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: firstname.lastname@example.org (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) 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