1. Introduction four extant species exist and they are restricted to a
umbrella-shaped indusia are less known amongpalaeontologists and neontologists, that specimen wasassumed to be a cap of a basidiocarpium similar to the
Review of Palaeobotany and PalynologMatoniaceae are already known from the Triassic andthey were widespread especially during the Jurassic andEarly Cretaceous (Tidwell and Ash, 1994; Skog, 2001).The absence of Cenozoic macrofossils and spores,however, is strongly emphasized in concluding treatises(e.g. Collinson, 2001). The only previous finding fromthe Palaeogene is a stem fossil from a conglomerate ofTasmania which has been reworked from much olderMesozoic sediments (Tidwell and Skog, 1992). Only
strongly equatorial relict area in Southeast Asia.Recently, several inclusions were found in Baltic and
Bitterfeld amber which provide fossil evidence ofCenozoic Matoniaceae. Cooperation with several ambercollectors allowed the examination of inclusions of alto-gether six indusia, 11 sporangia and numerous spores inBaltic amber, and three indusia in Bitterfeld amber.
The first amber inclusion of a matoniaceousindusium was found during palaeomycological investi-gations of Bitterfeld amber. Since the characteristic largeshow that the Matoniaceae became restricted to the equatorial Southeast Asia during the Neogene and not during the LateCretaceous or Palaeogene as indicated by the previous fossil record. The type specimen of Matonia striata (Drfelt et Striebich)Schmidt et Drfelt, comb. nov. was previously described as a fungus, Palaeocybe striata [Drfelt, H., Striebich, B., 2000.Palaeocybe striata, ein neuer fossiler Pilz in Bernstein des Tertir. Zeitschrift fr Mykologie 66, 27-34.], based on themorphological similarity of the fossil indusium with a cap of a basidiocarpium. The indusia show close similarities to those of theextant genera Matonia and Phanerosorus. 2006 Elsevier B.V. All rights reserved.
Keywords: Matoniaceae; Matonia; Palaeocybe; amber; Tertiarywhich was previously known from Mesozoic macrofossils andMatoniaceae occurred in European amber forests in the Eocene and at the OligoceneMiocene boundary. This is now revealedby resin-preserved indusia, sporangia and spores. These inclusions are the first Cenozoic fossil records of this archaic fern family,
spores and few relict species. The findings from non-tropical forestsEvidence of Cenozoic Matoniace
Alexander R. Schmida Museum fr Naturkunde der Humboldt-Universitt zu Berlin, I
b Martin-Luther-Universitt Halle, Institut fr Geobotanik und
Received 11 November 2005; received in rAvailable onlineCorresponding author. Tel.: +49 30 2093 8945.E-mail address: email@example.com
0034-6667/$ - see front matter 2006 Elsevier B.V. All rights reserved.doi:10.1016/j.revpalbo.2006.07.009from Baltic and Bitterfeld amber
, Heinrich Drfelt b
t fr Palontologie, Invalidenstr. 43, D-10115 Berlin, Germanynischer Garten, Neuwerk 21, D-06099 Halle/Saale, Germany
form 4 July 2006; accepted 10 July 2006eptember 2006
y 144 (2007) 145156www.elsevier.com/locate/revpalboextant genus Coprinus and consequently described as afungus (Drfelt and Striebich, 2000). Afterwards, a
further inclusion was found in the Hoffeins collectionand Gnter Krumbiegel (personal communication,2001) also believed a similar inclusion of his collectionto be a cap of a basidiomycete. Indeed, some featuressuch as the striation of the cap by small decurrent ribsand the wide tubes of the broken stalk may suggest abasidiocarpium. For the first time, Volker Arnold(personal communication, 2001) found indications forthe true nature of the inclusions. This was especiallybecause some indusia of his collection have syninclu-sions of sporangia and spores. Johanna H.A. vanKonijnenburg-van Cittert confirmed Arnold's identifi-cation. Basidia are not visible at the cross sections oftwo inclusions of Arnold's collection which weredamaged as the amber was ground and polished. Thisand especially the comparison with indusia of extantspecies provided evidence that the features of the fossilscorrespond to indusia of representatives of Matoniaceae.
2. Materials and methods
The following pieces of amber contain the micro-inclusions of Matoniaceae investigated:Collection Bernhard Striebich (Buxtehude, Germany),no. 21/8/56: piece of Bitterfeld amber; one indusium, i.e.the type specimen of Palaeocybe striata (Drfelt andStriebich, 2000); syninclusions are remnants of a Thujatwig, diverse small leaves, stellate hairs which originatemost probably from Fagaceae, a pseudoscorpion, detritusand several hairs from macrophytes.CollectionChristel andHansWerner Hoffeins (Hamburg,Germany), no. 809: piece of Bitterfeld amber; oneindusium.Collection Gnter Krumbiegel (Halle/Saale, Germany),no. KRU 2: piece of Bitterfeld amber; one indusium.Collection Volker Arnold (Heide, Germany), no. 1398:piece of Baltic amber; one immature closed indusium.
Plate I. Holotype of Matonia striata (collection Bernhard Striebich, no. 21/8/56). (see plate on page 147)
1. Piece of amber with the indusium.2. Detail of the indusium showing the almost hemispheric shape of the umbrella.3. Lower side of the umbrella with visible striation.45. Indusium in oblique lateral view with the stalk (4) and marginal region of the upper side (5) in focus.6. Detail of the margin with branching and anastomosing ribs.Scale bars represent 1 mm in 12, 500 m in 35 and 100 m in 6.
Plate II. Paratype of Matonia striata (collection Christel and Hans Werner Hoffeins, no. 809). (see plate on page 148)
dian ply forof th
d 6 w
146 A.R. Schmidt, H. Drfelt / Review of Palaeobotany and Palynology 144 (2007) 1451561. Upper side of the indusium.2. Lower side of the indusium. A bubble of air, detritus partic3. Polygonal apical cells at the top of the upper side of the um46. Details of the involute margin in upper view (4) and lowerScale bars represent 500 m in 12 and 100 m in 36.
Plate III. Specimens of Matonia striata from the collections Gnter Kr
1. Indusium of the collection Gnter Krumbiegel (no. KRU 22. Lower side of the closed immature sorus of the collection
involute margin and the centrally located break of the stalk39. Indusia, sporangia and spores of the collection Volker Arno3. Entire indusium (paratype).4. Reconstruction of an indusium which was damaged at its me
to the front. The cut surface shows that the umbrella is largeof the stalk; (1) cut surface in the median part, (2) undersidemargin of the indusium.
5. Almost closed sporangium.6. Opened sporangium.7. Streak of spores.89. Single spore in two different optical sections. Figs. 3, 5 ankindly provided by Volker Arnold (Heide, Germany).Scale bars represent 200 m in 12, 100 m in 37 and 10 m in 89.d optical distortions prevent the view inside.a.(56).
egel and Volker Arnold. (see plate on page 149)
r Arnold (no. 1398). The thin hyaline cell layer is visible between the
art at the amber surface during grinding. The indusium is slightly tiltedmed by just one cell layer, except for the part close to the initiationsitee indusium, (3) translucent back break of the stalk, (4) translucent back
ere obtained from different optical sections. Figs. 23 and 57 were
Plate I (caption on page 146).
147A.R. Schmidt, H. Drfelt / Review of Palaeobotany and Palynology 144 (2007) 145156
Plate II (caption on page 146).
148 A.R. Schmidt, H. Drfelt / Review of Palaeobotany and Palynology 144 (2007) 145156
Collection Volker Arnold (Heide, Germany), no. 1399:piece of Baltic amber; five indusia, two of them weredamaged during grinding of the amber; a removed celllayer of the underside of an indusium; 11 sporangia, twoof them damaged at the amber surface; more than 100spores; syninclusions are fungal hyphae, three remnantsof a moss, three pollen grains of Pinaceae, larvae of
aphids, an exuvia of an aphid, a beetle of the Aderidaefamily, an arthropod leg and detritus.
The ground and polished pieces of amber wereinvestigated under an incident light microscope (Stemi2000, Carl Zeiss, Germany) and under a transmittedlight, differential interference contrast microscope
149A.R. Schmidt, H. Drfelt / Review of Palaeobotany and Palynology 144 (2007) 145156Plate III (cap n page 146).
Plate IV. Indusia and sporangia of extant Matonia and Phanerosorus specimens.
1. Upper side of an indusium of M. pectinata.2. Lower side of the indusium of M. pectinata shown in 1. Note the prominent radial ridges at the underside of the umbrella (arrows).3. Irregular arcuate apical cells at the apex of an indusium of M. pectinata.4. Remnant of the receptacle at the lower side of the frond of M. pectinata. The tubular structure is visible.5. Two indusia of P. sarmentosus with attached (left) and largely removed (right) sporangia.6. Sporangia of P. sarmentosus in situ.Scale bars represent 100 m.
150 A.R. Schmidt, H. Drfelt / Review of Palaeobotany and Palynology 144 (2007) 145156
eobot(Axioplan, Carl Zeiss, Germany) with long distanceobjectives (10, 20 and 40) and alternative incidentlight. Photomicrographs were taken with a microscopecamera (MC 100, Carl Zeiss, Germany). Figs. 3, 5 and 6of Plate III were obtained from several optical sections.
In order to protect the amber from oxidation andbreakage, the polished amber piece of Hoffeins col-lection was embedded using polyester resin as describedby Hoffeins (2001).
Extant herbarium specimens of Matonia foxworthyiCopeland, 1908, Matonia pectinata R. Br. ex Wallich,1829, Phanerosorus major Diels, 1932, and P. sarmento-sus (Baker) Copeland, 1908, were investigated from thecollections of the Herbier National de Paris and theNational Herbarium of the Netherlands, Leiden. Matureindusia and sporangia from fertile specimens wereembedded in Canada balsam and investigated usingincident and transmitted light microscopes. Specimensexamined:Matonia foxworthyi: Sarawak, Borneo (LeidenL 0444391); Sarawak, Borneo (Paris P 00312355). M.pectinata: Sarawak, Borneo (Leiden L 0444390); MalayPeninsula (Leiden L 0353091); Malay Peninsula (Paris P00312360); Malay Peninsula (Paris P 00312361). Pha-nerosorus major: New Guinea (Paris H 2003/02160-1).P. sarmentosus: Sarawak, Borneo (Leiden L 0444399);Sarawak, Borneo (Paris P 00312364).
3. Age of the fossils
The material investigated originates from two famousEuropean amber deposits, Baltic and Bitterfeld amber.
The pieces of Baltic amber were found in the Kalinin-grad (Knigsberg) area on the eastern coast of the BalticSea (western Russia). The age of the Baltic amber rangesfrom Early to Late Eocene. Probably over millions ofyears during Middle and Late Eocene, large amounts ofresin were transported by rivers from Fennoscandia intomarine Baltic deposits. Single pieces of Baltic amber aretherefore not assignable to specific strata and horizons(Ritzkowski, 1999). Furthermore, the amber was rede-posited up to several times. The sediments containing themajority of Baltic amber in the Kaliningrad area are4738 million years old. Small amounts of Baltic amber,however, already occur in Lower Eocene deposits andthat material is up to 55 million years old. Therefore,Baltic amber potentially represents material of an inter-val of 9 to 17 million years (Ritzkowski, 1999).
The Bitterfeld amber was found in the Goitzscheminenear Bitterfeld (Sachsen-Anhalt, central Germany). Theage of the Cenozoic amber at this location was some-
A.R. Schmidt, H. Drfelt / Review of Palatimes disputed since a redeposition of Baltic amber or atleast a contemporaneous sedimentation of that amberwas assumed by Rschmann (1997) and Weitschat(1997). These studies emphasize the partly identity ofselected taxa of arthropods entrapped in amber. Theseterrestrial animals, however, have no stratigraphicrelevance and may have persisted morphologicallyunchanged over millions of years in the Palaeogeneeven at species level. Barthel and Hetzer (1982) firstlyprovided a detailed description of the Bitterfeld amberand assigned the amber-containing sediments stratigra-phically to the Early Miocene with an absolute age of 22million years. These authors and later also Kosmowska-Ceranowicz and Krumbiegel (1989) and Krumbiegel(1997) emphasized the independent origin of theBitterfeld amber and saw no connection with EoceneBaltic amber on sedimentological or palaeogeographicalgrounds. Modern studies (e.g. Knuth et al., 2002;Fuhrmann, 2004) confirm the independent origin of theBitterfeld amber. Detailed biostratigraphic investiga-tions recently favour an uppermost Chattian age (Blu-menstengel, 2004). Fuhrmann (2004) additionallyprovides a detailed reconstruction of the depositionalenvironment and local palaeogeography. The Bitterfeldamber occurs in the Bernsteinschluff Horizon, i.e. apart of the Upper Bitterfelder Glimmersand in theupper part of the Cottbus Formation. The amber bearingsediment has an absolute age of 25.323.8 million years(Knuth et al., 2002).
4. Systematic palaeontology
The inclusions are brown to reddish brown in colour.Sometimes, they appear very light due to a refractivefilm between the amber inclusion and resin (e.g. Plate II,16). This film was probably caused by shrinkage ofdrying resin during fossilisation.
The indusia are centrally stalked and umbrella-shapedto almost hemispheric (Plates I, 15, II, 12 and III, 1, 34). Their diameter ranges between 1.1 and 2.0 mm and theheight (margin to apex) between 0.3 and 1.0mm.The stalkis broken at the same level as the margin of the indusium(Plates I, 45 and III, 3) which is due to its attachment siteat the lower side of the frond (receptacle). The diameter ofthe stalk at the fracture is 210 to 400 m. According tothe situation in extant Matoniaceae, the sporangia wereattached to the stalk of the indusium close to its break.
The upper surface of the umbrella is almost regularlyconvex (Plates I, 2, 45 and II, 1) but sometimes six toeight (mostly seven) bumps are visible (Plate III, 1),which correspond to the former location of thesporangia developed below. The umbrella is membra-
151any and Palynology 144 (2007) 145156nous and just one cell layer thick except for the median
part which is multi-layered (Plate III, 4). The margin isinvolute (Plates I, 34 and II, 2, 56). The apex is oftenrecessed (Plate III, 1). The cells of the outer surface arelargely polygonal and rarely irregular arcuate and1050 m in size at the apex (Plate II, 3) and becamegradually more and more elongate toward the margin.Marginally, they form a structure of furrows and ribswhich are anastomosing and branching (Plates I, 6 and II,4). The furrows are due to the thickened cell walls ofthese radial cells and correspond to the striation of thelower side of the umbrella described by Drfelt andStriebich (2000). The distance between the ribs rangesbetween 15 and 40 m. The stalk is apically dilatedmerging regularly into the concave underside of theumbrella (Plate III, 34). The stalk is solidly cellular andconstructed of tubular...