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Review of Palaeobotany and Palynology 129 (2004) 81–101

A new contribution to the moss flora of Baltic and Saxon amber

J.-P. Frahm*

Botanisches Institut der Rheinischen, Friedrich-Wilhelm-Universitat, Meckenheimer Allee 170, D 53115 Bonn, Germany

Received 3 May 2002; accepted 28 November 2003

Abstract

So far, the largest collection of mosses from the Baltic and Saxon amber (Eocene, 37–57 million years B.P.) consisting of 55

specimens 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 amber

1. Introduction grad, also in Saxony (Bitterfeld) and scattered in

Baltic and Saxon amber is a valuable source for

fossils from many groups of organisms. It originated

in oak–pine 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-

tion 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.

doi:10.1016/j.revpalbo.2003.11.004

* Fax: +49-228-733-120.

E-mail address: [email protected] (J.-P. Frahm).

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 first

hypothesis, 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-

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).

Fig. 6. Hypnodontopsis conferta (Grohn 2042).

J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81–10182

ited 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 author’s 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 the

collectors.

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

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. Descriptions

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 similar

and consists of 13 leaves that are somewhat com-

planate. The laminal cells are rounded. All these

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

J.-P. Frahm / Review of Palaeobotany and Palynology 129 (2004) 81–101 83


fossil 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

leaf-border.

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,

1987–1994), 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 still—as all

extant species of this genus—in East Asia (Koponen,

1981).

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 5–7 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

(Koponen, 1981).

By a unique chance, two plants of different species

of Trachycystis were imbedded in Wichard 14. The

first plant is about 5 mm long and has 20 distant

leaves, 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

Caspary (1907).

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 5–6 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


few 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,

1928.

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

(2001b).

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 1.5–2 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. The

peristomes are no longer preserved. La 24 includes a

dozen 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. 29–31). Caspary

(1907, figs. 44–45) 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

Muscites confertus.

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. 44–45,

which belong to Hypnodontopsis conferta) as well as

Dicranites obtusifolius (Caspary, 1907, fig. 46). How-

ever, the few longitudinal rows typical for the fossil

Fig. 7. Hypnodontopsis conferta (Grabenhorst La 23), several fertile

plants.

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.


specimens 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-

known) species.

Hypnodontopsis fossilis was so far only known

from the type specimen (Frahm, 2000a).

2.5. Hypnodontopsis pilifer J.-P. Frahm sp. nov.

(Witsch 1)

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

(1–2 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

distinct rows.

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

be conspecific.

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

Hypnodontopsis.

Holotype: coll. Witsch #1. Etymology: The species

is named after its costa ending in a long hairpoint.

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 5–6

ordines.

The new species is easily distinguished from all

other species of the genus by its leaves ending in a

long hairpoint.

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

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. 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).


separate 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

2–3 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

capsule shape.

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 this

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

along the stem, longer teeth along the leaf margins

and smooth laminal cells.

The genus is mainly distributed in SE Asia and

Australasia.

2.8. Atrichum cf. rhystophyllum (C. Muller) Paris

(Grohn 2041)

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. 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.


pared 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

(Nyholm, 1971).

2.9. Haplocladium angustifolium (Hampe and C.

Muller) Brotherus (Grohn 2039, Damzen 120, Von

Holt 3)

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 Baltic

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

be 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-

ty—the 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. 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).


of 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

genus Campylopus.

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

La 10)

The preserved plant is 1 cm in length with creeping

stem and several erect, 3–4 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, 1987–1994), 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. Cellulae

laminae elongatae.

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

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, 1987–1994), 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. 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).


narrow 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 ovate–cordate

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 (1987–1994),

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 1.5–2 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 is

not visible. However, some of the branches end in

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. 7–8;

1999a: 228).

2.18. Brotherella sp. (Grohn 619)

Grohn 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



much 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

La 27)

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 1/4–1/3 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 finely

apiculate.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

costa.

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 a

longitudinally split and give the impression of a

prosenchymatous areolation.

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

La 1)

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

dicranaceous moss.

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

species.

2.29. Hypnites lanceolatus sp. nov. (Grabenhorst

La 25)

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

nd Palynology 129 (2004) 81–101 97

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.


of 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 (1987–1994, 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,

presumably elongate.

2.32. Hypnites complanatus sp. nov. (Hoffeins

1417/3)

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.

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 (2–3 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

acumen and thus not different from the normal

leaves. The sporophyte has a straight short seta

and an open ovate–cylindric 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 complanate–tristichous foliation,

which are 4–5 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

Japan.

Symphyodon is the only genus within the family

Symphyodontaceae. It can be supposed that this


Fig. 43. Symphyodon sp. (Grohn 2022).

J.-P. Frahm / Review of Palaeobotany a100

family 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.

3. Discussion

3.1. Size of plants

As already mentioned in previous publications,

all specimens of extant bryophyte species embedded

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,

1995).

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.

nd Palynology 129 (2004) 81–101


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 oak–pine forest. Interest-

ingly, many species of mosses (e.g., Haplocladium

angustifolium, Campylopodiella himalayana) are still

currently found in oak–pine 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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