Transcript
Page 1: New and Interesting Mosses from Baltic and Saxon Amber

Oikos Editorial Office

New and Interesting Mosses from Baltic and Saxon AmberAuthor(s): Jan-Peter FrahmSource: Lindbergia, Vol. 25, No. 1 (2000), pp. 33-39Published by: Oikos Editorial OfficeStable URL: http://www.jstor.org/stable/20150031 .

Accessed: 15/06/2014 12:18

Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at .http://www.jstor.org/page/info/about/policies/terms.jsp

.JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide range ofcontent in a trusted digital archive. We use information technology and tools to increase productivity and facilitate new formsof scholarship. For more information about JSTOR, please contact [email protected].

.

Oikos Editorial Office is collaborating with JSTOR to digitize, preserve and extend access to Lindbergia.

http://www.jstor.org

This content downloaded from 188.72.126.41 on Sun, 15 Jun 2014 12:18:00 PMAll use subject to JSTOR Terms and Conditions

Page 2: New and Interesting Mosses from Baltic and Saxon Amber

LINDBERGIA 25: 33-39. Lund 2000

New and interesting mosses from Baltic and Saxon amber

Jan-Peter Frahm

Frahm, J.-P. 2000. New and interesting mosses from Baltic and Saxon amber . -

Lindbergia 25: 33-39.

Eight mosses are recorded from Baltic and Saxon amber. Hypnodontopsisfossilis J.-P. Frahm is described as new. The extant species Trachycystis microphylla

(Dozy & Molk.) Lindb. is recorded for the first time as a fossil. Haplocladium

angustifolium (Hampe & C. M?ll.) Broth, is recorded once again in amber.

Additional specimens could be attributed to Dicranites casparyi R. Klebs, oth

ers could not be identified to genus.

J.-P. Frahm, Botanical Inst., Rheinische Friedrich-Wilhelms Universit?t,

Meckenheimer Allee 170, D-53115 Bonn, Germany.

Fossils in amber - in contrast to fossil plants and ani mals in other deposits

- are usually perfectly preserved and are therefore important sources for the reconstruc

tion of fossil floras and for the determination of ages of genera and species. The percentage of plant fossils in amber is, however, very poor: 99% of all fossils from Baltic amber consist of arthropods (Weitschat and Wichard 1998), less than 1% are phytogenic, and

bryophyte fossils account for an extremely small part of the plant fossils.

So far, eleven moss species from form genera, seven

extant species, nine apparently extinct species and numerous unidentified specimens are known from Baltic amber. In addition, two species from form gen era, three extant species and two species from extant

genera are known from Saxon amber (Frahm 1999a). The determination of fossil mosses from Baltic and

Saxon amber allows a more detailed knowledge of the flora of the 'amber forests', which were situated in the Eocene (38-54 MA BP, in contrast to previous estimates of a younger age) in the area of present Fennoscandia. In upper Eocene times the resin of co niferous trees was transported by rivers to secondary

deposits in an ocean south of these forests. According to the latest geological findings, the Saxon amber

originated by subsequent transport of the Baltic am

ber to Tertiary deposits in the Miocene. Plant fossils found in Baltic and Saxon amber thus belong to the same flora (Weitschat and Wichard 1998).

By courtesy of Mr Dirk Teuber (Bielefeld), Manfred Kutscher (Sassnitz) and Mrs and Mr Hoffeins (Ham

burg) I received some fossil mosses from Baltic and Saxon amber for determination. These contain sev

eral new and interesting records, which are compiled here. Unfortunately, only a small part of the fossil

mosses can be identified because characters necessary for determination are not visible, not preserved or not

present. Such unidentified specimens are, however, also listed to give an impression of the diversity of the moss flora of the Tertiary.

The specimens are kept in the collections of the collectors. B = Baltic amber, S = Saxon amber.

Trachycystis microphylla (Dozy & Molk.) Lindb.

(B: Teuber 1606, Fig. 1)

The piece of amber contains three mosses, two of which belong to the genus Trachycystis. One of the

Trachycystis specimens consists of the upper part of a Accepted 19 November 1999 ? LINDBERGIA 2000

LINDBERGIA 25:1 (2000) 33

This content downloaded from 188.72.126.41 on Sun, 15 Jun 2014 12:18:00 PMAll use subject to JSTOR Terms and Conditions

Page 3: New and Interesting Mosses from Baltic and Saxon Amber

plant with 20 erecto-patent leaves. The other also con

sists of an upper part of a plant, but with 13, more

patent leaves. The leaves are only 1.5-2 mm long and resemble those of the genus Mnium but have papil lose laminai cells.

The genus Trachycystis is known from the Tertiary of Europe from three species: Trachycystis antiquo rum (Cardot & Dixon) Kabiersch from the Pliocene of Reu ver (Reid and Reid 1915), Trachycystis szaferi Szafran from the Miocene of Poland (Szafran 1958) and Trachycystis flagellaris [Sull. & Lesq.] Lindb. from Baltic amber (Frahm 1994). Trachycystis antiq uorum and T. szaferi are regarded as extinct species, T. flagellaris occurs today in Eastern Asia. Another

species of Trachycystis, which is not identical to any of these three species, has been reported from Baltic amber (Frahm 1999a).

A key was given for two fossil and two of the three extant species of Trachycystis by Szafran (1958).

Keys, descriptions and illustrations for the three ex tant species of Trachycystis (T. flagellaris, T. micro

phylla [Dozy & Molk.] Lindb. and T. ussuriensis

[Maack & Regel] T. Kop.) were given by Koponen (1981) and Noguchi (1989). According to these au

thors, Trachycystis microphylla can be distinguished by unbordered leaves. The leaves are furthermore more

sharply pointed than those of the other species of

Trachycystis, which have blunt or Ungulate leaf api ces. The leaves are considerably shorter (1.5-2 mm) than those of extant specimens of the same species (3

mm), a phenomenon which has also been observed in other fossils of recent moss species. This may be at tributed to the fact that only small plants, which were

more easily blown into the resin than normally devel

oped plants, became embedded.

Conspicuously, the extant species of Trachycystis are growing on soil and rotten wood, but not epiphyti cally. This indicates that not only epiphytic species but also epigaeic and epilithic mosses were embed ded in resin. Fragments of these may either have been blown into the resin or covered by drops of resin.

Pleurocarpous moss (B: Teuber 1605, Fig. 2) The specimen consists of a single plant, which is bent over 90? in the upper third, and has 10 short branches. The leaves are narrowly lanceolate, finely pointed, and

apparently ecostate. They are distinctly patent, the leaf

apices are incurved. The specimen is apparently conspecific with Gr?hn 869, Wichard 326 and Teuber 1565 (Frahm 1999a).

Pleurocarpous moss (B: Teuber 1606a, Fig. 3, S: Hoffeins 962/2, Fig. 4, S: Kutscher 2b, Fig. 7-8) The piece of amber with two plants of Trachycystis microphylla (Teuber 1606) contains an apex of a stem or branch of a pleurocarpous moss with lanceolate leaves, which are ecostate, very narrow and long, 650

urn long and have prosenchymatous laminai cells. The basal leaves are widely patent, the upper leaves are convolute. Part of the plant is covered by air bubbles.

Kutscher 2b consists of two fragments of a pleuro carpous moss. The leaves are

conspicuously distant

but crowded at the apex, narrowly lanceolate, finely pointed and ecostate. They are distinctly concave and have long, narrow laminai cells.

Hoff eins 962/2 consists of two 7- and 12-mm-long fragments of a pleurocarpous moss, which is densely covered by air bubbles. A short stem 4 mm long with 15 leaves can be studied in detail. It is compl?nate

with narrowly lanceolate, ecostate leaves and prosen chymatous cells. All plants are very similar and seem to belong to the same species. They resemble Broth erella sp. from Baltic amber (cf. Frahm 1996a), which

has, however, sharply dentate leaf apices.

Dicranites casparyi R. Klebs (B: Hoff eins 874, Fig. 5, S: Kutscher 2c, 3b) Hoffeins 874 consists of an almost complete plant of 4 mm length with a 6-mm-long seta, however without

sporangium. The basal leaves are 1.6 mm long, the

upper ones 2 mm. A small branch with leaves only 1 mm long originates at the base of the stem. The leaves are narrowly lanceolate, the costa ends in the leaf apex. The leaf margins are slightly serrate near the leaf apex. The laminai cells are round to quadrate and arranged in distinct rows. The lamina is only 5-10 cell rows

wide on both sides of the costa. Kutscher 2c, 3b consists of the upper part of plants

with a tuft of narrowly lanceolate leaves. The species gives the impression of a pottiaceous

plant, but the species of Pottiaceae usually have basal laminai cells of different shape and the upper laminai cells are usually papillose or mamillose, which is not the case in these specimens. Therefore they may be better regarded as Dicranaceae. Conspicuous are the

smooth, quadrate laminai cells in a few rows on each side of the costa.

The specimens strongly resemble the illustrations of the four syntypes of Dicranites casparyi (cf. Figs

-> Fig. 1. Trachycystis microphylla (Dozy & Molk.) Lindb. (Teuber 1606). Fig. 2. Pleurocarpous moss (Teuber 1605).

Fig. 3. Pleurocarpous moss (Teuber 1606a). Fig. 4. Pleuro

carpous moss (Hoffeins 962/2).

34 LINDBERGIA 25:1 (2000)

This content downloaded from 188.72.126.41 on Sun, 15 Jun 2014 12:18:00 PMAll use subject to JSTOR Terms and Conditions

Page 4: New and Interesting Mosses from Baltic and Saxon Amber

LINDBERGIA 25:1 (2000) 35

This content downloaded from 188.72.126.41 on Sun, 15 Jun 2014 12:18:00 PMAll use subject to JSTOR Terms and Conditions

Page 5: New and Interesting Mosses from Baltic and Saxon Amber

42-45 in Caspary [1907]), which might be identical with Muscites confertus Goepp. & Ber. (Caspary 1907) as well as the specimens Kutscher 4,11, 12 and 14

(Frahm 1999a). Caspary (1907) denotes this species as the most common one in [Baltic] amber. The iden

tity of our specimens cannot be proved because the

specimens published by Goeppert and Berendt (1845) cannot be located. The name Dicranites casparyi is

given preference because it is more precise than Muscites confertus.

Haplocladium angustifolium (Hampe & C. M?ll.) Broth. (B: Wenzel s.n., S: Kutscher 3a, B:

Hoffeins 962/1 Fig. 6)

Wenzel s.n. consists of the apical part of a densely and regularly branched moss of 21 mm length. The

leaves are narrowly lanceolate and from an ovate base

gradually narrowed into a long and fine point. The

leaf border is narrowly recurved. The costa reaches the apex and is prominent at the back. The laminai cells are round.

Kutscher 3 a consists of a 1 -cm-long piece of a pleu rocarpous moss with five branches of 2-5 mm length,

which get shorter toward the stem tip. Stems and branches are narrowly foliate with lanceolate leaves. The costa reaches the leaf apex, the laminai cells are

round and the leaf border is revolute.

Hoffeins 962/1 contains a 7-mm-long fragment of a

pleurocarpous moss with creeping stem and seven,

quite regularly inserted branches of similar length which point in one direction. The leaves are narrow

and the margin is revolute along the whole leaf. The

leaves are 1 mm long and contracted from an ovate

base into a long tip. The costa vanishes in the leaf

apex. The laminai cells are rounded-quadrate.

All fossils apparently belong to the same species, which has frequently been found in Baltic amber

(Kutscher 15, ?17, Wichard 12, Gr?hn 1558 as

Leskeaceae, cf. Frahm [1999a], Hoff eins 483 as

isobryalean moss, cf. Frahm [1996a]). Some fragments (Teuber 1516 and 1507 [Frahm 1999a]) probably be

long to this species. Conspicuous in all these fossils is

the combination of characters such as lanceolate leaves

with roundish laminai cells, revolute leaf margins, and a costa filling the leaf apex and being prominent at

the back. This combination of characters seems to

match only Haplocladium angustifolium (Thuid

iaceae). This species is widely distributed in South

and Southeast Asia from India to Korea, but in Eu

rope it occurs only in the southern Alps, where it is

regarded as a relict from the Tertiary.

Hypnodontopsis fossilis spec, no v. (S: Kutscher

s.n., Fig. 9)

A single plant of an acrocarpous moss of 2.8 mm

length with sporophyte. The sporophyte is 1 mm long and consists of a 0.5-mm-long seta, which is conspicu

ously twisted, and a 0.5-mm-long cylindrical capsule with eight longitudinal furrows. The peristome can

be seen from the other side of the amber piece, which is cut and polished just above the mouth of the cap sule. It consists of eight teeth, presumably in pairs.

The leaves are up to 1.2 mm long, narrowly lanceo

late, twisted at the apex, with a costa which reaches the apex. The base of the leaf is somewhat widened.

The laminai cells are round in the upper part and rec

tangular at the base.

The sporophyte of this plant has an unusual combi nation of characters such as a short, twisted seta, an

empty capsule with eight furrows and a peristome of

eight paired teeth which are not split. This combina tion of characters fits -

according to my knowledge -

only the genus Hypnodontopsis. It was described more than 40 years ago by Iwatsuki and Noguchi. It is

monotypic with only Hypnodontopsis apiculata Iwats. & Nog. endemic to Japan (Noguchi 1989). The plants of H. apiculata are tiny, with only 0.7-1.2 mm long leaves, which applies also to the fossil plant; the leaves

are, however, Ungulate, whereas they are distinctly narrower in the fossil plant. Leaf-born brood bodies,

typical for H. apiculata, could not be observed in the fossil material. Occasionally, blackish-brown clusters can be seen at the leaf tips of the fossil plants, which could consist of brood bodies, but also of dirt.

It can be concluded from the sporophytic charac ters that the fossil plant is a species o? Hypnodontopsis, but from the gametophytic characters that the fossil is not identical with the extant H. apiculata. It can there fore be described as a new but apparently extinct spe cies:

Hypnodontopsis fossilis J.-P. Frahm sp. nov.

Planta 2,8 mm alta. Folia 1,0-1,2 mm longa, anguste

lanceolata, basi ovata, costa percurrente, apicibus

tortilibus. Cellulae lam?nales rotundae, basi rectan

gulares. Seta 0,5 mm longa, tortilis. Theca 0,5 mm

longa, striata. Peristomium cum 8 dentibus.

Holotype in the collection of M. Kutscher s. n., Sassnitz.

->

Fig. 5. Dicranites casparyi R. Klebs (Hoffeins 874). Fig. 6. Haplocladium angustifolium (Hampe & C. M?ll.) Broth.

(Hoffeins 962/1). Figs 7-8. Pleurocarpous moss (Kutscher

2b).

3 6 LINDBERGIA 25:1 (2000)

This content downloaded from 188.72.126.41 on Sun, 15 Jun 2014 12:18:00 PMAll use subject to JSTOR Terms and Conditions

Page 6: New and Interesting Mosses from Baltic and Saxon Amber

LINDBERGIA 25:1 (2000) 37

This content downloaded from 188.72.126.41 on Sun, 15 Jun 2014 12:18:00 PMAll use subject to JSTOR Terms and Conditions

Page 7: New and Interesting Mosses from Baltic and Saxon Amber

Fig. 9. Hypnodontopsis fossilis J.-P. Frahm based on a pho

tograph (Type).

Hypnaceae (S: Kutscher s. n.) An 8-mm-long part of a pleurocarpous moss with two small branches. The leaves are narrowly lanceolate,

longly and finely pointed, ecostate and have a pros enchymatous areolation.

The ecostate leaves and prosenchymatous laminai cells suggest that the plant belongs to the Hypnaceae. Sematophyllaceae, which resemble the Hypnaceae in

many respects, have serrate or shorter leaf apices. The

leaves are only slightly curved in the apex. It can there fore not be decided whether it is an orthophyllous form of a species of Hypnum or a representative of another

genus. A Hypnum sp. has been recorded twice from Baltic amber (Frahm 1996b, 1999a).

Pleurocarpous moss (S: Kutscher 3c) The specimen consists of several stems of 5-8 mm

length. The stems are densely and almost vermicularly foliate, the leaves are

suddenly contracted from a

broadly oval base. Neither laminai cells nor the pres ence or absence of a costa is visible. Therefore the

specimen must remain unidentified.

Brotherella sp. (S: Kutscher 2a) An apical part of the stem of a pleurocarpous moss with 24 leaves. The leaves are narrowly lanceolate, ecostate, and have a prosenchymatous areolation. The

leaf apices are frequently twisted. This diagnosis may apply to many pleurocarpous mosses; however, the leaf apices are coarsely serrate. The specimen is ap

parently identical with Hoffeins 187 and 357, which were named as Brotherella sp. because of the serrate

leaf apices (Frahm 1996b). It strongly resembles the

descriptions and illustrations of Muscites serratus and Muscites tortifolius. The type of the former consists

of a similar apical part of a stem.

Besides the numerous unidentified specimens or speci mens which can only be attributed to families, the fol

lowing species are so far known from Baltic and Saxon amber.

Form genera

Dicranites casparyi R. Klebs (Caspary 1907, Frahm

1999a, this paper) Dicranites grollei J.-P. Frahm (Frahm 1999b) Dicranites obtusifolius Casp. & R. Klebs (Caspary

1907) Dicranites subflagellare (Goepp. & Middend.) Char.

(Caspary 1907, Frahm 1999a) Muscites convolutus M?gdefrau (M?gdefrau 1957) Muscites apiculatus Goepp. & Berendt (Goeppert &

Berendt 1845) Muscites confertus Goepp. & Berendt (Goeppert &

Berendt 1845) Muscites dubius Goepp. & Berendt (Goeppert &

Berendt 1845, Goeppert 1853) Muscites elegans Goepp. & Berendt (Goeppert 1853) Muscites hirsutissimus Goepp. & Berendt (Goeppert

& Berendt 1845, Goeppert 1853) Muscites pilifer J.-P. Frahm (Frahm 1999b) Muscites serratus Goepp. & Berendt (Goeppert &

Berendt 1845, Goeppert 1853, Caspary 1907) Muscites tortifolius Casp. & R. Klebs (Caspary 1907,

Frahm 1999a)

Extant species Dicranum fuscescens Sm. (Goeppert 1853) Fabroniacf. ciliaris (Brid.) Brid. (Frahm 1999a) Haplocladium angustifolium (Hampe & C. M?ll.)

Broth. (Frahm 1999a, this paper)

38 LINDBERGIA 25:1 (2000)

This content downloaded from 188.72.126.41 on Sun, 15 Jun 2014 12:18:00 PMAll use subject to JSTOR Terms and Conditions

Page 8: New and Interesting Mosses from Baltic and Saxon Amber

Hymenostomum microstomum Hedw. [Weissia brachy

carpa (Nees & Hornsch.) Jur.] (Goeppert 1853) Hypnum squarrosum Hedw. [Rhytidiadelphus squar

rosus (Hedw.) Warnst.] (Goeppert 1853) Phascum cuspidatum Hedw. (Goeppert 1853)

Trachycystis flagellaris (Sull. & Lesq.) Lindb.

[Muscites hauchecornei Casp. & R. Klebs] (Caspary 1907, Frahm 1994)

Trachycystis microphylla (Dozy & Molk.) Lindb. (this

paper) Cf. Merrilliobryumfabronioides Broth. (Frahm 1996b)

Extinct species Dicranum simplex Goepp. & Menge nom. nud.

(Dicranella simplex Broth, nom. nud.) (Goeppert 1853)

Dicranum subflagellare Goepp. & Menge nom. nud.

(Goeppert 1853) Dicranum subpellucidum Goepp. & Menge nom. nud.

[Dichodontium subpellucidum] (Goeppert 1853) Dicranum subscoparium Goepp. & Menge nom. nud.

(Goeppert 1853) Grimmia subelongata Goepp. & Menge nom. nud.

(Goeppert 1853) Hypnodontopsis fossilis J.-P. Frahm (this paper) Polytrichum subseptentrionale Goepp. & Menge nom.

nud. (Goeppert 1853) Polytrichum subundulatum Goepp. & Menge nom.

nud. (lAtrichum undulatum (Hedw.) P. Beau v.)

(Goeppert 1853) Polytrichum suburnigerum Goepp. & Menge nom.

nud. (Pogonatum suburnigerum Dix. nom. nud.)

(Goeppert 1853) Trichostomum substrictum Goepp. & Menge nom.

nud. (Goeppert 1853) The identifications of Goeppert (1853) seem in part

to be dubious, but can no longer be verified since the Goeppert collections are lost.

Extant genera

Barbella sp. (Frahm 1996b, 1999a) Bescherellea sp. (Frahm 1999a) Brotherella sp. (Frahm 1996a, this paper) Campylopodiella sp. (Frahm 1996a, 1999a)

Campylopus sp. (Frahm 1996b) Fabronia sp. (Frahm 1999a)

Hypnum sp. (Frahm 1996b, 1999a)

Mastopoma sp. (Frahm 1999a)

Symphyodon sp. (Frahm 1999a)

Trachycystis sp. (Frahm 1999a)

With the records published in this paper, the number of mosses known from Baltic and Saxon amber is

raised to 13 species in form genera, nine extant spe cies, ten extinct species, and ten unidentified species in extant genera.

Part of the taxa are still known from Europe, others are extinct in Europe but are still known from East or

Southeast Asia.

References

Caspary, R. 1907. Die Flora des Bernsteins. - Abhandl.

Preu?. Geolog. Landesanstalt N.F. 4. Berlin.

Frahm, J.-P. 1994. Die Identit?t von Muscites hauchecornei

Caspary & Klebs (Musci) aus Baltischem Bernstein. -

Nova Hedwigia 58: 239-243. - 1996a. Mosses newly recorded from Saxonian amber. -

Nova Hedwigia 63: 525-527. - 1996b. Laubmoose aus Baltischem Bernstein.

-

Palaeontogr. Abt. B 241: 127-135. - 1999a. Die Laubmoosflora des Baltischen und Bitter

felder Bernsteins. - Mitt. Geolog.-Pal?ontol. Staats

institut 83: 219-238. -

1999b. Neue bemerkenswerte Laubmoosfunde aus B al

tischem Bernstein. - Haussknechtia Beih. 9: 129-132.

Goeppert, H.R. 1853. ?ber die Bernsteinflora. - Monats

berichte der K?niglich preu?ischen Akademie der Wis senschaften zu Berlin 1853: 450-477.

- and Berendt G.C. 1845. Der Bernstein und die in ihm

befindlichen Pflanzenreste der Vorwelt. - Berlin.

Koponen, T. 1981. A synopsis of Mniaceae (Bryophyta) VI. Southeast Asian taxa. - Acta Bot. Fenn. 117: 1-34.

M?gdefrau, K. 1957. Flechten und Moose in Baltischem

Bernstein. - Ber. Dtsch. Bot. Ges. 70: 433-435.

Noguchi, A. 1989. Illustrated mossflora of Japan. Part 3. -

Hattori Botanical Laboratory, Nichinan.

Reid, C. and Reid, E.M. 1915. The Pliocene floras of the

Dutch-Prussian border. - M. Nijhoff, Gravenhage.

Szafran, B. 1958. Trachycystis zaferi, a new species of moss

from the Miocene of Poland. - Acta Soc. Bot. Pol. 20:

247-250.

Weitschat, W and Wichard, W 1998. Atlas der Pflanzen

und Tiere im Baltischen Bernstein. - Dr. Pfeil, M?nchen.

LINDBERGIA 25:1 (2000) 39

This content downloaded from 188.72.126.41 on Sun, 15 Jun 2014 12:18:00 PMAll use subject to JSTOR Terms and Conditions