全 文 ：古生物学报 , 39(1):81-91(2000 年 1 月)
Acta Palaeontologica Sinica , 39(1):81-91(Jan., 2000)
　　收稿日期:1999-07-02
　　＊所长基金和国家自然科学基金 9390010项目资助。
＊＊Email:Guo.S.X@jlonline.com
辽宁西部晚侏罗世晚期义县组的似麻黄属植物＊
郭双兴＊＊ 　吴向午
(中国科学院南京地质古生物研究所　南京 210008)
　　提要　《辽西义县组单子叶植物化石的发现》中被归入被子叶植物门单子叶植物纲莎草科和禾本科的化石 ,经
与现代麻黄科植物对比研究后 ,发现它们应归入裸子植物门麻黄科似麻黄属。讨论似麻黄属植物的命名 、地理分
布和生态环境。
关键词　似麻黄属　盖子植物纲　裸子植物门　晚侏罗世晚期　义县组　辽宁西部
　　半个多世纪前 ,矢部长克和远藤诚道(Yabe and
Endo , 1935)曾在热河凌源(现属辽宁西部)早白垩
世? 狼鳍鱼层(现名义县组)发现一些保存欠佳的植
物化石 ,并鉴定为被子植物门单子叶植物纲眼子菜
科的 Potamogeton jeholensis Yabe et Endo 和 Pota-
mogeton ?sp.。
近年来 ,曹正尧 、吴舜卿等(1997 , 1998)在辽宁
北票晚侏罗世晚期义县组中发现大量植物化石 ,其
中有一些被他们鉴定为被子植物门单子叶植物纲莎
草科和禾本科的新属 、种。这些白垩纪以前的所谓
被子植物化石引起了国内学者的广泛关注 。然而 ,
现经核查这些植物化石的特征与现代莎草科和禾本
科的特征并不相符。化石与禾本科共同的特征仅是
茎具节和节间 ,而两者的其它特征差别甚远 ,化石与
莎草科的特征则毫无共同之处。实际上 ,这些化石
与现代麻黄科的特征颇为接近 。它们应是裸子植物
麻黄科的化石。
麻黄科属于盖子植物纲(Chlamydospermopsi-
da),又称买麻藤纲(Gnetopsida)或买麻藤目(Gne-
tales)。该纲或目还包含另外两科 ,即百岁兰科和买
麻藤科 ,它们都是单属科 ,是裸子植物门中最进化的
类群 。该纲植物化石极其稀少 。目前已知麻黄科约
有 16个化石种 ,它是此纲中最原始的类群。百岁兰
科尚未见其化石报道 。买麻藤科仅记录一种化石种
(Jongmans und Dijkst ra , 1973)。该科植物的叶片
和具网状结构的脉序与被子植物门双子叶植物纲一
致 ,二者的叶相(foliar physiognomy)特征很难区分。
此外 ,美国弗吉尼亚早白垩世 Potomac群发现的具
有网状脉的 Drewria 属被归入买麻藤科(Crane and
Upchurch , 1987)。
麻黄科现代植株呈灌木 ,亚灌木 ,藤本或偶草本
状 。茎细 ,圆柱状 ,直立或匍匐 ,具节和节间 ,节间具
多数细纵沟槽:茎上的小枝对生或轮生 。苗期的初
生叶为针形或线形 ,以后退化成膜质叶或鳞片状叶 ,
叶交互对生 ,轮生或 2—3(4)片合成鞘状生于茎节
基部或下部 ,顶端具三角状齿 ,叶有 1—3条平行脉。
花单性 ,雌雄异株 ,稀同株 ,球花卵圆形或椭圆形 ,生
于枝顶或叶腋;雄球花单生 ,或数个丛生或 3—5个
组成复穗花序 ,具 2—8 对生或 2—8 轮生(每轮 3
片)的苞片 ,圆形或倒卵形 ,雄球花具膜质假花被 ,每
个苞片生 1雄花;雌球花亦具 2—8对生或 2—8轮
生(每轮 3片)的苞片 ,仅顶端 1 —3 苞片生雌花;雌
花具顶端开口的革质假花被 ,包于胚珠外 ,珠被上部
延伸成直或弯的珠被管 。种子 1—3粒。子叶 2枚。
麻黄科现只有麻黄属(Ephedra Tourn ex
Linn.),含 40余种 ,广布于亚洲中 、西部 ,欧洲南部 ,
非洲北部 ,北美西南部 ,南美西部和南部的干旱 、荒
漠地区;我国有 12种 4变种 ,除长江下游及珠江流
域外 ,全国各地均有分布 ,以西北和西南各地种类较
多 ,常生于干旱山地 ,土壤贫瘠的山坡及荒漠戈壁 ,
是一类耐干旱的植物(郑万钧 、傅立国 ,1978;郑万钧
主编 ,1983;侯宽昭等 , 1982)。
盖子植物纲 Chlamydospermopsida (买麻藤纲 Gen-
topsida)
麻黄目 Ephedrales
麻黄科 Ephedraceae
似麻黄属 Ephedri tes Saporta , 1891 (non Goeppert
et Berendt , 1845)
特征　茎具小枝 ,茎枝通常具分散的条纹和关
节 ,小坚果(指雌球花)成双对生 ,其表面具明显的联
合缝 ,背腹面类似 ,基部强烈结合在一起 ,初生的苞
片密集 ,小坚果(雌球花)的变态叶后来开放 ,互生分
离 ,脱落。
讨论 　似麻黄属(Ephedrites)系 Goeppert 和
Berendt (1845 ,见 Berendt ,1845)依据产自德国北部
中新世植物而创立的 ,原模式种 Ehedri tes johnianus
Goeppert et Berendt 的标本曾被 Goeppert(1853)和
Conw entz(1886)分别改置于 Ephedra 和被子植物
桑 寄 生 科 (Loranthaceae)Patzea 属 , 因 此 ,
Ephedri tes成了裸名 ,虽然如此 ,至今仍有很多学者
沿用此属名 ,将一些与现代麻黄类似而又有所区别
的植物化石归入似麻黄属(Ettingshausen , 1890;
Saporta , 1891 , Seward , 1919;吴向午等 , 1986 ;李佩
娟等 , 1988;周志炎 , 1995)。特别是 Sapo rta(1891 ,
p.22)给似麻黄属以新的属征 。这个新属征虽然也
未完整地反映似麻黄属的特征 ,并在术语和特征的
认识上同现代植物还有一些差别 ,但这个属征大致
与现代麻黄属特征接近 ,可作为似麻黄属的新属征 。
Saporta(1891 ,p.26)并根据他的新属征将产自法国
侏罗纪的标本建立一新种(Ephedrites armail len-
sis),并对产自西伯利亚东部侏罗纪的 E .antiquus
Heer(Heer , 1876 , p.82)重新记述和讨论。这表明
Saporta 有意将其作为本属的新的模式种(ty pe
species)对待。 E .ant iquus具有保存很好的茎枝和
种子 ,本文也认为此种可选作似麻黄属的模式种 。
基于似麻黄属名习用已久 ,虽然最初的模式标本被
改归它属 ,但 Saporta (1891 , p.22)重新补述和讨论
了似麻黄属的属征已被后来学者接受 ,并使用至今 ,
本文也以 Saporta 的似麻黄属属征为准。似麻黄属
名语义明白 ,能望文生义 ,本文建议应予以保留 ,若
改为其他名称 ,在使用上会产生诸多不便。
陈氏似麻黄(新组合)Ephedrites chenii (Cao et
Wu)Guo et WuX.W.Comb.nov.
(图版Ⅰ ,图 1—7;图版Ⅱ ,图 1—8)
1997 　Liaoxia chenii Cao et Wu , 曹正尧等 , 1764页 ,图版Ⅰ , 图 1 ,
2 , 2a—2c。
1997 　Erogrosites changii Cao et Wu , 曹正尧等 , 1765 页 ,图版 Ⅱ ,
图 1—3 , 2 , 2a—2c;插图 1。
1998　Liao xia cheni i Cao et Wu , Cao et al ., p.231 , pl.Ⅰ , figs.1 , 2 ,
2a—2c.
1998 　Erogrosi tes changi i Cao et Wu , Cao et a l., p.231—232 , pl.
Ⅱ , f igs.1—3 , 2a—2c;Tex t-fig.1.
增订特征　植物茎丛生 ,保存长 3 —15cm ,茎枝
具节和节间;节略膨大 ,宽 1.5—6.5mm , 节间长
10 —35mm ,宽 1—5mm ,具纵沟槽;茎枝直伸或略弯
曲;侧枝以 30°—60°角自主枝节上伸出 ,交互对生。
叶自节基部伸出 ,对生 ,线形 ,长 15—30mm ,宽1mm
左右。雌球花椭圆形 ,倒卵形或圆形 ,长 4—7mm ,
宽 2—4mm ,通常单生于小枝顶端或近顶端 ,雌球花
具 4—8对交互对生的苞片 ,苞片卵形至长卵形;顶
端锐尖 ,长2 —5mm ,宽1 —2mm 。种子? 长椭圆形 ,
长 2.5mm ,宽 1mm 。
讨论　本文研究的标本共 8件 ,其中 PBJ31是
本所东方标本中心收藏的标本 ,其余 7件是曹正尧 、
吴舜卿(1997)研究过的标本 ,除 2 件(PB 17805 ,PB
17806即本文图版Ⅰ ,图 8—10)被原作者鉴定为“单
子叶植物” ,实际上它们属于松柏类 ,可能与苏铁杉
有关 ,本文不予描述外 ,其余 6件均是似麻黄属的球
花序和茎枝标本。这些标本与现代麻黄属的特征基
本一致 ,即茎枝直立 ,具明显的节和节间 ,节间具纵
沟槽 ,小枝对生 ,球花生于小枝顶端或近顶端;但当
前化石具一线形叶与现代多数麻黄具膜质叶或鳞片
叶 ,叶基部联合成叶鞘的特征有所区别 ,化石的线形
叶可能反映一种原始性状。德国早侏罗世的麻黄科
分子 Piroconi tes kuespert ii Gothan 的苞叶亦成带
状 ,并具多数平行脉(Van Konijnenburg-van Ci ttert ,
1992;Kirchner ,1992;Crane ,1996)。现代麻黄也有
一些具细长针形叶和线形叶的种 ,如 Ephedra fol i-
ata , E .f ragilis , E .al tissima和E .chi lensis ,其中
有的种其叶长30mm ,宽 1—1.5mm ,具 2—3条平行
脉 (Sew ard , 1919 ;Foster and Gifford , 1974 )。
Ephedra vulgaris 和 E .alt issima 的实生苗的几对
初生叶 ,也具针形或线形叶(Rendle , 1953)。因此 ,
当前具线形叶的标本归入似麻黄属是易于理解的。
化石的雌球花与现代麻黄也很相似。化石的雌球花
常生于小枝顶端或近顶端 ,这与分布在我国新疆西
部 、俄罗斯 、印度和巴基斯坦现代的细子麻黄
(Ephedra regeliana Florin)和生于中国北部和西部
及中亚的单子麻黄(Ephedra monosperma Gmel.ex
82 　　 　　　古　生　物　学　报 第 39卷
Mey)的特征颇相类似(郑万钧 、傅立国 , 1978 ;郑万
钧主编 ,1983)。化石中仅发现一粒 ,有可能是陈氏
似麻黄的种子? 因种子顶端延伸的珠被管未被保留
而存疑 ,然而 ,这种顶端延伸的珠被管在搬运和埋藏
过程中也极易受损而失落 。
麻黄科麻黄属的化石现知有 11 种之多 。通常
学者将发现于第三纪的 ,并与现代麻黄特征颇相一
致的化石归入麻黄属 ,目前麻黄属的化石已记录 9
种 ,它们产自美国早第三纪 ,德国 、意大利 、瑞士等地
第三纪 ,澳大利亚早第三纪和智利第三纪地层 。此
外尚有 2 个未定名的种(Jongmans und Dijkst ra ,
1974 ,p.352—353)。
一些产自中生代的与麻黄属接近的化石 ,或少
数虽产自第三系 ,因各种原因不能确信它们可以归
入现 代 麻 黄 的 化 石 , 常 被 归 入 似 麻 黄 属
(Ephedri tes)。似麻黄属现已记录 5 种(Jongmans
und Dijkst ra ,1974)。其中 2种 E .sinensis Wu , He
et Mai和 E .exhibens Wu , He et Mei产自我国青
海柴达木盆地下侏罗统小煤沟组(吴向午等 , 1986)。
其余 3种 ,如 E .ant iquus Heer(1876)是茎枝和雌球
花化石 ,发现于俄罗斯西伯利亚的侏罗纪地层中 ,是
保存最完好的标本;E .armai llensis Saporta 是雌球
果化石 , 发现于法国的侏罗纪地层中(Saporta ,
1891 ;Seward , 1919;Jongmans und Dijkst ra , 1974);
E .sotzkianus Unger常见于法国 ,奥地利 ,瑞士 ,意
大利和罗马尼亚等地区的第三纪地层中 ,是欧洲的
广布种(Jongmans und Dijkst ra ,1974)。我们没有机
会去核查所有似麻黄属种的鉴定是否都是正确的 ,
但是 ,本文描述的标本确实与现代麻黄属的特征非
常相似 ,因此 ,当前标本归属裸子植物似麻黄属是适
宜的 ,它们不属被子植物中的单子叶植物是肯定无
疑的 。
曹正尧 、吴舜卿将当前的部分化石标本(曹正尧
等 ,1997 , 1764—1765页;Cao et al .,1998 ,p.231 ,图
版Ⅰ ,图 1—2 , 2a—c即本文图版Ⅰ ,图 1—4)归入莎
草科的新属种(Liaoxia cheni i Cao et Wu),并与鳞
子莎属(Lepidosperma)和墨莎草属(Gnhnia)相比
较。莎草科的茎秆实心 ,常三(4 —5)棱状或偶圆柱
状 ,茎无节和节间;叶片狭长 ,或偶缺 ,常三列 。而目
前化石的茎具明显的节和节间这个重要特征排除它
们与莎草科的关系。莎草科化石已报道 10余属 ,最
早出现于古新世(LaMo tte ,1952)。该科的枝叶化石
不易确定其属种的分类位置 ,其茎叶化石常归入形
态属似莎草属(Cyperacites =Cyperi tes)。莎草科现
有 70—90 属 , 约 4 000 种(Hayw ood , 1978;Cron-
quist , 1981), 中国有 31 属 , 670 种(侯宽昭等 ,
1982)。
曹正尧 、吴舜卿同时又将另一部分化石标本(曹
正尧等 ,1997 , 1765页;Cao et al ., 1998 , p.231 ,图
版Ⅱ ,图 1—3 ;插图 1 即本文图版 Ⅰ ,图 5—7 ;图版
Ⅱ , 图 3—8)归入禾本科的新属种 Eragrosites
changii Cao et Wu),并与画眉草属(Eragrostis)相比
较 。禾本科的茎虽具节和节间与当前化石类似 ,但
禾本科的茎常中空 ,茎节上的侧枝通常不对称 ,即侧
枝长短不齐 ,且无对生性;叶形各种各样 ,通常披针
形 ,均具叶鞘和叶舌 ,叶互生非对生 ,常二列;花的苞
片通常 2—3片 ,非对生 。这些禾本科的重要特征在
目前化石中均未得到证实 ,故当前化石不应归入禾
本科。禾本科的化石已报道 20 余属(LaMot te ,
1952)。依据此科的茎叶化石很难鉴定其属的分类
位置 ,除非果实 ,表皮和叶内植硅石同时发现。据
Cronquist(1981)报道 ,禾本科最早的化石记录是晚
白垩世 Senonian 期 ,从始新世开始逐渐丰富起来。
禾本科现有 500—660余属 ,8 000—10 000种 ,是单
子叶植物中最大的科之一(Hayw ood , 1978;Cron-
quist ,1981),中国有 225 属 , 1 200 余种(侯宽昭等 ,
1982)。
此外 ,曹正尧 、吴舜卿还将正 、反面的两个枝叶
化石(曹正尧等 , 1997 ,1965页;Cao et al ., 1998 ,p.
232 ,图版Ⅰ ,图 3 , 4 , 4a即本文图版 Ⅰ ,图 8—10)称
做未定名的单子叶植物也是值得怀疑的。它们应属
松柏类 ,可能与苏铁杉类 ? (Podozam ites ?)的小枝
有关。
段淑英(1997)也在此同一产地和同层位报道一
“被子植物”新属种梁氏朝阳序(Chaoyangia liangi i
Duan)。这类化石也不像是被子植物 , 很可能是盖
子植物纲的植物。
三木茂(Miki , 1964 , p.3—22)记录一个产自中
国东北中生代狼鳍鱼层(现辽宁义县组)的标本 ,并
依此建立新属种 Amphiephedra rhamnoides Miki ,
他认为化石与麻黄属接近 ,具有特别短的枝 ,但枝上
无与现代麻黄近似的叶。因其标本保存破碎 ,特征
不明显 ,难以确定其分类位置。
原河北凌源(现属辽宁省)早白垩世? 狼鳍鱼层
的标本被定名为被子植物 Potamogeton jeholensis
Yabe et Endo 和 Potamogeton ?sp.(Yabe and Endo ,
1935 ,p .274 —276),其特征或多或少地与当前标本
有些接近 ,它们也许与似麻黄属有关 。可是后来它
83第 1 期 郭双兴等:辽宁西部晚侏罗世晚期义县组的似麻黄属植物　　　　
们又被 Miki 改归毛莨属 Ranunculus jeholensis
(Yabe et Endo)Miki(1964 , p.19)。其真实性质和
归属有待确证。
Velanovsky 和 Viniklar (1926)根据发现于捷克
白垩 纪地层 的化石 建立 拟麻 黄新 属种 , 即
Ephedropsis strobil ifera ,后来此种被归入杉科(An-
drews , 1970),这个属名虽具有麻黄(Ephedr-)的词
干 ,但它与麻黄科无关。
德国早侏罗世发现的 Piroconites kuespert ii
Gothan ,其苞叶带状 ,长 9 —15cm ,宽 4—5cm , 具有
多数平行的不分叉的脉 ,每厘米有 12 —15 条;小孢
子叶有凹的新月形的基部;雌球花鳞片长 3.5 —
5.5cm ,宽 2.0—3.5cm(Van Konijenburg-van Cit-
tert , 1992;Kirchner , 1992)。因其花粉与麻黄粉类
似而归入麻黄科 。此种特征与当前的标本差别甚
远。
最近 , 澳大利亚的维多利亚早白垩世晚期
Koonw arra 化石层发现的麻黄科植物 Leongathia
elegans Krassilov ,Dilcher and Douglas(1998),它的茎
枝有节和节间 ,并具纵条纹;叶线形 , 4 叶或偶尔 2
至 3叶交互对生排列于节上 ,叶鞘退化 ,这些特征与
我国的标本有些类似 ,但当前标本的其它特征与前
者有明显区别。
Crane(1987 , 1996)指出麻黄类的花粉化石三叠
纪时在北半球已经广泛分布 ,在我国华北下三叠统
也有记载(Ouyang and Norris , 1988),侏罗纪较少 ,
但继续发展到白垩纪 ,中白垩世在低纬度较前更加
丰富。据我国的孢粉学者黎文本 、尚玉珂和刘兆生
等面告 ,我国辽宁侏罗纪义县组发现过类似于现代
麻黄属具粗条纹饰的花粉化石 ,其它地区侏罗纪地
层中也曾发现过此类麻黄花粉化石 。
在本文的研究工作中 ,有关化石种的命名问题
曾与本所周志炎教授和南京大学张永辂教授讨论
过 ,并得到他们的有益建议。本文即将完稿时 ,恰逢
瑞典自然历史博物馆古植物学部主任 Else M arie
Friis教授来本所访问 。她仔细地观察了本文研究
的标本 ,对其命名 ,被子植物起源和早期演化等问题
同作者进行了广泛讨论和交流 ,事后 ,又赠送有关学
术论文。江苏省及中国科学院植物研究所提供查阅
腊叶标本 ,宋之耀拍摄照片 ,本所东方标本中心张志
平借用一件标本 。笔者对他们的帮助表示衷心感
谢。
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EPHEDRITES FROM LATEST JURASSICYIXIAN FORMATION IN
WESTERN LIAONING , NORTHEAST CHINA
GUO Shuang-Xing and WU Xiang-Wu
(Nanjing Inst itute of Geology and Palaeontology , Chinese Academy of Sciences , Nanjing 210008, China)
Key words:Ephedrites , Chlamydospermopsida , Gymnosperm , Latest Jurassic , Yixian Formation , Western Liaoning
　　Over half of a century ago , some fossil plants in
bad preservation discovered from the Lower Creta-
ceous? Lycoptera Bed in Lingyan County of Jehol
(now Western Liaoning)was identified as Potamoge-
ton jeholensis Yabe et Endo and Potamogeton sp.
(Yabe and Endo , 1935 , p .274—276)belonging to
family Po tamogetonaceae of monocotyledon of an-
giospermae.This report had not att racted at tention
from palaeobo tanists.
In recent years , a g reat number of fossil plants
have been found from the Latest Jurassic Yixian For-
mation in Yixian county of Liaoning Province ,
Northeast China.Among them , some of specimens
w ere identified as monocoty ledons of angiosperm by
Cao Zhengyao , Wu Shunqing and others (1997 ,
1998). These “ monocoty ledons ” fossils were
spearately named tw o new genera Liaox ia of Cyper-
aceae and Eragrosites of Gramineae(Poaceae)(Cao
Zhengyao , Wu Shunqing and others , 1997 , 1998).
However many characters of these so-called mono-
coty ledonous fossils are actually close to the living
Ephedra of gymnospermae rather than monocotyle-
donous plants of angiospermae.Their morphological
characters of stems , branches , leaves and inf ructes-
cences are much different from both families Cype-
raceae and Gramineae.
Ephedraceae belongs to Chlamydospermopsida
(Gnetopsida or Gnetales)which is composed of three
families , the other two families being Welw itschi-
aceae and Gnetaceae.Each family contains only one
genus.This class is the most advanced g roup in gym-
nospermae.The fossl plants of this class are rare.So
far as we know there are about 16 fossil species of
Ephedraceae.Fossil of Welw itschiaceae has not been
known yet.Only one fossil species of Gnetaceae has
been discovered in the passed years (Jongmans und
Dijkst ra , 1974).Recently , a new fossil genus
Drewria with ret iculate veins w as att ributed to Gne-
taceae.It w as discovered f rom the Early Cretaceous
Potomac Group in Virginia of the United States
85第 1 期 郭双兴等:辽宁西部晚侏罗世晚期义县组的似麻黄属植物　　　　
(Crane and Upchurch , 1987).The Gnetaceae leaves
w ith reticulate veins are co rresponding to those of di-
co tyledons.Both of them are hardly to distinguish in
their foliar phy siognomy from each other.
The living ephedraceans are represented by
w oody shrubs , subshrubs , vines and occasionally
herbs.They are characterized by the follow ing fea-
tures:stem slender , cylindraceous , upright and pros-
trate;nodes and internodes w ith longi tudinal ridges;
branches diverging f rom stem in pairs or whorled.
Primary leaves subulifo rm or linear in seedling , later
reduced to scale or membranous leaf , opposite or in
w horl of 2 to 3 (4)with their bases combining
togerther to fo rm sheath at nodes , and triangularly
teethed at apex , 1—3 parallel-veined.Flower nor-
mally dioecious , raely monoecious or hermaphrodite ,
born on teminal of branches or axillary;male st robile
solitary , some clustered , or 3—5 forming spiculate
spike, 2—8 opposi te or 2—8 whorled bracts (three
bracts per w heel), rounded or obovate , one male
flow er per bract;male flower wi th pseudo-perianth in
membrane;female strobile 2—8 opposite or 2—8
whorled bracts (three bracts per w heel);female
flow er only born at apical 1—3 bracts , female f low er
w ith an apical o ra formed by cy stic-coriaceous false
perianth surrounding ovule;integument w ith elon-
gate , erect or curve tube in its apex , Seed 1—3.
Coty ledon tw o.
The ex tant genus Ephedra has over 40 species.
It is widely dist ributed in arid and desert areas of
Central and West Asia , South Europe , North Africa ,
southw estern part of North America , southern and
w estern parts of Latin America.There are 12 living
species and 4 varieties in China.With exception of
v alleys of the Yangtze and Zhujiang Rivers , they are
w idely distributed everyw here of China.However ,
they are mainly grow ing in droughty land , barren
slopes , Gobi and desert of northwestern and south-
western parts of China (Zheng and Fu , 1978;
Zheng , 1983 ;Huo , 1982).
Class Chlamydospermopsida(Gentopsida)
Order Ephedrales
Family Ephedraceae
Genus Ephedrites Saporta , 1891 , non Goeppert et
Berendt , 1845
Diagnosis:Rami remulique plerumque dist racti
striat i articulati;nuculæ binæ geminatim appositæe ,
facie commissuralli plana adpresse convenientes , basi
ext rema cohaerentes , bracteis primum stipatæ;
nuculæ bracteaeque post anthesim ab alterutra liberæ,
deciduæque.
Discussion:The above diagnosis of Ephedrites
was given as a new definit ion by saporta(1891).The
original diagnosis of Ephedri tes established by Goep-
pert et Berendt (1845 see Berendt , 1845)was based
on the Miocene plants f rom no rthern Germany .Lat-
er , the specimens of the type species Ephedri tes joh-
nianus Goeppert et Berendt were separately trans-
ferred to the living Ephedra and Patzea of Loran-
thaceae of angiospermae (Conwentz , 1886).The
genus Ephedrites becomes an empty name (nomen
nudum).However , the fossil plants mo re or less sim-
ilar to the living Ephedra have still been at tributed to
Ephedrites by many palaeobo taists since then (Et-
tingshausen , 1890;Saporta , 1891;Sew ard , 1919;
Wu et al ., 1986;Li et al ., 1988;Zhou , 1995),
especially af ter saporta (1891 , p.22)given a new
definition and supplemented new characters of
Ephedrites.He has redescribed and discussed the
Jurassic species E .ant iquus Heer from easten Siberia
in detail at the same time.The generic name
Ephedrites used in the present article is in Saporta s
sense with the well-preserved E .antiquus Heer as
the type species.Sapo rta (1891 , p.26)also estab-
lished a new species Ephedri tes armail lensis based on
some specimens collected f rom Jurassic in France.
Ephedri tes chenii (Cao et Wu)Guo et Wu X.W.
comb.nov.
(Pl.Ⅰ , figs.1—7;Pl.Ⅱ , figs.1—8)
1997　Liaoxia chenii Cao et Wu , p.1764—1765 , pl.Ⅰ , figs.1 , 2 ,
2a—2c.
1997　E rag rosites changi i Cao et Wu , p.1765, pl.Ⅱ , figs , 1—3 , 1a ,
2a—2c;text-fig.1.
1998　Liao xia cheni i Cao et Wu , p.231 , pl.Ⅰ , figs.1 , 2 , 2a—c.
1998　Eragrosites changii Cao et Wu , p.231—232 , pl.Ⅱ , figs.1—3 ,
1a , 2a—2c;text-fig.1.
Emended diagnosis:Stem cluster , upright or
slight ly curved , 3 —15 cm long , w ith nodes and in-
ternodes;nodes a li tt le expanded , 1.5—6 .5mm
w ide;internodes 10—35 mm long , 1—5 mm w ide ,
w ith longitudinal ridges and g rooves;stem and
branches st raight or a lit tle curved;lateral branches
diverging from stem nodes at angles of 30°to 60°,
oppsite-decussate.Leaves diverging f rom base of
node , linear , opposite in pair , 15—30 mm long and 1
86 　　 　　　古　生　物　学　报 第 39卷
mm w ide.Female st robile elliptical , obovate and sub-
rotund , 4—7 mm long and 2—4 mm wide , borne at
terminals or near apices of branches;female f low er
w ith 4—8 decussate bracts;bracts ovate to narrow-
ovate , acute at apex , 2—5 mm long , 1—2mm wide;
seed ? elliptical , 2 .5 mm long , 1 mm wide.
Discussion:In total , 8 specimens may be at-
tributed to the present species.Among them , one
specimen (PBJ31)is collected by Oriental Collections
of Fossil and Craft in our Institute.Other 7 speci-
mens were studied by Cao and Wu , S .Q .et al .,
(1997 , 1998).Tw o (PB 17805 , PB 17806)of 7
specimens w ere at tributed to “monocotyledons” by
Cao and Wu , S .Q .et al ., (1997 , 1998).In fact ,
the tw o specimens should be referable to conifers and
might be related to Podozam ites .The o ther 5 speci-
mens are generally identical w ith living Ephedra in
the follow ing characters:stem upright , stem and
branches w ith distinct nodes and internodes , inter-
nodes w ith longitudinal ridges and g rooves , and
branches opposite in pai r.However , one of the pre-
sent fossils w ith a linear leaf (pl.Ⅰ , figs.1 , 2)is
somewhat different from those most living species of
Ephedra with scale and membranous leaves combined
in sheath at base.This linear leaf might reflect a
primi tive characteristic of Ephedra .The Early Juras-
sic ephedralean fossils , described as Piroconi tes (male
reproductive o rgan), and Bernet tia (female repro-
ductive organ) are also believed to have paired
Desm iophylum-lide leaves (Van Konijnenburg-van
Cit tert , 1992;Kirchner , 1992 ;Crane , 1996).Even
so , such linear leaves are also found in some extant
species of Ephedra , e.g .Ephedra chilensis , E .fo-
liata and E .f ragi lis bearing linear leaves 30 mm
long and 1—1.5 mm wide , and with 2—3 parallel
veins (Sew ard , 1919;Foster and Giffo rd , 1974).In
addi tion , such linear leaves are also present in the
seedling of Ephedra alt issima and E .vulgaris (Ren-
dle , 1953).Therefore , the present fossil specimens
are morphologically close to Ephedra .The st robiles
at tached at the top or near the top of branches in the
present specimens are also similar to those of extant
species of Ephedra .These characteristics can be seen
in living Ephedra regeliana Florin grow ing in Xin-
jiang , Northwest China , Kazakhstan , Pakistan and
India and Ephedra monosperma Gmel.ex Mey.
g row ing in North and West China and Russia(Cheng
and Fu , 1978;Cheng Wanchun (Ed.in Chief),
1983).There is only one doubtful seed discovered in
the present specimens.Its apical-st retched tube of in-
tegument w as not preserved.
There are about 11 fossil species of Ephedra of
Ephedraceae so far as w e have known.Usually , the
fossil plants from Tertiary are quite similar to moderm
Ephedra and are at tributed to Ephedra Tournef .ex
Linn.9 fossil species of Ephedra have been recorded
from the Early Tertiary strata in the United States
and Chile and from Tertiary in Germany , Italy ,
Switzerland and Australia.In addi tion , there are 2
undeterminable species (Jongmans und Dijkstra ,
1974 ,p .352 —353).
The fossil plants found from M esozoic , or even
from Tertiary , which are hardly ident ical w ith
Ephedra or in poor preservat ion were alw ays at-
t ributed to the fossil genus Ephedri tes .So far , there
are 5 species of Ephedri tes (Jongmans und Dijkstra ,
1974).Ephedrites antiquus Heer (1876)is repre-
sented by some branches and strobiles in good preser-
vation discovered f rom the Jurassic st rata of Siberia ,
Russia.E .armail lensis Sapo rta is composed of fossil
st robiles collected f rom the Jurassic strata in France
(Saporta , 1891;Sew ard , 1919;Jongmans und Dijk-
st ra , 1974).E .sotzkianus Unger is a widespread
species discovered f rom Tetrary st rata of France ,
Australia , Italy , Switzerand and Rumania of Europe
(Jongmans und Dijkst ra , 1974).In China there are
two species , Ephedrites sinensis Wu , He et Mai and
E .exhibens Wu , He et Mai (1986)collected f rom
the Lower Jurassic Xiaomeigou Fo rmation , Qinghai ,
West China.We have no oppo rtuni ty to examine all
species of Ephedrites identified formerly and check
whether they are entirely correctly identified or not.
Anyhow , the present specimens are most clos to mod-
ern Ephedra in their characters of stems and stro-
biles.They can be undoubtedly and aff irmatively at-
t ributed to Ephedri tes of Gymnospermae instead of
monoco tyledons of angiospermae.As stated above ,
we proposed to use the name Ephedri tes in Saporta s
(1891)sense and no t in the original sense of Goep-
pert and Benerendt (1845).Else M .Friis (personal
communication)suggests to give a new genus name
fo r such fossils.No mat ter w hat name should be used
fo r them , the resemblance between the present speci-
mens and living Ephedra is definitive.
Cao and Wu , S Q .et al ., w rongly compared
the present specimens with Cyperaceae of mono-
87第 1 期 郭双兴等:辽宁西部晚侏罗世晚期义县组的似麻黄属植物　　　　
co tyledon and assigned some specimens (in pl.1 ,
figs.1 —4)to a new genus and species Liaox ia chenii
Cao et Wu(1997 , p.1764—1765;1998 , p.231 , pl.
Ⅰ , figs.1 ,2 2a—c)of Cyperaceae.However , these
specimens are not similar to Cyperaceae at all , whose
stem is solid , thriquet rous , w ithout nodes and inter-
nodes.The family Cyperaceae consists of 4 000 living
species belonging to 70—90 genera (Hayw ood ,
1978 ;Cronquist , 1981).There are 31 surviving
genera and 670 species in China (How et al .,
1982).Over 10 fossil genera of this family have been
recorded.They first appeared in Palaeocene (LaM-
ot te , 1952).One common genus of this family is a
form genus Cyperaci tes (Cyperites).
The other specimens(in pl.Ⅰ , figs.5—7 ;pl.
Ⅱ , figs.3—8)were at tributed to another one new
genus and species Eragrosites changii Cao et Wu
(1997 , p .1765;1998 ,p .231—232;pl.Ⅱ , figs.1 —
3 ,1a , 2a—c;tex t-f ig ure 1)of Gramineae (Poaceae).
The specimens have a litt le analogy to Gramineae just
for their stem w ith nodes and internodes , but in other
characters , they are all dif ferent from Gramineae.
The stems of Gramineae are alw ays hollow .The lat-
eral branches of Gramineae are asymmetrical , alw ays
w ith twig s of unequal leng th , and not opposite.The
leaves are mainly lanceolate , alternate and dist ichous
w ith sheath and lingule.The f lowers are w ith 2—3
not opposite bracts. These main characters of
Gramineae are not seen in the present specimens.So
these fossil specimens should not be at t ributed to
Gramineae.Gramineae (Poaceae)is one of largest
family in monocoty ledons consisting of 500 —660 liv-
ing genera and 8 000—10 000 species (Hayw ood ,
1978 ;Cronquist , 1981;How and others , 1982).
Over 20 fossil genera of this family have been record-
ed.Acco rding to Cronquist (1981), the earliest fossil
record of Gramineae w as found from Senonian stage
of Late Cretaceous.Since Eocene on this family g rad-
ually became f lourished.
In addition , some fossil specimens (in pl.I ,
figs.8 —10) were assigned to monocoty ledonous
leaves by Cao and Wu (1997 , p.1765;1998 , p.232 ,
pl.Ⅰ , f ig s.3 ,4 ,4a).In fact , they are of conifer and
might belong to Podozamites.
Duan Shuy ing (1998)has also got a specimen
from the same formation and same locality.She also
considered it to be an oldest angiosperm in the wo rld
and gave a new genus and species Chaoyangia liangi
Duan for it.Her specimen might also be not related
to angiosperm.It may be a taxon of Chlamydosper-
mopsida(Gnetopsida).
A new ephedroid plant species , Leongathia ele-
gans discovered recently f rom the low er Aptian Koon-
warra fossil Bed in Victoria of Aust ralia w as studied
by Krassilov , Dilcher and Douglas(1998).The Aus-
t ralian species is characterized by slender shoots , lon-
gitudinally ribbed stems , linear leaves , four-leaved ,
occasionally tw o-three leaved , whorled arrangement
on nodes , with reduced sheaths.These characters are
superficially similar to the present fossil specimens ,
but they are much different f rom the present speci-
mens in the st robiles.
Miki(1964 ,p.13—22)has also described a new
genus and species Amphiephedra rhamnoides based
on a specimen found from the Lecoptera Bed in South
M anchuria(Northeast China).The Lecoptera Bed is
actually the same st ratum of the present fossil plants.
Miki considered that his fossil plant bears a resem-
blance to Ephedra .However , Amphiephedra rham-
noides is too bad in preservation without enough good
characters of stems , leaves and st robiles to erect a
new genus and species.
Yabe and Endo (1935 , p.274—276 , pl.1)have
ever described tw o species , Potamogeton jeholensis
Yabe et Endo and Potamogeton ? sp.collected f rom
the Lower Cretaceous ? Lycoptera Bed in Lingyuan
county of Jehol (now Liaoning), Northeast China.
Both species may be collected f rom the same st rata
and same locality of the present specimens.They are
litt le bi t similar to the present specimens.They are
perhaps relevant to Ephedrites , but they w ere trans-
ferred to Ranunculus jeholensis (Yabe et Endo)by
Miki(1964 ,p .19).
Velanovsky and Viniklar (1926)created a new
genus and species Ephedropsis strobi li fera based on
fossil plants f rom Cretaceous strata in Czech , but this
species has been at tributed to Taxodiaceae(Andrew s ,
1970).
Piroconi tes kuespert ii Go than found from the
low er Jurassic of Germany is a ephedroid plant.It s
bracts are 9 —15 cm long and 4—5 cm w ide.The
bracts show numerous parallel , unforked veins (12—
15 per cm).Its microsporophy lls show a concave ,
crescent-shaped base.It srobolaceus scales are 3.5—
5.5 cm in length and 2.0—3.5 cm in width (Van
Konijenburg-van Ci ttert , 1992 ;Kirchner , 1992),
88 　　 　　　古　生　物　学　报 第 39卷
these characters are very dif ferent from the present
specimens.
Crane(1987 ,1996)has indicated that Ephedra-
like pollen fossils first became common during the
Triassic and are widely distributed in Northern Hemi-
sphere , and has been recorded from the Lower Trias-
sic in North China(Ouyang and Norris , 1988).Dur-
ing the Jurassic , Ephedra-like pollen fossils are less
common.However , the pollen record still extended
into Cretaceous. During the Mid-Cretaceous ,
ephedroid pollen became more abundant in low er lati-
tude areas.Recently , Chinese palynologists Li Wen-
ben , Shang Yuke and Liu Zhaosheng told us that fos-
sil Ephedra-like pollen have been found from the
Jurassic Yixian Formation in Liaoning and from the
Jurassic strata in o ther areas of China.
＊　　＊　　＊　　＊　　＊　　＊
Acknowledgements:In preparation of this paper ,
Professor Zhou Zhiy an of our Institute and Professo r
Zhang Yonglu of Nanjing University have discussed
w ith the authors about the taxonomy and nomencla-
ture of the fossils and have of fered some valuable sug-
gestions.When this paper nearly be finished , Profes-
sor Else M arie Friis , the Sw edish Museum of Natural
Histo ry came to China and visited our Institute.She
had a close inspection on the present fossil specimens
and discussed w ith us about them.She has presented
her contributions , copied relevant papers and offered
useful suggestions.This study w as supported by the
g rant of Director of Nanjing Inst itute of Geology and
Palaeontology , Chinese Academy of Sciences and by
the g rant 9390010 of National Natural Science Foun-
dation of China.We would like to thank all of them
sincerely.
图版说明(Plate Explanation)
所有标本均保存在中国科学院南京地质古生物研究所。除注明
放大倍数者外 ,图片均代表标本的原大(All specimens are stored in
the collection house of Nanjing Institute of Geology and Palaeontology ,
Chinese Academy of Sciences.With the exception of the enlarged pho-
tographs , other specimens are natural size)。
图 版 Ⅰ(Plate Ⅰ)
1—7.陈氏似麻黄 Ephedr ites chen ii (C ao et Wu)Guo et Wu X.W.
1.标本原大(Natural size)。登记号:PB 17800 , Holotype。
2.系图 1放大 , ×3 ,示图 1下部茎节上的线形叶(Enlarged from
f igure 1 , ×3 , show ing the linear leaf on the lower part of spec-
imen)。
3.系图 1 放大 , ×3 示枝中部茎节 , 节间和球花序(Enlarged
f rom f igure 1 , × 3 , show ing the middle part of branches ,
nodes , internodes and st robiles)。
4.系图 1放大 , ×2 , 示枝顶球花序(Enlarged f rom f igu re 1 , ×
2, showing the st robiles of apical branches)。
5.标本原大(Natural size),正面。登记号:PB17802。
6.系图 5放大 , ×2 , 示枝和球花序(Enlarged f rom f igu re 5 , ×
2, showing the branches and st robiles)。
7.系图版Ⅱ , 图 6中间标本放大, ×2(Enlarged from plate Ⅱ
f irure 6 in the middle part of specimen , ×2)。
8—10.苏铁杉 ? (未定种)(Podozami tes? sp.)
8.标本原大(Natural size),正面。登记号:PB 117805。
9.系图 8放大 , ×3 ,示枝叶(Enlarged f rom figure 8 , ×3 , show-
ing branches and leaves)。
10.系图 8标本原大 , 反面(The counterpart of figure 8 , natu ral
size)。登记号:PB 17806。
图 版 Ⅱ(Plate Ⅱ)
1—8.陈氏似麻黄 Ephedr ites chen ii (C ao et Wu)Guo et Wu X.W.
1.标本原大(PB J31)(Natural size)。登记号:PBJ31。
2.系图 1放大 , ×2 ,示茎枝 , 节和节间(Enlarged f rom f igu re 1 ,
×2., show ing b ranches , nodes and internodes)。
3.标本原大(Natural size)。登记号:PB 17801。
4.系图 3放大 , ×3 , 示枝和球花序(Enlarged f rom f igu re 3 , ×
3, show ing branches and st robiles)。
5.标本原大(Natural size)。登记号:PB 17802。
6.标本原大(Natural size)。登记号:PB 17804。
7.标本原大 ,右侧(Natural size , right part)。登记号:PB 17803。
8.标本原大 ,左侧(Natural size , left part)。登记号;PB 17803。
89第 1 期 郭双兴等:辽宁西部晚侏罗世晚期义县组的似麻黄属植物　　　　


Chaoboridae .Its st range wing venation should be w rong
drawn.Both the two , Paratendipedidae and Sinotendi-
pedidae , are based on short-horned flies rather than repre-
sentatives of Chironomoidea.Zhang and other(1993)in-
dicated that Protabanidae is erected based on a cicada , not
a short-horned fly .It should be pointed out that there
have still been numerous so-called new families and new
genera established based on this sort of “ standard” in
Hong s papers and books besides these taxa mentioned
above .This author will give comments regarding them in
other articles.
The genus Sunaphis Hong et Wang , 1990 from the
Laiyang Formation in Laiyang of Shandong province ,
China has been placed in the extant f amily Aphididae.
Owing to possessing a close similarity in the main charac-
teristics , especially in the pattern of antenna (secondary
rhinaria ranging irregularly)to those of Sinaphididae , it
can now be t ransferred to Sinaphididae.
They have been thrown into self-contradictory over
the descriptions and text-figures about the Petiolaphis
Hong et Wang , 1990 and the Petiophioides Hong et
Wang , 1990 f rom the same locality like Sunaphis.This
author has failed to identify w hat the basic character actu-
ally is.For example , in description , Petiophis possesses
the antennal secondary rhinaria ranging irregularly ;but in
text-figure , it appears to be annular.Both genera do not
belong to the extant family Hormaphididae .Nevertheless ,
until further investigations of t he specimens are possible ,
they are of uncertain at familial status.
The species Expansaphis laticosta Hong et Wang ,
1990 was also derived f rom the identical location and st ra-
ta (Laiyang Formation)and placed in the genus Ex-
pansaphis Hong et Wang , 1990 within Oviparosiphidae.
There has been some confusion concerning its description
and text-figure.For instance , in the former the antennal
secondary rhinaria are of t ransverse arrangement , but in
the latter ranging irregularly.It is difficult to see f rom the
illustration how it might be related to the extinct family
Oviparosiphidae , let alone the genus Expansaphis.Its
taxonomic positionis uncertain not only at generic but at
familial levels before a reexamination of the type specimen
can be made .
Lin (1980)described a new genus Penaphis Lin ,
1980 from the Shouchang Formation in Zhejiang
province , China , which he considered to be a member of
the extant family Aphididae .Jarzembowski (1989) re-
garded Oviparosiphidae as a junior synonym of Callaphidi-
dae , and Penaphis could be transferred to Callaphididae.
Lin (1995)chimed in with Jarzembowski s (1989)opi-
nion.Hong (1998)agreed that Penaphis is a representa-
tive of Callaphididae.However , Zhang and others(1989)
recognized the mistaken classification and placed it in the
Oviparosiphidae.Ren (1995)placed Penaphis also into
this extinct family .Carpenter (1992), Ren(1995), An-
sorge (1996), and Heie and Wegierek (1998)admitted
the family Oviparosiphidae being well founded , respec-
tively .The present w riter believes that , with a great deal
of primeval characteristics , Oviparosiphidae is easily dis-
tinguished from Callaphididae:7-segmented antenna with
annlar , secondary rhinaria not only on the 3rd but also on
the following segments , in fore wing Rs straight and long
arising f rom middle of pterostigma and ending near wing
top , M arising f rom base of Pt , both CuA1 and CuA2 ori-
ginating independently f rom a commen stem Sc +R+M
but for a rather short distance each other , ovipositor large
whereas cauda and caudal plate absent or poorly deve-
loped.Meanw hile , Penaphis bears close resemblance in
forewing venation to those of Oviparosiphidae: the
straight and long Rs , arising f rom middle of Pt , M arising
f rom base of Pt , both CuA1 and CuA2 arising f rom almost
same point on Sc+R+M ;and then it may be placed in
Oviparosiphidae (see Text-figures 1 , 2).
Heie(1985)listed 26 plesiomorphous characters for
reconstruction of a primitive aphid.Here are provided
some additional items which are most likely of primeval
features:the last segment of antenna normal , not subdi-
vided(processus terminalis wanting or poorly developed);
the antennal secondary rhinaria usually same or similar in
shape , size , and arrangement on the 3rd and the following
segments;in forewing Pt laying near middle of wing ;Rs
elongated and st raight , ending near wing top.
Lin (1995)and Hong (1998)respectively regarded
the Yixian Formation in Liaoning province to be Upper
Jurassic—Lower Cretaceous , and the Laiyang Formation
in Shandong province to be Lower Cretaceous and younger
than the former.But they did not provide any direct evi-
dence of stratigraphical dating .Recently , an important
and famous fossil bird , the Confuciusornis santus Hou et
al ., has been discovered from the above-mentioned st rata
(Hou , 1997);thus both should be the same in geological
age.In addition , a geog raphically widespread species of
fossil dragonfly , Aeschnidum heishankowense (Hong),
has already been recognized by the present author
(Zhang , in press), which exists simultaneously in the two
strata.It is related to the Aeschnidum densum Hagen
f rom the Lower Tithonian of Solhofen , Germany , and
thus the Yixian and Laiyang formations can be regarded as
the Middle-Upper Tithonian deposits.
150 　　　 　　古　生　物　学　报 第 39卷