in part) to that
2.29. Hypnites lanceolatus sp. nov. (Grabenhorst
The amber includes two fragments of a pleuro-
carpous moss of 3 and 5 mm length, as well as
numerous single, resupinate leaves (Figs. 38 and
39). The leaves are ecostate, have a smooth margin
and possess narrow, elongate laminal cells.
Although quite distinct, this specimen cannot be
attributed to any genus at all. Because of the ecostate
leaves and elongate laminal cells, it belongs to the
Hypnaceae and is accordingly described here in the
form genus Hypnites.
Holotype: Grabenhorst La 25. Etymology: The
species is named after its conspicuously narrowly
lanceolate leaves. Diagnosis: Plantae cum foliis resu-
pinatis, valde angustis, lanceolatis, ecostatis, margin-
ibus integerrimis, cellulis linearibus.
2.30. Aptychella sp. (Kernegger 1992/51)
Kernegger 1992/51 includes fragments of three
different moss species (Figs. 40 and 41). One is part
2.33. Unknown moss (Grabenhorst 11)
The amber contains a rosette of four leaves and
one sporophyte (Fig. 42). The leaves are up to 4
mm long, have a broad ovate base and are con-
tracted into a long acumen (23 times as long as
the basal lamina). They have a percurrent costa,
which fills the apex. The leaves could (especially in
connection with the sporophyte) be perichaetial
leaves because of their clasping bases and long
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 8110198of a stem of a pleurocarpous moss with erect patent
leaves. The leaves are from ovate base finely apiculate
and concave. The laminal cells are elongate oval, a
costa could not be recognized. The leaves at the
branch tip are distinctly smaller and appressed. The
leaves seem to be arranged in three rows.
Another fragment of a pleurocarpous moss is 2 mm
long; the shape of leaves and laminal cells are the
same as described before; however, the plant has a
different appearance. It may possibly be identical with
the first species.
The tip of the stem in the first specimen (Fig. 41) is
conspicuously caudate by appressed leaves, a charac-
teristic of the genus Aptychella (Sematophyllaceae),
see also Noguchi (19871994, fig. 477). For that
reason, these specimens are tentatively placed in this
genus, which comprises 11 species in the Neotropics
and (mainly) SE Asia (Brotherus, 1924).
Besides, there is a short apical part of a pleuro-
carpous moss with two short branches and narrow
lanceolate leaves, which cannot be identified.
2.31. Pleurocarpous moss (Grohn 617)
A 2-mm-long apical part of a presumably pleuro-
carpous moss. The leaves are from lanceolate base
tapering into a very long acumen, which is from 1/3 to
2/3 as long as the lamina and in part coarsely dentate.
A costa seems to lack; laminal cells are hardly visible,
2.32. Hypnites complanatus sp. nov. (Hoffeins
A 1.5-mm-long apical part of a moss stem with
12 leaves, which ends in a flagelliform apex. The
leaves are lanceolate, without costa and with pros-
enchymatous cells. They are distantly arranged and
conspicuously complanate. Such mosses were more
often found in the Baltic amber (Frahm 1999a: 235,
2000a: 38, 2000b: 126), but remained undeter-
mined. For an easier identification, these plants
shall be described here in a form genus.
Holotype: Hoffeins 1417/3. Etymology: The spe-
cies is named after its conspicuously complanate
leaves. Diagnosis: Plantae foliis lanceolatis, valde
distantibus, complanatis, ecostatis, cellulis laminali-bus elongatis.acumen and thus not different from the normal
leaves. The sporophyte has a straight short seta
and an open ovatecylindric capsule. A peristome
is not visible. Although the plant is almost com-
plete, I have no suggestion regarding its identity.
2.34. Symphyodon sp. (Grohn 2022)
Grohn 2022 includes six different pieces of a
similar moss with complanatetristichous foliation,
which are 45 mm in length (Fig. 43). The leaves
are ovate with serrate margins. The preservation is
not so good that details of the cell-structure or
costa-structure can be recognized. Because of the
unique shape of the leaves with serrate margins
and the complanate foliation, the specimen is,
however, identical with Grohn 641 and perhaps
also Grohn 611 (Frahm, 1999a). Because of the
unique combination of characters (ovate, compla-
nate serrate leaves) and details of the lamina (elon-
gate rhombic cells without costa), Grohn 641 was
placed into the genus Symphyodon. The genus com-
prises eight species (Brotherus, 1924), which occur
today in SE Asia from India over the Philippines to
Symphyodon is the only genus within the family
Symphyodontaceae. It can be supposed that this
Fig. 37. Muscites serratus (Grohn 615).
Fig. 38. Hypnites lanceolatus sp. nov. (Grabenhorst La 25).
Fig. 39. Hypnites lanceolatus sp. nov. (Grabenhorst La 25).
Fig. 40. Aptychella sp. (Kernegger 1992/51).
Fig. 41. Aptychella sp. (Kernegger 1992/51b).Fig. 42. Unknown moss (Grabenhorst 11). Scale = 1 mm.
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101 99
3.1. Size of plants
J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81101100As already mentioned in previous publications,
all specimens of extant bryophyte species embeddedfamily was richer in genera and species before the
Tertiary, and that the extant eight species show the
decrease of diversity within the family by extinction.
Fig. 43. Symphyodon sp. (Grohn 2022).in amber are distinctly smaller than those of today.
This concerns also the specimens described here.
The reduction rate of the size is about 50%.
Commonly, it is argued that only small plants were
preserved in amber, and that normal plants in
Tertiary should have had the same size as today.
It is, however, remarkable that all mosses studied
so far are smaller today without a single exception,
and it is unlikely that not even one normal-sized
plant was preserved amongst the dozens specimens
studied so far. Furthermore, amongst these speci-
mens are plants with sporophytes, which should be
fully developed plants. And finally, differences in
size were also observed by earlier authors, e.g.,
Monkemeyer (1927). Monkemeyer wrote: Im all-
gemeinen sind sie [the mosses] kraftiger geworden.
So ist z.B. Scorpidium scorpoides julaceum aus
dem Braunkohlenrevier von Bohlen bei Leipzig
nur etwa 1/3 so stark als die heutige Pflanze.3.2. Age of species
It may sound surprising for nonbryologists that
many of the mosses from the Eocene belong to extant
species. However, it has to be considered that bryo-
phytes are a quite conservative group of plants. The
oldest fossil known from the Devonian (Pallavicinites
devonicus [Hubener] Schuster) resembles already ex-
tant genera such as Pallavicinia or Symphyogyna.
Bryophytes from the late Palaeozoic can easily be
attributed to extant orders, those of the Mesozoic to
extant genera. Besides fossil evidences, we have
molecular evidence that bryophytes have a very slow
rate of evolution. There are taxa disjunct in southern
Chile and New Zealand, thus former Gondwana
elements and separated for the past 80 million years,
which have no means of long-distance dispersal.
These taxa have either few differences on a species
level or even have identical base sequences of molec-
ular markers, which usually show differences on a
species level (Blocher and Frahm, 2002).
It also has to be kept in mind that there is quite a
number of extant ferns and flowering plants, mainly
from the Macaronesian Islands or the Mediterranean,
which are known as fossils from the Tertiary, even
back to the Oligocene (subtropical tertiary relicts,
e.g., Culcita macrocarpa, Davallia canariensis,
Hymenophyllum tunbrigense, Ilex canariensis, Oco-
tea foetens, Laurus canariensis amongst others; Mai,
3.3. Phytogeographical aspects
The phytogeographical elements of the mosses
found in the Baltic and Saxon amber are the same
as the phanerogamic flora of that period, which
included Mediterranean, (S)E Asian and subtropical
North to Central American elements. This paper
added more examples of Eocene species that are
still found in Japan, Korea, China or Taiwan
(Gingko effect). The main reason is that bryophytes,
higher plants as well as other organisms, cannot be
seen isolated but formed synusiae, which persisted
more or less unchanged until the present. A major
difference, however, is that the conformity between
the extant and fossil flora of flowering plants
mostly concerns genera and not species because
flowering plants have developed into new species.
In contrast, the conformity between the extant and
fossil moss flora is much higher due to a much
slower rate of evolution.
The amber forest was an oakpine forest. Interest-
ingly, many species of mosses (e.g., Haplocladium
angustifolium, Campylopodiella himalayana) are still
currently found in oakpine forests in central Amer-
ica, southern Europe and (S)E Asia. Apparently, the
mosses persisted in this kind of habitat over millions
of years more or less unchanged, whereas the phaner-
ogams developed into new species.
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A new contribution to the moss flora of Baltic and Saxon amberIntroductionDescriptionsTrachycystis flagellaris (Sullivant and Lesquereux) Lindberg (Grhn 2023, Grabenhorst La 3, Wichard 14 p.pte.)Trachycystis microphylla (Ddozy and Molkenboer) Lindberg (Hoffeins 1161.3, Wichard 14 p.pte.)Hypnodontopsis conferta (Goeppert and Berendt) J.-P. Frahm (Grhn 616, Hoffeins 1417/1, 2, Grabenhorst La 23, 24, Von Holt 2)Hypnodontopsis fossilis J.-P. Frahm (Hoffeins 1417/2, Grabenhorst La 7, 14)Hypnodontopsis pilifer J.-P. Frahm sp. nov. (Witsch 1)Campylopodiella sp. (Grhn 2028, Grabenhorst La 13, 15, 19, 21, 22)Rhizogonium sp. (Hoffeins 1416)Atrichum cf. rhystophyllum (C. Mller) Paris (Grhn 2041)Haplocladium angustifolium (Hampe and C. Mller) Brotherus (Grhn 2039, Damzen 120, Von Holt 3)Hypnum palaeocircinale sp. nov. (Hoffeins 1417b, Kernegger 1992/27, Grhn 2034)Campylopus sp. (Grabenhorst La 7)Eurohypnum revolutum sp. nov. (Grabenhorst La 10)Tristichella glabrescens Iwats., Sematophyllaceae (Grabenhorst La 12)Sematophyllaceae (Grabenhorst La 12)Ctenidium capillifolium (Mitt.) Broth. (von Holt 4)Campylium cf. squarrosulum (Besch. and Card.) Kanda (von Holt 6)Barbella sp. (von Holt 1, 5)Brotherella sp. (Grhn 619)Brachythecium sp. (Grabenhorst La 18)Bartramia sp. (Grabenhorst La 18)Muscites pilifer J.-P. Frahm (Grabenhorst La 27)Hypnaceous moss (Grabenhorst La 20)Drepanocladus sp. (Kernegger 1998/117)Hypnaceae (Hoffeins 1343/1)Echinodium sp. (Kutscher 15, Witsch 6)Plagiotheciaceae (Grhn 618, 2033)Muscites serratus Goepp. and Berendt (Grabenhorst La 26, Grhn 615)Dicranites subflagellare Caspary and Klebs (Hoffeins 1343/3, Kernegger 1994/259, Grabenhorst La 1)Hypnites lanceolatus sp. nov. (Grabenhorst La 25)Aptychella sp. (Kernegger 1992/51)Pleurocarpous moss (Grhn 617)Hypnites complanatus sp. nov. (Hoffeins 1417/3)Unknown moss (Grabenhorst 11)Symphyodon sp. (Grhn 2022)
DiscussionSize of plantsAge of speciesPhytogeographical aspects