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The First Record of a Fossil Hornwort (Anthocerotophyta) from DominicanAmberAuthor(s) Jan-Peter FrahmSource The Bryologist 108(1)139-141 2005Published By The American Bryological and Lichenological Society IncDOI httpdxdoiorg1016390007-2745(2005)108[139TFROAF]20CO2URL httpwwwbiooneorgdoifull1016390007-2745282005291085B1393ATFROAF5D20CO3B2
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0007-274505$0450
The Bryologist 108(1) pp 139 141Copyright q 2005 by the American Bryological and Lichenological Society Inc
The First Record of a Fossil Hornwort (Anthocerotophyta)from Dominican Amber
JAN-PETER FRAHM
Nees Institut fur Biodiversitat der Pflanzen Rheinische Friedrich-Wilhelms-Universitat Meckenheimer Allee 17053115 Bonn Germany
Abstract A hornwort is reported from Dominican amber (EocenendashOligocene) The specimenshows the horn-like sporophytes typical for this group of plants and can be placed into theDendrocerotaceae although an exact placement in Dendroceros or Megaceros cannot be made itis the first record of a fossil hornwort
Keywords Anthocerotophyta amber Dominican Republic fossil hornwort
There are only few records of fossil bryophytesfrom the tropics as compared to temperate regions(Krassilov amp Schuster 1984 Miller 1984) Duringthe past twenty years bryophytes embedded in am-ber from the Dominican Republic with an age of20ndash40 million years (EocenendashOligocene) have be-come a rich source of information on fossil neo-tropical mosses (Frahm 1993 1996 2001 Frahmamp Reese 1998) and liverworts (Grolle 19831984ab 1987 1990 1993 Grolle amp Braune 1988Gradstein 1993)
Among the material of fossil mosses from Do-minican amber that I received for study from theSmithsonian Institute (Washington DC) there wasa specimen consisting of a thallus with conspicuouslong-cylindric horn-like sporophytes The fragmentis 7 mm long The thallus is 30 3 35 mm largeand ends in a stalk that is 4 mm length The horn-like structures are 18 mm long (Figs 1ndash4) Thisspecimen was forwarded to Riclef Grolle for evalu-ation and publication but reached him only shortlybefore his death Because of the importance of thisspecimen I have proceeded to publish this record
It can be concluded from the thalloid organiza-tion of the plant along with the horn-like structuresresembling sporophytes that this fragment repre-sents a fossil hornwort Further interpretationshowever are difficult to make and it remains un-clear whether the stalk-like structure at the base ofthe thallus is part of the hornwort or not This struc-ture can be interpreted as a midrib from which thelower parts of the thallus have been eroded Fur-thermore the translucent thallus could indicate thatit is unicellular Both the midrib and unicellularundulate wings of the thallus indicate that this fossilrepresents a thallus of the genus Dendroceros An-other characteristic for this genus the presence oftrigones in the thallus cells is not visible Dendro-
ceros is a pantropical genus which in contrast toall other genera of hornworts occurs epiphytic oreven epiphyllous In the Neotropics there is onlyone species D crispus (Sw) Dumort (Gradsteinet al 2001) The plants are 2ndash5 mm wide and inthis regard resemble the fossil material The spo-rophytes are described as 1ndash5 cm in length whenmature and this is not the case in the fossil planthowever these are apparently young and not openThe extant species grows epiphytic in submontaneand lower montane rainforests conditions suitablefor amber forests Dendroceros crispus is said tobe variable and several closely related taxa havebeen described the status of which is not clear(Gradstein et al 2001)
If the stalk-like structure is not part of the horn-wort and thus does not represent a midrib a rep-resentative of the genus Megaceros could be con-sidered Both genera are closely related and unitedin the family Dendrocerotaceae and the order Den-drocerotales Hassel de Mendendez (1962) even in-cluded Megaceros within Dendroceros but latertreated both genera separately (Hassel de Menendez1987) Hasegawa (1980) recognized Dendrocerosas a distinct genus although there are a few inter-mediate species but later placed the two genera indifferent families (Hasegawa 1988) Both generashare capsules without stomata unispiral elatersand greenish spores Megaceros differs from Den-droceros by the presence of several chloroplasts inthe thallus cell a character not visible in the fossilspecimen The unistratose thallus eventual pres-ence of a midrib together with a strap-shaped thal-lus short sporophytes and corticolous growthmake it more probable that the fossil specimen be-longs to Dendroceros compared to Megaceros thathas broadly radiating thalli and very long sporo-phytes
140 [VOL 108THE BRYOLOGIST
FIGURES 1ndash3 Specimen of the Dendrocerotaceae (An-thocerotophyta) from Dominican amber (US 522733) mdash1ndash2 Total plant mdash 3 Upper part of plant
Even if the generic placement (in Dendrocerosor Megaceros) cannot be solved the fossil recordfits well to the moss genera recorded from Domin-ican amber including Adelothecium CalymperesClastobryum Entodon Hypnum Leucobryum Mit-tenothamnium Mniomalia Octoblepharum Or-thostichopsis and Syrrhopodon (Frahm 19931996 2001 Frahm amp Reese 1998) or liverwortgenera such as Archilejeunea BlepharolejeuneaBryopteris Ceratolejeunea Cyrtolejeunea Dre-
panolejeunea Leucolejeunea Lopholejeunea Mar-chesinia Mastigolejeuneaa Neurolejeunea andStictolejeunea (Grolle 1983 1984ab 1987 19901993 Grolle amp Braune 1988 Gradstein 1993)
This record of a hornwort from the Tertiaryseems to be the first record of a complete fossilhornwort plant According to Krassilov and Schus-ter (1984) lsquolsquono credible reports of Anthocerotaefrom pre-cretaceous sediments are known How-ever reports of spores of Anthoceros (Phaeoceros)from the North American Cretaceous are nowknownrsquorsquo There is a questionable record of a horn-wort from the late Cretaceous or Tertiary that wasdescribed as Notothylacites filiformis Nemejc amp Pa-cltova (Oostendorp 1987) This specimen has beendescribed as having a midrib (that is not present inNotothylas) and the illustration suggests the midribis rather a Riccia-like thallus The thallus is de-scribed as 12ndash16 mm wide and the capsules as450ndash500 3 430ndash450 mm (thus almost round) with-out a columella all of which makes placement inthe Anthocerotae much less probable Krassilovand Schuster (1984) related this fossil to theSphaerocarpales or Proto-Ricciineae Another an-thocerotalean record exists from the Plio-Pleisto-cene of Taiwan (Shaw 1994) based only on sporesAdditionally only a sporophyte resembling that ofNotothylas has been reported before from the lateCretaceousPaleocene (Chitaley amp Yawale 1980)
ACKNOWLEDGMENTS
I wish to thank Barbara Crandall-Stotler S RobbertGradstein Karen Renzaglia and an unknown reviewer forcomments on the fossil
LITERATURE CITED
CHITALEY S amp N R YAWALEW 1980 Notothylites ni-rulaei gen et spec nov a petrified sporogonium fromthe Deccan Intertrappean beds of Mahgaonkalan MPIndia Botanique 9 111ndash118
FRAHM J-P 1993 Mosses in Dominican amber Journalof the Hattori Botanical Laboratory 74 249ndash259
mdashmdashmdash 1996 New records of fossil mosses from Domin-ican amber Cryptogamie Bryologie Lichenologie 17231ndash236
mdashmdashmdash 2001 New records of mosses from Dominicanamber Tropical Bryology 20 39ndash42
mdashmdashmdash amp W D REESE 1998 Calymperes palisotii (Mus-ci Calymperaceae) found in Dominican amber THE
BRYOLOGIST 101 131ndash132GRADSTEIN S R 1993 New fossil Hepaticae preserved
in amber of the Dominican Republic Nova Hedwigia57 353ndash374
mdashmdashmdash S P CHURCHILL amp N SALAZAR-ALLEN 2001Guide to the bryophytes of tropical America Memoirsof the New York Botanical Garden 86 1ndash577
GROLLE R 1983 Leucolejeunea antiqua n sp das ersteLebermoos aus Dominikanischem Bernstein Stutt-garter Beitrage zur Naturkunde Ser B 96 1ndash9
mdashmdashmdash 1984a Cyrtolejeunea suzannensis spec novCryptogamie Bryologie Lichenologie 5 27ndash32
2005] 141FRAHM FOSSIL HORNWORT
mdashmdashmdash 1984b Bryopteris und Cyclolejeunea fossil inDominikanischem Bernstein Journal of the HattoriBotanical Laboratory 56 271ndash280
mdashmdashmdash 1987 Radula steerei sp nov a further hepaticin Dominican amber Memoirs of the New York Bo-tanical Garden 45 271ndash280
mdashmdashmdash 1990 Leucolejeunea antiqua (ein Lebermoos ausDominikanischem Bernstein) erstmals mit GynoziumPerianth und Androzium Nova Hedwigia 50 473ndash479
mdashmdashmdash 1993a Ein autozisches Lebermoos mit Perianthin Dominikanischem Bernstein Drepanolejeuneaeogena spec nov Nova Hedwigia 57 375ndash380
mdashmdashmdash 1993b Bryopteris bispinosa spec nov (Lejeu-neaceae) ein weiteres Lebermoos in Dominikanis-chem Bernstein Journal of the Hattori Botanical Lab-oratory 74 71ndash76
mdashmdashmdash amp W BRAUNE 1988 Bazzania oleosandashein Leber-moos mit erhaltenen Olkorpern in DominikanischemBernstein Beihefte zu Nova Hedwigia 90 101ndash108
HASEGAWA J 1980 Taxonomical studies on Asian An-
thocerotaceae II Journal of the Hattori Botanical Lab-oratory 47 287ndash309
HASSEL DE MENENDEZ G G 1962 Estudio de las Antho-cerotales y Marchantiales de la Argentina Opera Lil-liona 7 1ndash297
mdashmdashmdash 1988 A proposal for a new classification of thegenera within the Anthocerotophyta Journal of theHattori Botanical Labotarory 64 71ndash86
KRASSILOV V A amp R M SCHUSTER 1984 Paleozoic andMesozoic fossils pp 1172ndash1193 In R M Schuster(ed) New Manual of Bryology Hattori Nichinan
MILLER N G 1984 Tertiary and Quaternary fossils pp1194ndash1232 In R M Schuster (ed) New Manual ofBryology Hattori Nichinan
OOSTENDORP C 1987 The Bryophytes of the Palaeozoicand the Mesosoic Bryophytorum Bibliotheca 34 Cra-mer Vaduz
SHAW C-L 1984 Fossil sporomorphs of some Antho-cerotae in the Plio-Pleistocene of South Taiwan Yu-shania 1 17ndash28
ms received June 23 2004 accepted Oct 4 2004
0007-274505$0450
The Bryologist 108(1) pp 139 141Copyright q 2005 by the American Bryological and Lichenological Society Inc
The First Record of a Fossil Hornwort (Anthocerotophyta)from Dominican Amber
JAN-PETER FRAHM
Nees Institut fur Biodiversitat der Pflanzen Rheinische Friedrich-Wilhelms-Universitat Meckenheimer Allee 17053115 Bonn Germany
Abstract A hornwort is reported from Dominican amber (EocenendashOligocene) The specimenshows the horn-like sporophytes typical for this group of plants and can be placed into theDendrocerotaceae although an exact placement in Dendroceros or Megaceros cannot be made itis the first record of a fossil hornwort
Keywords Anthocerotophyta amber Dominican Republic fossil hornwort
There are only few records of fossil bryophytesfrom the tropics as compared to temperate regions(Krassilov amp Schuster 1984 Miller 1984) Duringthe past twenty years bryophytes embedded in am-ber from the Dominican Republic with an age of20ndash40 million years (EocenendashOligocene) have be-come a rich source of information on fossil neo-tropical mosses (Frahm 1993 1996 2001 Frahmamp Reese 1998) and liverworts (Grolle 19831984ab 1987 1990 1993 Grolle amp Braune 1988Gradstein 1993)
Among the material of fossil mosses from Do-minican amber that I received for study from theSmithsonian Institute (Washington DC) there wasa specimen consisting of a thallus with conspicuouslong-cylindric horn-like sporophytes The fragmentis 7 mm long The thallus is 30 3 35 mm largeand ends in a stalk that is 4 mm length The horn-like structures are 18 mm long (Figs 1ndash4) Thisspecimen was forwarded to Riclef Grolle for evalu-ation and publication but reached him only shortlybefore his death Because of the importance of thisspecimen I have proceeded to publish this record
It can be concluded from the thalloid organiza-tion of the plant along with the horn-like structuresresembling sporophytes that this fragment repre-sents a fossil hornwort Further interpretationshowever are difficult to make and it remains un-clear whether the stalk-like structure at the base ofthe thallus is part of the hornwort or not This struc-ture can be interpreted as a midrib from which thelower parts of the thallus have been eroded Fur-thermore the translucent thallus could indicate thatit is unicellular Both the midrib and unicellularundulate wings of the thallus indicate that this fossilrepresents a thallus of the genus Dendroceros An-other characteristic for this genus the presence oftrigones in the thallus cells is not visible Dendro-
ceros is a pantropical genus which in contrast toall other genera of hornworts occurs epiphytic oreven epiphyllous In the Neotropics there is onlyone species D crispus (Sw) Dumort (Gradsteinet al 2001) The plants are 2ndash5 mm wide and inthis regard resemble the fossil material The spo-rophytes are described as 1ndash5 cm in length whenmature and this is not the case in the fossil planthowever these are apparently young and not openThe extant species grows epiphytic in submontaneand lower montane rainforests conditions suitablefor amber forests Dendroceros crispus is said tobe variable and several closely related taxa havebeen described the status of which is not clear(Gradstein et al 2001)
If the stalk-like structure is not part of the horn-wort and thus does not represent a midrib a rep-resentative of the genus Megaceros could be con-sidered Both genera are closely related and unitedin the family Dendrocerotaceae and the order Den-drocerotales Hassel de Mendendez (1962) even in-cluded Megaceros within Dendroceros but latertreated both genera separately (Hassel de Menendez1987) Hasegawa (1980) recognized Dendrocerosas a distinct genus although there are a few inter-mediate species but later placed the two genera indifferent families (Hasegawa 1988) Both generashare capsules without stomata unispiral elatersand greenish spores Megaceros differs from Den-droceros by the presence of several chloroplasts inthe thallus cell a character not visible in the fossilspecimen The unistratose thallus eventual pres-ence of a midrib together with a strap-shaped thal-lus short sporophytes and corticolous growthmake it more probable that the fossil specimen be-longs to Dendroceros compared to Megaceros thathas broadly radiating thalli and very long sporo-phytes
140 [VOL 108THE BRYOLOGIST
FIGURES 1ndash3 Specimen of the Dendrocerotaceae (An-thocerotophyta) from Dominican amber (US 522733) mdash1ndash2 Total plant mdash 3 Upper part of plant
Even if the generic placement (in Dendrocerosor Megaceros) cannot be solved the fossil recordfits well to the moss genera recorded from Domin-ican amber including Adelothecium CalymperesClastobryum Entodon Hypnum Leucobryum Mit-tenothamnium Mniomalia Octoblepharum Or-thostichopsis and Syrrhopodon (Frahm 19931996 2001 Frahm amp Reese 1998) or liverwortgenera such as Archilejeunea BlepharolejeuneaBryopteris Ceratolejeunea Cyrtolejeunea Dre-
panolejeunea Leucolejeunea Lopholejeunea Mar-chesinia Mastigolejeuneaa Neurolejeunea andStictolejeunea (Grolle 1983 1984ab 1987 19901993 Grolle amp Braune 1988 Gradstein 1993)
This record of a hornwort from the Tertiaryseems to be the first record of a complete fossilhornwort plant According to Krassilov and Schus-ter (1984) lsquolsquono credible reports of Anthocerotaefrom pre-cretaceous sediments are known How-ever reports of spores of Anthoceros (Phaeoceros)from the North American Cretaceous are nowknownrsquorsquo There is a questionable record of a horn-wort from the late Cretaceous or Tertiary that wasdescribed as Notothylacites filiformis Nemejc amp Pa-cltova (Oostendorp 1987) This specimen has beendescribed as having a midrib (that is not present inNotothylas) and the illustration suggests the midribis rather a Riccia-like thallus The thallus is de-scribed as 12ndash16 mm wide and the capsules as450ndash500 3 430ndash450 mm (thus almost round) with-out a columella all of which makes placement inthe Anthocerotae much less probable Krassilovand Schuster (1984) related this fossil to theSphaerocarpales or Proto-Ricciineae Another an-thocerotalean record exists from the Plio-Pleisto-cene of Taiwan (Shaw 1994) based only on sporesAdditionally only a sporophyte resembling that ofNotothylas has been reported before from the lateCretaceousPaleocene (Chitaley amp Yawale 1980)
ACKNOWLEDGMENTS
I wish to thank Barbara Crandall-Stotler S RobbertGradstein Karen Renzaglia and an unknown reviewer forcomments on the fossil
LITERATURE CITED
CHITALEY S amp N R YAWALEW 1980 Notothylites ni-rulaei gen et spec nov a petrified sporogonium fromthe Deccan Intertrappean beds of Mahgaonkalan MPIndia Botanique 9 111ndash118
FRAHM J-P 1993 Mosses in Dominican amber Journalof the Hattori Botanical Laboratory 74 249ndash259
mdashmdashmdash 1996 New records of fossil mosses from Domin-ican amber Cryptogamie Bryologie Lichenologie 17231ndash236
mdashmdashmdash 2001 New records of mosses from Dominicanamber Tropical Bryology 20 39ndash42
mdashmdashmdash amp W D REESE 1998 Calymperes palisotii (Mus-ci Calymperaceae) found in Dominican amber THE
BRYOLOGIST 101 131ndash132GRADSTEIN S R 1993 New fossil Hepaticae preserved
in amber of the Dominican Republic Nova Hedwigia57 353ndash374
mdashmdashmdash S P CHURCHILL amp N SALAZAR-ALLEN 2001Guide to the bryophytes of tropical America Memoirsof the New York Botanical Garden 86 1ndash577
GROLLE R 1983 Leucolejeunea antiqua n sp das ersteLebermoos aus Dominikanischem Bernstein Stutt-garter Beitrage zur Naturkunde Ser B 96 1ndash9
mdashmdashmdash 1984a Cyrtolejeunea suzannensis spec novCryptogamie Bryologie Lichenologie 5 27ndash32
2005] 141FRAHM FOSSIL HORNWORT
mdashmdashmdash 1984b Bryopteris und Cyclolejeunea fossil inDominikanischem Bernstein Journal of the HattoriBotanical Laboratory 56 271ndash280
mdashmdashmdash 1987 Radula steerei sp nov a further hepaticin Dominican amber Memoirs of the New York Bo-tanical Garden 45 271ndash280
mdashmdashmdash 1990 Leucolejeunea antiqua (ein Lebermoos ausDominikanischem Bernstein) erstmals mit GynoziumPerianth und Androzium Nova Hedwigia 50 473ndash479
mdashmdashmdash 1993a Ein autozisches Lebermoos mit Perianthin Dominikanischem Bernstein Drepanolejeuneaeogena spec nov Nova Hedwigia 57 375ndash380
mdashmdashmdash 1993b Bryopteris bispinosa spec nov (Lejeu-neaceae) ein weiteres Lebermoos in Dominikanis-chem Bernstein Journal of the Hattori Botanical Lab-oratory 74 71ndash76
mdashmdashmdash amp W BRAUNE 1988 Bazzania oleosandashein Leber-moos mit erhaltenen Olkorpern in DominikanischemBernstein Beihefte zu Nova Hedwigia 90 101ndash108
HASEGAWA J 1980 Taxonomical studies on Asian An-
thocerotaceae II Journal of the Hattori Botanical Lab-oratory 47 287ndash309
HASSEL DE MENENDEZ G G 1962 Estudio de las Antho-cerotales y Marchantiales de la Argentina Opera Lil-liona 7 1ndash297
mdashmdashmdash 1988 A proposal for a new classification of thegenera within the Anthocerotophyta Journal of theHattori Botanical Labotarory 64 71ndash86
KRASSILOV V A amp R M SCHUSTER 1984 Paleozoic andMesozoic fossils pp 1172ndash1193 In R M Schuster(ed) New Manual of Bryology Hattori Nichinan
MILLER N G 1984 Tertiary and Quaternary fossils pp1194ndash1232 In R M Schuster (ed) New Manual ofBryology Hattori Nichinan
OOSTENDORP C 1987 The Bryophytes of the Palaeozoicand the Mesosoic Bryophytorum Bibliotheca 34 Cra-mer Vaduz
SHAW C-L 1984 Fossil sporomorphs of some Antho-cerotae in the Plio-Pleistocene of South Taiwan Yu-shania 1 17ndash28
ms received June 23 2004 accepted Oct 4 2004
140 [VOL 108THE BRYOLOGIST
FIGURES 1ndash3 Specimen of the Dendrocerotaceae (An-thocerotophyta) from Dominican amber (US 522733) mdash1ndash2 Total plant mdash 3 Upper part of plant
Even if the generic placement (in Dendrocerosor Megaceros) cannot be solved the fossil recordfits well to the moss genera recorded from Domin-ican amber including Adelothecium CalymperesClastobryum Entodon Hypnum Leucobryum Mit-tenothamnium Mniomalia Octoblepharum Or-thostichopsis and Syrrhopodon (Frahm 19931996 2001 Frahm amp Reese 1998) or liverwortgenera such as Archilejeunea BlepharolejeuneaBryopteris Ceratolejeunea Cyrtolejeunea Dre-
panolejeunea Leucolejeunea Lopholejeunea Mar-chesinia Mastigolejeuneaa Neurolejeunea andStictolejeunea (Grolle 1983 1984ab 1987 19901993 Grolle amp Braune 1988 Gradstein 1993)
This record of a hornwort from the Tertiaryseems to be the first record of a complete fossilhornwort plant According to Krassilov and Schus-ter (1984) lsquolsquono credible reports of Anthocerotaefrom pre-cretaceous sediments are known How-ever reports of spores of Anthoceros (Phaeoceros)from the North American Cretaceous are nowknownrsquorsquo There is a questionable record of a horn-wort from the late Cretaceous or Tertiary that wasdescribed as Notothylacites filiformis Nemejc amp Pa-cltova (Oostendorp 1987) This specimen has beendescribed as having a midrib (that is not present inNotothylas) and the illustration suggests the midribis rather a Riccia-like thallus The thallus is de-scribed as 12ndash16 mm wide and the capsules as450ndash500 3 430ndash450 mm (thus almost round) with-out a columella all of which makes placement inthe Anthocerotae much less probable Krassilovand Schuster (1984) related this fossil to theSphaerocarpales or Proto-Ricciineae Another an-thocerotalean record exists from the Plio-Pleisto-cene of Taiwan (Shaw 1994) based only on sporesAdditionally only a sporophyte resembling that ofNotothylas has been reported before from the lateCretaceousPaleocene (Chitaley amp Yawale 1980)
ACKNOWLEDGMENTS
I wish to thank Barbara Crandall-Stotler S RobbertGradstein Karen Renzaglia and an unknown reviewer forcomments on the fossil
LITERATURE CITED
CHITALEY S amp N R YAWALEW 1980 Notothylites ni-rulaei gen et spec nov a petrified sporogonium fromthe Deccan Intertrappean beds of Mahgaonkalan MPIndia Botanique 9 111ndash118
FRAHM J-P 1993 Mosses in Dominican amber Journalof the Hattori Botanical Laboratory 74 249ndash259
mdashmdashmdash 1996 New records of fossil mosses from Domin-ican amber Cryptogamie Bryologie Lichenologie 17231ndash236
mdashmdashmdash 2001 New records of mosses from Dominicanamber Tropical Bryology 20 39ndash42
mdashmdashmdash amp W D REESE 1998 Calymperes palisotii (Mus-ci Calymperaceae) found in Dominican amber THE
BRYOLOGIST 101 131ndash132GRADSTEIN S R 1993 New fossil Hepaticae preserved
in amber of the Dominican Republic Nova Hedwigia57 353ndash374
mdashmdashmdash S P CHURCHILL amp N SALAZAR-ALLEN 2001Guide to the bryophytes of tropical America Memoirsof the New York Botanical Garden 86 1ndash577
GROLLE R 1983 Leucolejeunea antiqua n sp das ersteLebermoos aus Dominikanischem Bernstein Stutt-garter Beitrage zur Naturkunde Ser B 96 1ndash9
mdashmdashmdash 1984a Cyrtolejeunea suzannensis spec novCryptogamie Bryologie Lichenologie 5 27ndash32
2005] 141FRAHM FOSSIL HORNWORT
mdashmdashmdash 1984b Bryopteris und Cyclolejeunea fossil inDominikanischem Bernstein Journal of the HattoriBotanical Laboratory 56 271ndash280
mdashmdashmdash 1987 Radula steerei sp nov a further hepaticin Dominican amber Memoirs of the New York Bo-tanical Garden 45 271ndash280
mdashmdashmdash 1990 Leucolejeunea antiqua (ein Lebermoos ausDominikanischem Bernstein) erstmals mit GynoziumPerianth und Androzium Nova Hedwigia 50 473ndash479
mdashmdashmdash 1993a Ein autozisches Lebermoos mit Perianthin Dominikanischem Bernstein Drepanolejeuneaeogena spec nov Nova Hedwigia 57 375ndash380
mdashmdashmdash 1993b Bryopteris bispinosa spec nov (Lejeu-neaceae) ein weiteres Lebermoos in Dominikanis-chem Bernstein Journal of the Hattori Botanical Lab-oratory 74 71ndash76
mdashmdashmdash amp W BRAUNE 1988 Bazzania oleosandashein Leber-moos mit erhaltenen Olkorpern in DominikanischemBernstein Beihefte zu Nova Hedwigia 90 101ndash108
HASEGAWA J 1980 Taxonomical studies on Asian An-
thocerotaceae II Journal of the Hattori Botanical Lab-oratory 47 287ndash309
HASSEL DE MENENDEZ G G 1962 Estudio de las Antho-cerotales y Marchantiales de la Argentina Opera Lil-liona 7 1ndash297
mdashmdashmdash 1988 A proposal for a new classification of thegenera within the Anthocerotophyta Journal of theHattori Botanical Labotarory 64 71ndash86
KRASSILOV V A amp R M SCHUSTER 1984 Paleozoic andMesozoic fossils pp 1172ndash1193 In R M Schuster(ed) New Manual of Bryology Hattori Nichinan
MILLER N G 1984 Tertiary and Quaternary fossils pp1194ndash1232 In R M Schuster (ed) New Manual ofBryology Hattori Nichinan
OOSTENDORP C 1987 The Bryophytes of the Palaeozoicand the Mesosoic Bryophytorum Bibliotheca 34 Cra-mer Vaduz
SHAW C-L 1984 Fossil sporomorphs of some Antho-cerotae in the Plio-Pleistocene of South Taiwan Yu-shania 1 17ndash28
ms received June 23 2004 accepted Oct 4 2004
2005] 141FRAHM FOSSIL HORNWORT
mdashmdashmdash 1984b Bryopteris und Cyclolejeunea fossil inDominikanischem Bernstein Journal of the HattoriBotanical Laboratory 56 271ndash280
mdashmdashmdash 1987 Radula steerei sp nov a further hepaticin Dominican amber Memoirs of the New York Bo-tanical Garden 45 271ndash280
mdashmdashmdash 1990 Leucolejeunea antiqua (ein Lebermoos ausDominikanischem Bernstein) erstmals mit GynoziumPerianth und Androzium Nova Hedwigia 50 473ndash479
mdashmdashmdash 1993a Ein autozisches Lebermoos mit Perianthin Dominikanischem Bernstein Drepanolejeuneaeogena spec nov Nova Hedwigia 57 375ndash380
mdashmdashmdash 1993b Bryopteris bispinosa spec nov (Lejeu-neaceae) ein weiteres Lebermoos in Dominikanis-chem Bernstein Journal of the Hattori Botanical Lab-oratory 74 71ndash76
mdashmdashmdash amp W BRAUNE 1988 Bazzania oleosandashein Leber-moos mit erhaltenen Olkorpern in DominikanischemBernstein Beihefte zu Nova Hedwigia 90 101ndash108
HASEGAWA J 1980 Taxonomical studies on Asian An-
thocerotaceae II Journal of the Hattori Botanical Lab-oratory 47 287ndash309
HASSEL DE MENENDEZ G G 1962 Estudio de las Antho-cerotales y Marchantiales de la Argentina Opera Lil-liona 7 1ndash297
mdashmdashmdash 1988 A proposal for a new classification of thegenera within the Anthocerotophyta Journal of theHattori Botanical Labotarory 64 71ndash86
KRASSILOV V A amp R M SCHUSTER 1984 Paleozoic andMesozoic fossils pp 1172ndash1193 In R M Schuster(ed) New Manual of Bryology Hattori Nichinan
MILLER N G 1984 Tertiary and Quaternary fossils pp1194ndash1232 In R M Schuster (ed) New Manual ofBryology Hattori Nichinan
OOSTENDORP C 1987 The Bryophytes of the Palaeozoicand the Mesosoic Bryophytorum Bibliotheca 34 Cra-mer Vaduz
SHAW C-L 1984 Fossil sporomorphs of some Antho-cerotae in the Plio-Pleistocene of South Taiwan Yu-shania 1 17ndash28
ms received June 23 2004 accepted Oct 4 2004
