Evidence of Cenozoic Matoniaceae from Baltic and Bitterfeld amber

nology 144 (2007) 145–156www.elsevier.com/locate/revpalbo

Review of Palaeobotany and Paly

Evidence of Cenozoic Matoniaceae from Baltic and Bitterfeld amber

Alexander R. Schmidt a,⁎, Heinrich Dörfelt b

a Museum für Naturkunde der Humboldt-Universität zu Berlin, Institut für Paläontologie, Invalidenstr. 43, D-10115 Berlin, Germanyb Martin-Luther-Universität Halle, Institut für Geobotanik und Botanischer Garten, Neuwerk 21, D-06099 Halle/Saale, Germany

Received 11 November 2005; received in revised form 4 July 2006; accepted 10 July 2006Available online 20 September 2006

Abstract

Matoniaceae occurred in European amber forests in the Eocene and at the Oligocene–Miocene boundary. This is now revealedby resin-preserved indusia, sporangia and spores. These inclusions are the first Cenozoic fossil records of this archaic fern family,which was previously known from Mesozoic macrofossils and spores and few relict species. The findings from non-tropical forestsshow 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 (Dörfelt et Striebich)Schmidt et Dörfelt, comb. nov. was previously described as a fungus, Palaeocybe striata [Dörfelt, H., Striebich, B., 2000.Palaeocybe striata, ein neuer fossiler Pilz in Bernstein des Tertiär. Zeitschrift für 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; Tertiary

1. Introduction

Matoniaceae 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

⁎ Corresponding author. Tel.: +49 30 2093 8945.E-mail address: [email protected]

(A.R. Schmidt).

0034-6667/$ - see front matter © 2006 Elsevier B.V. All rights reserved.doi:10.1016/j.revpalbo.2006.07.009

four extant species exist and they are restricted to astrongly equatorial relict area in Southeast Asia.

Recently, several inclusions were found in Baltic andBitterfeld 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 largeumbrella-shaped indusia are less known amongpalaeontologists and neontologists, that specimen wasassumed to be a cap of a basidiocarpium similar to theextant genus Coprinus and consequently described as afungus (Dörfelt and Striebich, 2000). Afterwards, a

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146 A.R. Schmidt, H. Dörfelt / Review of Palaeobotany and Palynology 144 (2007) 145–156

further inclusion was found in the Hoffeins collectionand Günter 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.

Plate I. Holotype of Matonia striata (collection Bernhard Striebich, no. 21/8

1. Piece of amber with the indusium.2. Detail of the indusium showing the almost hemispheric shape of t3. Lower side of the umbrella with visible striation.4–5. Indusium in oblique lateral view with the stalk (4) and marginal r6. Detail of the margin with branching and anastomosing ribs.Scale bars represent 1 mm in 1–2, 500 μm in 3–5 and 100 μm in 6.

Plate II. Paratype of Matonia striata (collection Christel and Hans Werner H

1. Upper side of the indusium.2. Lower side of the indusium. A bubble of air, detritus particles an3. Polygonal apical cells at the top of the upper side of the umbrell4–6. Details of the involute margin in upper view (4) and lower viewScale bars represent 500 μm in 1–2 and 100 μm in 3–6.

Plate III. Specimens of Matonia striata from the collections Günter Krumbi

1. Indusium of the collection Günter Krumbiegel (no. KRU 2).2. Lower side of the closed immature sorus of the collection Volke

involute margin and the centrally located break of the stalk.3–9. Indusia, sporangia and spores of the collection Volker Arnold (n3. Entire indusium (paratype).4. Reconstruction of an indusium which was damaged at its median p

to the front. The cut surface shows that the umbrella is largely forof the stalk; (1) cut surface in the median part, (2) underside of thmargin of the indusium.

5. Almost closed sporangium.6. Opened sporangium.7. Streak of spores.8–9. Single spore in two different optical sections. Figs. 3, 5 and 6 w

kindly provided by Volker Arnold (Heide, Germany).Scale bars represent 200 μm in 1–2, 100 μm in 3–7 and 10 μm in 8–9.

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 (Dörfelt 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 Günter 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.

/56). (see plate on page 147)

he umbrella.

egion of the upper side (5) in focus.

offeins, no. 809). (see plate on page 148)

d optical distortions prevent the view inside.a.(5–6).

egel and Volker Arnold. (see plate on page 149)

r Arnold (no. 1398). The thin hyaline cell layer is visible between the

o. 1399).

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. 2–3 and 5–7 were

Plate I (caption on page 146).

147A.R. Schmidt, H. Dörfelt / Review of Palaeobotany and Palynology 144 (2007) 145–156

Plate II (caption on page 146).



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

Plate III (caption o

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

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 μm.



(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 (Königsberg) 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 are47–38 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-times disputed since a redeposition of Baltic amber or at

least a contemporaneous sedimentation of that amberwas assumed by Röschmann (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 25.3–23.8 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,1–6). This film was probably caused by shrinkage ofdrying resin during fossilisation.

The indusia are centrally stalked and umbrella-shapedto almost hemispheric (Plates I, 1–5, II, 1–2 and III, 1, 3–4). 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, 4–5 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 μm. 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, 4–5 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-nous and just one cell layer thick except for the median


part which is multi-layered (Plate III, 4). The margin isinvolute (Plates I, 3–4 and II, 2, 5–6). The apex is oftenrecessed (Plate III, 1). The cells of the outer surface arelargely polygonal and rarely irregular arcuate and10–50 μm 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 Dörfelt andStriebich (2000). The distance between the ribs rangesbetween 15 and 40 μm. The stalk is apically dilatedmerging regularly into the concave underside of theumbrella (Plate III, 3–4). The stalk is solidly cellular andconstructed of tubular cells. These cells are 20–35 μm indiameter and their walls ca 5–8 μm thick (see also PlateIV, 4). Sporangia of immature sori of extant species areprotected by a thin layer which extends from the marginof the persistent part of the umbrella to the base of thestalk of the indusium. This layer is only preserved in theimmature closed indusium of piece no. 1398 (Plate III, 2)and removed in piece 1399 of Arnold's collection. In

Table 1Morphological comparison of the specimens of fossil indusia investigated (?piece of amber)

CollectionStriebich no.21/8/56, PlateI, 1–6

Hoffeins no.809, Plate II,1–6

Krumbiegelno. KRU 2,Plate III, 1

Arnold no.1398 (closeindusium),Plate III, 2

Shape of theumbrella

Almosthemispheric,regularlyconvex

Almosthemispheric,regularlyconvex

Umbrella-shaped,irregularlybumpy,recessed

Umbrella-shaped

Shape of themargin

Involute Involute ? Involute

Diameter [mm] 1.6 2.0×1.9 ca 2 1.9×0.76Height (margin toapex) [mm]

0.8 ca 1 ? ca 0.32

Marginal ribdistance [μm]

25–35 ca 30 ca. 30 ca 20

Cell diameter(apex) [μm]

? ca 20–50 ? 10–20

Stalk diameter atthe fracture[μm]

400 400 ? 240×200

Diameter of thetubular cells atthe stalk fracture[μm]

25–35 20–30 ? ?

Wall diameter ofthe tubular cellsat the stalkfracture [μm]

? 5–8 ? ?

extant specimens, this layer is probably one cell layerthick and removes from the margin to expose the maturesporangia, which are shown in Plate IV, 5–6.

The features of the fossil indusia investigated areshown in Table 1.

We can estimate that a sorus bore six to eight, mostlyseven, sporangia. Most of the 11 sporangia preserved inamber are opened and up to 660 μm long (Plate III, 6).Few closed or almost closed sporangia are 560–620 μmin size (Plate III, 5). The number of annulus cells rangesbetween 14 and 19.

More than 100 spores are preserved. They are arrangedin clusters and streaks (Plate III, 7). The spores arerounded triangular in polar view and have an equatorialdiameter of 35–50 μm. Superficially, most spores appearvery light in colour which is due to a refractive filmbetween the inclusions and the resin. Few spores are bettervisible and show a trilete aperture (Plate III, 8–9).

Field observations by Walker and Jermy (1982)revealed that the indusia of the Matoniaceae do notshrink during drying. So we can assume that the her-barium specimens investigated are adequate for com-parison with the fossils.

=feature not visible because of the orientation of the inclusion in the

dArnold no.1399, PlateIII, 3

Arnold no.1399 notillustrated



Arnold no.1399,Plate III, 4

Hemispheric,bumpy

Umbrella-shaped,slightlybumpy

Umbrella-shapedwithbumps

Umbrella-shapedwithbumps

Umbrella-shaped,regularlyconvex

Involute Probablyinvolute

? Involute Partlyinvolute

1.3×1.2 1.2×1.1 1.3×1.2 1.3 1.30.34 0.33 0.56 0.53 0.54

20–30 20–30 15–40 ca 30 ?

20–40 20–30 10–25 ? ?

240 ? ? ? 210

? ? ? ? ?

? ? ? ? ?


Generally, we could not securely distinguish theindusia of the four extant species of Matoniaceae whichare nowadays accepted for the genera Matonia andPhanerosorus (Kato, 1989, 1993). As shown in Table 2,the features of the fossil indusia largely correspond tothose of the extant genera Matonia and Phanerosorus.However, most fossils have a nearly perfect umbrella tohemispheric shape. This is because the bumps at theformer locations of the sporangia are less prominent infossil specimens. Between the bumps at the upper side,radial ridges are prominent at the underside of theumbrella of extant specimens (Plate IV, 2). These ridgeswere never found in the fossils. The cells of the outersurface of extant indusia are largely irregular arcuate atthe apex and just few polygonal cells occur betweenthem (Plate IV, 3). Furthermore, the fossil indusia havean involute margin whereas that of extant specimens isusually elongate. The fossil indusia appear to be morerobust than extant ones which always have one toseveral radial cracks (Plate IV, 1–2).

The assignment of the inclusions to the Matoniaceaenecessitates a nomenclatural change. We follow the con-vention to assign all matoniaceous fossils with indusia

Table 2Morphological comparison of the indusia, sporangia and spores of the extan

Fossils

IndusiumShape of the umbrella Umbrella-shaped to hemispher

with slight bumpsProminent radial ridges

at the undersideAbsent

Shape of the margin InvoluteDiameter [mm] 2.0–1.2×1.9–1.1Height (margin to

apex) [mm]1.0–0.3

Marginal ribdistance [μm]

15–40

Cell diameter and shape(apex) [μm]

10–50,largely polygonal

Stalk diameter at thefracture [μm]

210–400

Diameter of the tubular cellsat the stalk fracture [μm]

20–35

Wall diameter of the tubular cellsat thestalk fracture [μm]

5–8

SporangiaNumber of sporangia

in each sorus(6–) 7 (–8)

Number of annulus cells 14–19Maximum diameter of

closed sporangia [μm]560–620

Spore diameter [μm] (35–) 40 (–50)

similar to that of extant representatives to the genus Ma-tonia (see Appert, 1973; Harris, 1980; van Konijnenburg-van Cittert, 1993). We provide two paratypes additionallyto the holotype described by Dörfelt and Striebich (2000).

Family Matoniaceae Presl, 1847Genus Matonia R. Br. in Wallich, 1829Type species: Matonia pectinata R. Br. in Wallich, 1829

Matonia striata (Dörfelt et Striebich) Schmidt etDörfelt, comb. nov. (Plates I, II and III); Plate I showsthe holotype).Basionym: Palaeocybe striata gen. et sp. nov. Dörfeltet Striebich 2000, Zeitschrift für Mykologie 66: 31,Figs. 1–6.Holotype: Oligocene amber inclusion, collection Bern-hard Striebich (Buxtehude, Germany) no. 21/8/56(Plate I, 1–6).Paratypes: Oligocene amber inclusion, collectionChristel and Hans Werner Hoffeins (Hamburg, Ger-many) no. 809 (Plate II, 1–6); and Eocene amber in-clusions, collection Volker Arnold (Heide, Germany)no. 1399 (Plate III, 3).

t genera Matonia and Phanerosorus with the fossils

Extant Matonia ExtantPhanerosorus

ic Umbrella-shaped withprominent bumps

Umbrella-shaped withprominent bumps

Present Present

Elongate Elongate1.3–1.1×1.2–0.9 1.8–1.1×1.8–1.10.3–0.4 0.4–0.52

20–40 10–40

50–30×20–15,largely arcuate

50–20×10–17,largely arcuate

370–250×300–160 400–160×340–100

(9–) 15–25 (–35) 20–30

3–5 3–5

5–10 7–10

17–25 20–26340–510 370–550

50–65 45–60


Type locality: Former coal and amber mine Goitzscheeast of the city of Bitterfeld, central Germany.Stratigraphic range and age: Mid Eocene (Blue Earthfrom the Kaliningrad area, 47–38 million years) toOligocene–Miocene boundary (Upper Chattian, “Bern-steinschluff” Horizon of the Upper Cottbus Formation,25.3–23.8 million years).Etymology: The specific epithet was given by Dörfeltand Striebich (2000) to the proposed fungal inclusionbecause of the striation of the upper and lower side ofthe umbrella which is caused by the shape of radialelongate cells.Emended diagnosis: Fossil matoniaceous fern indusi-um, differentiated from those of extant Matonia andPhanerosorus species by an almost hemispheric shape,by the lack of prominent radial ridges at the underside ofthe umbrella, by an involute margin and by largelypolygonal cells at the apex.Emended description:Centrally stalked umbrella-shapedto almost hemispheric indusium of 1.1 to 2.0 mmdiameter and 0.3–1.0 mm height; margin membranousand involute.

5. Palaeoecology

The Baltic amber forests grew in Scandinavia in atemperate to subtropical climate. Members of anarctotertiary flora of a circumboreal temperate climaticzone such as deciduous trees and members of apalaeotropical flora with evergreen trees and numerouspalms grew together in that area (Kohlman-Adamska,2001). The main resin-producing plants are stilldisputed (see Weitschat and Wichard, 1998). Based onthe analysis of macroinclusions and amber-preservedpalynomorphs, Kohlman-Adamska (2001) provides ahypothetical reconstruction of the plant communities ofresin-abundant forests and distinguishes between asso-ciations of higher and lower mountain altitudes as wellas river valleys. The coniferous forests of highermountain altitudes may have consisted of Pinaceaesuch as Abies, Larix and Picea and of Cupressaceae.Pine-palm-oak forests should have grown in lower andlowest mountain altitudes and consisted, apart fromPinus species, mainly of palms and Fagaceae includingQuercus, Castanea and Fagus. Representatives of thefamilies Aceracae, Aquifoliaceae, Cycadaceae, Laur-aceae, Magnoliaceae and Pittosporaceae occurred inthese forests as well. Humid river valleys weredominated by Cupressaceae, Myricaceae, Clethraceae,Connaraceae and Salicaceae.

The central German Bitterfeld amber forest grewin the uppermost Oligocene. Geinitziaceae such as

Cupressospermum saxonicum, Pinaceae and Fagaceae(Quercus spp.) were already described from that resin(Barthel and Hetzer, 1982). The Thuja syninclusionclose to the Matonia striata indusium of the Strie-bich collection shows presence of Cupressaceae aswell.

A rich flora of liverworts is well known from bothamber deposits (Grolle and Meister, 2004). Ferns,however, are generally very rarely preserved in fossilresins. The only ferns described from Baltic amber weredescribed by Goeppert and Berendt (1845) and byCaspary (1887). To date no fern inclusion becameknown from Bitterfeld amber. Indusia, sporangia andspores might have been more easily transported by windto fresh resin outflows than remnants of fronds.

At least some habitats within the amber forests musthave been suitable for matoniaceous ferns. They mayhave grown at the forest floor, at steep slopes or at rockslike extant species. Mature indusia got attached to freshresin outflows. Extant indusia which are fallen off oftenstill contain sporangia and explain the occurrence ofsporangia and spores close to the indusia in Arnold'sspecimen no. 1399.

6. Mesozoic and Cenozoic Matoniaceae

The fossil history of the Matoniaceae extends at leastinto the Early Mesozoic. The family was cosmopolitanin the Mid-Mesozoic and shows recently a narrowendemism (Tidwell and Ash, 1994).

Matoniaceae are recorded by fronds, permineralisedrhizomes, indusia or spores. Leaves and branching typeof most fossil representatives resemble the fronds of theextant genus Matonia closely. The modern genus Pha-nerosorus, however, has no close fossil relatives(Walker and Jermy, 1982). Already Hirmer andHörhammer (1936) emphasized the apomorphic fea-tures of that genus and assumed Phanerosorus to bevery young.

The oldest leaf fossils attributed to theMatoniaceae areLate Triassic (Carnian) specimens of the genus Phlebob-teris fromUSA, Greenland and Europe (Tidwell and Ash,1994). AntarcticMiddle Triassic permineralised rhizomeswere tentatively assigned to matoniaceous ferns (Millayand Taylor, 1990). Recently, peltate indusia withaffiliation toMatoniaceaewere described from theMiddleTriassic of Antarctica (Kalvins et al., 2004).

Matoniaceous ferns were common and most diversefrom the Lower Jurassic to the Lower Cretaceous ofNorth America, Europe and Asia. Further importantfinds from Greenland, Antarctica, Australia, Africa andMadagascar show that the Matoniaceae occurred on


every part of Pangaea. Generally, leaf fossils are assignedto the genera Aninopteris, Delosorus, Matonia, Mato-nidium, Nathorstia, Phlebobteris, Piazopteris, Sele-nocarpus and Weichselia. Among those, the genusPhlebobteris was widely distributed and contains atleast 10 species (Tidwell and Ash, 1994; Givulescu andPopa, 1998; Skog, 2001).

The Lower Liassic Selenocarpus has crescent-shapedsori without indusia. Phlebobteris and similar taxa showround sori without indusia. The Lower Jurassic to LowerCretaceous genusMatonidium has subpeltate indusia. Apeltate indusium is developed in the genera Matoniaand Phanerosorus. Based on this presence and absenceof an indusium and its size, an evolutionary lineage wasassumed which resulted in an increasing protection ofthe sporangia by an indusium (Hirmer and Hörhammer,1936; Harris, 1961). According to this theory, anelaboration of the receptacle through time resulted inthe development of a peltate indusium. However, thefinding of a peltate indusium in the Middle Triassic byKalvins et al. (2004) shows that this development musthave taken place very early. Kalvins et al. (2004)therefore postulated that the Matoniaceae originate fromPermian or Early Triassic.

The genus Matonia firstly appears in the LowerJurassic. AlreadyM. braunii (Goeppert) Harris from theLower Jurassic of Germany and Greenland and theMiddle Jurassic of England as well as the MiddleJurassic M. mesozoica Appert from Madagascar pos-sessed indusia without any doubt (Appert, 1973; Harris,1980). Also the Lower Cretaceous M. brownii (Rush-forth) van Konijnenburg-van Cittert is placed into thisgenus on the basis of the presence of a large, peltateindusium (van Konijnenburg-van Cittert, 1993). TheLate Triassic Phlebobteris muensteri (Schenk) Hirmeret Hörhammer was proposed to be conspecific with M.braunii (Harris, 1980); however, this was not acceptedby later examinations (van Konijnenburg-van Cittert,1993) because peltate indusia are not visible.

Apart from macrofossils described from foliage,permineralised rhizomes and spores are importantrecords showing a wide distribution of the family.Matoniaceous genera were described on the basis ofpermineralised rhizomes. Some of them such as theSiberian Matoniopteris, the Tasmanian Heweria andTasmanopteris and the Antarctic Soloropteris show awider Mesozoic distribution as indicated by leaf fossils(Tidwell and Ash, 1994).

The latest Mesozoic records of Matoniaceae arespores of Narthorstia galleyi (Miner) van Konijnen-burg-van Cittert from the Cenomanian of the USA (vanKonijnenburg-van Cittert, 1993) and Matonia-like

permineralised rhizomes from the Japanese Turonianto Santonian (Nishida, 1991). Spores from Matoniaceaewere also reported from the Maastrichtian of Argentina(Papú, 2002).

The records of Matoniaceae from European Eoceneand Upper Oligocene amber forests provide the missingCenozoic fossils of this archaic fern family. The amberinclusions of Matonia striata show that the family wasstill present in Europe during the whole Palaeogene andthat the formation of the Southeast Asian relict area ofthe family happened much later than previouslyproposed.

7. Conclusions

The finds of Eocene and uppermost Oligocene Mato-niaceae close a frequently discussed gap in the fossilrecord of matoniaceous ferns. The period of restrictionof the Matoniaceae to the extant relict area in SoutheastAsia can therefore now be confined into the Neogene.

The palaeogeographical context of the previous fossilrecord of the Matoniaceae (see Skog, 2001) and themodern equatorial relict area indicate that this family isuntypical for non-tropical coniferous and mixed forestssuch as the European Palaeogene amber forests. It istherefore likely that the inclusions of Bitterfeld amberdocument the latest matoniaceous ferns in Europe sincewe can not expect younger finds in Europe according tothe continuous cooling in the Neogene. Maybe Matoniastriata was already a relict species in the EuropeanPalaeogene.

Acknowledgements

Especially, we would like to express our gratitude tothe amber collectors Volker Arnold (Heide), Christel andHans Werner Hoffeins (Hamburg), Brigitte and GünterKrumbiegel (Halle/Saale) and Bernhard Striebich (Bux-tehude) for the kind permission to investigate their fossilsand for an excellent cooperation over years. We thankManfred Barthel (Berlin), Horst Blumenstengel (Jena),Wilfried Krutzsch (Berlin), Kerstin Schultz (Jena),Johanna H.A. van Konijnenburg-van Cittert (Utrecht)and Hans-Joachim Zündorf (Jena) for helpful informationand advice. Special thanks to the curators Marc Pignal(National Herbarium of Paris) and Gerhard Thijsse(National Herbarium of the Netherlands, Leiden) for theloans of extant species. The work on amber-microfossilsis supported by the German Research Foundation, projectnumber SCHM 2152/1-1.

The amber collector Volker Arnold now kindlydecided to donate the amber pieces described above to


the Museum für Naturkunde der Humboldt-Universitätzu Berlin for final deposition: collection no. MB. Pb.2007/161 (no. 1398 of the collection Volker Arnold) andcollection no. MB. Pb. 2007/162 (no. 1399 of thecollection Volker Arnold and paratype of Matoniastriata).

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Evidence of Cenozoic Matoniaceae from Baltic and Bitterfeld amber