全 文 ：古生物学报 , 47(2):256-259(2008 年 6 月)
Acta Palaeontolo gica Sinica , 47(2):256-259(June , 2008)
　　收稿日期:2007-04-26
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中泥盆世草本石松植物原始鳞木属的命名问题＊
徐洪河1)　王　祺2)
1)中国科学院南京地质古生物研究所 ,南京 210008 , honghexu@hotmail.com;
2)中国科学院植物研究所系统与进化植物学国家重点实验室 , 北京 100093 , happy king@ibcas.ac.cn
　　提要　草本石松植物原始鳞木属(Protolepidodend ron)一名 , 最初由 Krejcˇi 根据波希米亚泥盆纪地层中一种
具叶的茎干标本所建立 ,当时为了强调其与石炭纪鳞木属(Lepidodendron)石松植物的相似性 ,取名为 P roto-Lep i-
dodend ron。后来 , 这种以具有二分叉的叶为特征的草本石松植物相继在捷克 、德国 、比利时 、美国 、中国 、哈萨克斯
坦 、俄罗斯等地的早 、中泥盆世地层中被发现。值得一提的是 , 对该属及模式种的命名问题存在一定的争议 ,同时
存在多种作者引证 ,甚至还存在不同拼写的种本名。 根据国际植物学命名法规 , 对化石植物的命名必须依据模式
标本。目前 , P rotolepi dodendron 的模式标本保存在捷克共和国首都布拉格的国家博物馆。通过对模式标本进行
的重新观察 ,并发现 Gothan(1921)首次根据模式标本明确了 Protolep idodendron 的属征 ,是该属最早的合格发表
作者;后来 , 模式标本又经过 Kräusel和Weyland(1929)的研究 , 唯一合法的种 P.scharyanum 被确立为模式种 , 并
提出了种征。对 Protolepidodendron 属及其模式种 ,尽管后来有大量研究和报道 , 但都不是基于原始的模式标本。
因此 ,根据国际植物学命名法规 , 该属及其模式种的合法名称应为 Protolepidodendron Krejcˇ i ex Go than 和 P.
scharyanum Krejcˇi ex K räusel et Wey land。
关键词　原始鳞木属　中泥盆世　模式种　合格发表名称
A NOMENCLATURAL NOTE ON THE MIDDLE DEVONIAN
HERBACEOUS LYCOPSID PROTOLEPIDODENDRON
XU Hong-he1) and WANG Qi2)
1)N anj ing Insti tute o f Geo logy and Palaeon tolog y , the Ch inese A cademy o f Sciences , Nanj ing , 210008 , honghexu@hotmai l.com;
2)S tate K ey Laboratory o f S ystemat ic and E vo lut ionary Botany , Inst itute o f Botan y ,
th e Chinese Academ y o f S ciences , Bei j ing 100093 , happy king@ibcas.ac.cn
　　Abstract　Studies on the type specimens of the Middle Devonian herbaceous lycopsid Protolepidodendron are re-
viewed.According to the ICBN , the valid name o f the genus and the type species should be Protolepidodendron Kre j
cˇi ex Gothan , and P .scharyanum Krejcˇi ex Kräusel et Weyland.
　　Key words　Protolepidodendron , Middle Devonian , type species , nomenclatural validity
　 　 Herbaceous lycopsid Protolepidodendron w as
frequently repo rt ed in the Middle Devonian w orldwide
(see Halle , 1936;Stockmans , 1939 , 1940;Leclercq ,
1940;Obrhel , 1961;1968;Gensel and Andrews ,
1984;Jurina , 1988;Cai and Wang , 1995).However ,
authorships of the genus and i ts type species have not
been recognized consistently.There are two ortho-
graphical variants of the type species , namely the origi-
nal spelling P .scharianum (see K rejcˇi , 1880;Obrhel ,
1968;Banks , 1960;Grierson and Banks , 1963;Chal-
oner , 1967;Meyer-Berthaud et al., 2003)and more
widely used name P .scharyanum (see Stur , 1882;
Potonié and Bernard , 1904;Lang , 1926;Hirmer ,
1927;Kräusel and Weyland , 1929;1932;Halle ,
1936;Stockmans , 1939;1940;Leclercq , 1940;Em-
berger , 1944;Obrhel , 1957;Sze , 1960;Dou , 1983;
Cai and Wang , 1995).
Krej cˇi(1880)reported some plant fossils from the
Srbsko Fo rmation , Hostí n , Cent ral Bohemia , Czech
Republic.The new plant s that show a similarity to the
dis tal branch of Carboniferous Lepidodendron Stern-
berg w ere named Proto-Lepidodendron schar ianum
and P.dusl ianum.Krej cˇi described the fo rmer plant
as small leafy twig with small , oval-lanceolate leaves ,
leaf less stem with spirally arranged fusiform leaf scars
(“ Blatt narben” in German).The specific epithet
schar ianum is in memory of M rs.J.M .Schary , who
collected specimens.Krejcˇi gave no generic description
or illust ration and , according to A rticles 41.2 , 43.1 of
the ICBN (McNeill et al., 2006), both designations
were invalid.
Stur(1882 , P l.1 , Fig.1)illust rated K rej cˇi s
(1880)specimen (Tex t-figure 1)and described bifur-
cated leaves.Stur also noted the tight ly arranged leav-
es on the stem but doubted it s relationship with Lepi-
dodendron and t ransferred P.scharianum to Chauvinia
scharyana (algae), and P.duslianum to Barrandeina
duslianum .In cit ations of Krej cˇ i s designation P.
schar ianum , t he spelling of t he specific epi thet w as
slightly changed , i.e.feminine inst ead of neut ral and
`-yana instead of `-ianum .
Potonié and Bernard (1904)described new speci-
mens from the same horizon.The vascular nature of
C.schary ana was confirmed and an orthographical va-
riant , Protolepidodendron schary anum Krej cˇ i w as
used referring a lycopsid with dis tally once-forked leav-
es.Meanwhile , another new species P.kar lsteinii
Potonié et Bernard was established to accommodate
similar plants with simple leaves.Potonié and Bernard
(1904 , Fig.102)illust rated and described Krej cˇi s o-
riginal specimen as Stur(1882)show n , but also no ge-
neric diagnosis was provided.In Potonié and Bernard s
(1904)opinions , Protolepidodendron should include
those plant s with Lepidodendron-like s tem and once-
forked or simple leaves.Among the three species
(Protolepidodendron scharianum , P .duslianum , P.
kar lsteini i)erected by K rej cˇi (1880)and Potonié and
Bernard (1904), only P .scharianum was later accept-
ed (Kräusel and Weyland , 1929;Obrhel , 1957).
Potonié (1912)described new specimens of P .schary-
anum , to w hich distal once-forked leaves were referred
again.
Gothan (1913 , Fig.18 , p.32)quot ed Potonié
and Bernard s (1904)illust ration and at tributed once-
f orked leaves to the distinguished character of the ge-
nus Protolepidodendron af ter making a comparison be-
tween Protolepidodendron and Lepedodendron.Goth-
an (1921 ) re-w rote Potonié s Lehrbuch der
Paläobotanik , in w hich the genus Protolepidodendron
Krejcˇi was diagnosed as :`simple or branched small ax-
es with Lepidodendron-like s tem , but apparent ly lack
of special leaf cushions;leaves f orked dis tally;proba-
bly no relationship with Lepidodendron.Middle Devo-
nian , Bohemia (T ranslat ed f rom German , Gothan ,
1921) .This is the earliest valid publication of the ge-
neric name Protolepidodendron.
Based on K rej cˇ i s original description , Arber
(1921), however , redefined the concept of the genus
Protolepidodendron , to which lanceolate leaves and
spirally arranged leaf bases were referred rather than
bifurcated leaves.
Kräusel and Weyland (1929)studied Protolepi-
dodendron f rom the Middle Devonian of Germany and
Bohemia.Both species of Protolepidodendron erected
by Pot onié and Bernard (1904)were t ransferred t o P .
schary anum (Stur)Krej cˇi and a detailed specific de-
scription was given.Later , Kräusel and Wey land
(1932) studied Protolepidodendron specimens f rom
the Middle Devonian of Elberfeld , Germany.They
designated once-fo rked leaves as the essential feature of
t he genus and recons tructed P .scharyanum as a w hole
herbaceous lycopsid.Both the spelling and reconst ruc-
tion have been widely used by subsequent authors and
in textbooks.Kräusel and Weyland s (1932)work
was only based on German fossils , so it is not appro-
priate to cite as P.schary anum Kräusel and Wey land
as Bonamo , Banks and G rierson (1988)did.
Obrhel(1961)formally designated the K rejcˇi s o-
riginal specimen as the type specimen and P .scharia-
num as the type species.The nomenclature of the spe-
cies was discussed;it is suggested that only P.scharia-
num should be the correct name.All occurrences of the
species by then w ere lis ted.Later , Obrhel (1968)
s tudied the geological age of the Srbsko Formation
(Givetian)and summarized Silurian and Devonian flo-
ras of Cent ral Bohemia.The conception of“ t he Proto-
257　第 2 期 徐洪河等:中泥盆世草本石松植物原始鳞木属的命名问题　　　　
lepidodendron flora”was first proposed.
Jurina and O rlova(2005)thought that the genus
should be cited as Protolepidodendron Krej cˇi 1880 ex
Potonié 1921.However , author of Lehrbuch der
Paläobotanik , who supplied the generic diagnosis of
Protolepidodendron Krejcˇi , is W .Gothan rather than
H.Potonié who died in 1917 (Noé , 1921).
Acco rding t o the ICBN (McNeill et al ., 2006),
the identity of the genus Protolepidodendron should be
based on the type specimen , which is deposited at t he
National Museum of P rague.Therefore , the author-
ships of t he names should be cited as Protolepidoden-
dron Krej cˇi ex Gothan 1921 and Protolepidodendron
scharyanum Krejcˇi ex K räusel et Wey land , 1929.The
orthographical variant P.schar ianum is not a validly
published designation hence no status under the ICBN
(McNeill et al ., 2006 , Art.12.1)despit e that it pre-
dat es P.schary anum.The nomenclatural hist ory of
Protolepidodendron and the type species P.schary a-
num is listed below :
Protolepidodendron Krejcˇi , 1880 ex Gothan , 1921
1880　Proto-Lepidod end ron , Krej cˇi , p.203.(invalidly pu blish ed ,
w ithou t generic diagnosis and il lust ration)
1882　Chauvin ia Bo ry , Stur , p.333 , pl.1 , fi g.1.(synonym)
1913　Protolepidodendron , Gothan , p.30 , f ig.18.(fi rst at t rib ute
forked leaves to the character of the genus)
1921　Proto lep idodendron , Gothan , p.225.(wi th generic diagno-
si s , valid name)
Type species 　 Protolepidodendron scharyanum
Krej cˇi ex Kräusel et Weyland , 1929(Text-figure 1-A ,
B)
1880　Proto-Lep id odend ron schar ianum Krej cˇi , pp.203.(invalid
design tion)
1882　Chauvinia scharyana S tu r , p.333 , pl.1 , fig.1.(synonym)
1904　P rotolep idod end ron scharyanum K rej cˇi , Potonié and Ber-
nard , p.42 , fi g.102.(invalid designt ion)
1929　Proto lep idoden dron sch aryanum Krej cˇi , Krä usel et Wey-
land , p.25.(valid n ame)
1961　Proto lep idoden dron schar ianum Krej cˇi , Ob rhel , p.15.(in-
valid design tion)
1967　Protolepidodendron schar ianum Potonié et Bernard , Chalo-
ner , p.455 , fig.317F.(invalid designt ion)
　　Type locality　Hostí n , the Czech Republic.
Horizon 　Srbsko Formation , late Middle Devoni-
an (Givetian)(Obrhel , 1961).
Type specimen　 D-372.Text-figure 1-A.
Repository　The National Museum , Prague , t he
Czech Republic.
Generic and specific diagnoses　Herbaceous lycop-
sid with Lpeidodendron-like s tem;stem at least 10 cm
T ext-figu re 1　Protolepidod end ron sch aryanum Krej cˇi ex
K rä usel et Weyland.
A.Ty pe specim en deposited at th e National Museum of Prague ,
the Czech Republic.D-372.Arrow indicates th e dis tal bifu rca-
t ed leaf.S cale bar = 10 mm.
B.Th e i llus t rat ion of the type specimen cited by S tu r (1882 ,
Pl.1 , Fig.1), similar d raw ings al so see Fig.102 of Potonié
and Bernard(1904), Fig.18 of Gothan(1913)and Fig.317F
of Ch aloner (1967).
long , 2.5 -3 mm wide.Dis tally once-f orked leaves
tightly inserted in a tight ly spi ral fashion.
Acknowledgments　We thank M .Libertí n , cura-
to r of palaeobotany at the National Museum of Prague
258 古　生　物　学　报 第 47 卷　　　　　　
for providing access to the type specimen;A.Jurina ,
Moscow State University and M .Raskatova , Vo ronez
St ate University , for helpful discussion.
参 考 文 献 (References)
Arber E A N , 1921.Devonian f loras , a study of the origin of cormo-
phy ta.C amb ridge:Cam bridge Universi ty Pres s.42.
Bank s H P , 1960.Notes on Devonian ly copods.Senckenbergiana
Lethaea , 41:59—88.
Bonamo P M , Banks H P , Grierson J D , 1988.Leclercqia , H askin-
sia , and the role of leaves in delineat ion of Devonian lycopod
genera.Botanical Gazet te , 149:222—239.
C ai Chong-yang , Wang Yi , 1995.Devonian f loras.In:Li Xing-xu e
et a l.(eds.), Fos si l f loras of China through the geological a-
ges.Guangzh ou:Guan gdon g Science and T echnology Press.
28—77.
C haloner W G , 1967.Lycophy ta.I n:Bou reau E , Jover-Art S ,
H eg O A et al.(ed s.), T raité de Palé obotanique II.Paris:
M asson et Cie.435—802.
Dou Ya-wei(窦亚伟), 1983.Palaeobotany .In:Geological Su rvey-
ing T eam and Inst itute of Geoscience , Xinjiang Bureau of Geol-
ogy , Geological Surveying Department , Xinjiang Bureau of Pe-
t roleum (eds.), Palaeontological Illust ration s of Northwest
C hina , Xinjiang Uygur Autonom ous Region , Vol., 2.Beijing:
Geological Pub lishin g H ouse.561—614(in Chinese).
Emberger L , 1944.Les p lan tes fossiles dans leurs rapp orts avec les
vé gé t aux vivants.Pari s:Mason et Cie.1—492.
Gensel P G , And rew s H N , 1984.Plant lif e in the Devonian.New
York:Praeger Publi shers.1—381.
Gothan W , 1913.En tw ick lungs geschich te der Pf lanzenwelt.In:
Gothan W , Pilger R , Winkler H (eds.), Das Leben der Pflanz e
6(3).S tu ttgart , Kosm os:Fran ckhsche Verlagshandlung.3—
116.
Gothan W , 1921.Potonié s Lehrbu ch der Paläobotanik , Zwet te
Anflage(2nd edit ion).Berlin:Verlag von Gebrǜder Bornt rae-
ger.225—226.
G rierson J D , Banks H P , 1963.Lycopods of th e Devonian of New
York S tate.Palaeon tographica Americana , 4:220—295.
Halle T G , 1936.On Drepanophycus , Protolepidod end ron , and
Proto pterid um , w ith notes on th e Palaeozoic f lora of Yunnan.
Paleon tologia S inica , 1A:1—28.
Hirm er M , 1927.H andbuch der Palä ob otanik , I.Thallop hyta ,
Bryophy ta , Pteridophyta.Mǜnchen und Berlin:R.Olden-
bou rg.1—708.
Jurina A L , 1988.The Middle and Late Devonian flora of North
Eurasia.Trudy Palaeon tologicheskigi Ins tituta , 227:1—175(in
Rus sian).
Jurina A L , Orlova O A , 2005.Ly copsid Proto lep idoden dron
schar ianum Krej cˇi(1880)ex Krä usel et Weyland , 1929 , th e
collection of th e Nat ional Museum of Prague:h olotype and th e
prob lem of the autho rship.T heses of report s , the XVII In ter-
national Botanical Conference.50—52(in Russian).
Krä usel R , Weyland H , 1929.Beit räge zu r Kenntnis der Devonf lora
III.Abhandlungen der Senck enb ergis chen Naturfo rschenden
Gesell schaf t , 41:315—360.
Krä usel R , Weyland H , 1932.Pf lanzenreste aus dem Devon.IX ,
P rotolep id odend ron.S encken bergiana , 14:391—403.
Krej cˇ i J , 1880.Noti z ǜber die Res te von Landpf lanzen in der
bö hmischen Silu rformat ion. Sit zung sberichte der Kö niglich-
Bö hmis chen Gesel lsch af t der Wis senschaf ten , 201—204.
Lang W H , 1926.Con t rib utions to the study of the Old Red S and-
stone Flora of S cot land .IV.On a specimen of P rotolep idoden-
dron from th e Middle Old Red Sandstone of C aithness.T ransac-
ti ons of the Royal Society of Edin burgh , 54:785—799.
Leclercq S , 1940.Cont ribut ion à l é tude de la flore du Dé vonien de
Belgiqu e.Mém oires du Acadé mie Royale de Belgique , 12:1—
65.
McNeill J , Bar rie F R , Burdet H M et al., 2006.Internat ional Code
of Botanical Nomenclature(Vienna Code), adopted by the Sev-
en teen th Internat ional Botanical Congress Vienna , Au st ria , Ju-
ly 2005(Regnum Vegetabi le , 146).A.R.G.Gan tner Verlag ,
Ruggell , Liechtenstein.1—568.
Meyer-Berthaud B , Fai ron-Dem aret M , Steemans P et al., 2003.
T he plant Lecler cqia (Lycopsida) in Gondw ana:implications
for reconst ructing Middle Devonian palaeogeography .Geolog-
ical Magazine , 140:119—130.
Noé A C , 1921.New text books of paleobotany.Botanical Gazette ,
71:238—239.
Obrhel J , 1957.T he Flora of the Daleje-S t rata of the Cent ral Bohe-
mian Devonian.Sborní k U′st rˇ edníh o ú stavu geologickéh o , 23:
523—531(in C zechish).
Obrhel J , 1961.Die Flora der S rb sko-Schichten(Givet)des mit tel-
boehmisch en Devons.Sborní k U′st rˇední h o ú stavu geologické ho ,
26:7—46.
Obrhel J , 1968.Die Si lur- und Devonflora des Barrandiums.
Paläeontologi sche Abhandlungen , 2B:661—701.
Potonié H , 1912.G rundlinien der Pflanzen-Morphologie im lichte
der Palaeontologie , Jena , Verlag von Gu stav Fischer.110.
Potonié H , Bernard C , 1904.Flora Dévonienne de l é t age H-h1 de
Barrande.Prague , Raimund Gerh ard and Wolfgan g Gerhard.
39—43.
Stockman s F , 1939.Note Pré liminai re sur des empreintes vé gé t ales
du Dé vonien infé rieur de la Belgique.Bullet in du Musé e Royal
d Histoi re Natu relle de Belgique , 15:1—6.
Stockman s F , 1940.Vé gé taux évodé voniens de la Belgique.
Mé moires du Musée Royal d H istoire Naturelle de Belgique ,
93:1—90.
Stu r D , 1882.Die Silurf lora der tage H-h1 in Böhm en.S it zungs-
berichte der Kaiserlichen Akademie der Wissenschaf ten , 84:
330—391.
Sze H-C(斯行健), 1960.On some Devonian plants f rom the Dzuan-
garia Basin.Acta Palaeontologica Sinica , 8(3):205—216(in
Chinese w ith Engli sh ab st ract).
259　第 2 期 徐洪河等:中泥盆世草本石松植物原始鳞木属的命名问题