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

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    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

    J.-P.

    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.

    doi:10.1016/j.revpalbo.2003.11.004

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

    E-mail address: frahm@uni-bonn.de (J.-P. Frahm).ora of Baltic and Saxon amber

    hm*

    sitat, Meckenheimer Allee 170, D 53115 Bonn, Germany

    www.elsevier.com/locate/revpalbo

    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. 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 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

    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,

    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,

    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 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

    (Koponen, 1981).

    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

    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 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,

    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 1.52 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

    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. 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

    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

    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-

    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

    (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

    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 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

    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.

  • 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

    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 thisalong 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. 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

    (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 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

    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, 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;

    1999a: 228).

    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

    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 1/41/3 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

    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 and Palynology 129 (2004) 81101 97longitudinally 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

  • 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,

    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.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

    Japan.

    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.

    3. Discussion

    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,

    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.

  • 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.

    stein. Cryptogamie 21, 121132.

    Frahm, J.-P., 2001a. Neue laubmoosfunde aus sachsischem und

    baltischem Bernstein. Nova Hedwig. 72, 271281.

    Frahm, J.-P., 2001b. Hypnodontopsis confertus from Baltic amber.

    Trop. Bryol. 20, 7982.

    Goeppert, H.R., 1853. Uber die Bernsteinflora. Monatsberichte der

    Koniglich Preuischen. Akademie der Wissenschaften, Berlin,

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    Goeppert, H.R., Berendt, G.C., 1845. Der Bernstein und die in Ihm

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    Jovet-Ast, S., 1967. Bryophyta. In: Boureau, E. (Ed.), Traite de

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    Koponen, T., 1981. A synopsis of Mniaceae (Bryophyta) VI South-

    east Asian taxa. Acta Bot. Fenn. 117, 134.

    Kutscher, M., 1999. Bernstein. Verlin der Freunde und Forderer des

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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

    References