全 文 ：植 物 分 类 学 报 44 (2): 197–203(2006) doi: 10.1360/aps040120
Acta Phytotaxonomica Sinica http: //www.plantsystematics.com
———————————
2005-02-03收稿, 2005-05-03收修改稿。
基金项目: 国家自然科学基金(40402001); 中国科学院知识创新方向性项目(KSCX2-SW-108) (Supported by the National
Natural Science Foundation of China, Grant No. 40402001, and the Pilot Project of Knowledge Innovation, the Chinese
Academy of Sciences, Grant No. KSCX2-SW-108)。
* E-mail: happyking@ibcas.ac.cn; Tel.: 010-62836478.
山东中新世山旺植物群中豆荚属的名实问题
王 祺*
(系统与进化植物学国家重点实验室, 中国科学院植物研究所 北京 100093)
On the identity of Podogonium Heer 1857, nom. illeg.
(Leguminosae) from the Miocene Shanwang
flora of Shandong
WANG Qi*
(State Key Laboratory of Systematic and Evolutionary Botany, Institute of Botany, the Chinese Academy of Sciences,
Beijing 100093, China)
Abstract The nomenclature and taxonomy of Podogonium Heer 1857, nom. illeg., a legume
fossil widely distributed in the Neogene of Eurasia, were briefly reviewed. Herendeen’s
viewpoint that Podogonium Heer 1857 is a later synonym of Podocarpium A. Braun ex
Stizenberger 1851 is supported. The genus possibly contains only one species, Podocarpium
podocarpum (A. Braun) Herendeen 1992. Podogonium Heer 1857 should be no longer used
as a conserved name. The specimens from the Miocene Shanwang flora of Shandong, which
had been previously referred to Podogonium oehningense (Koenig) Kirchh. and P. knorrii
(Braun) Heer, were re-investigated. The results suggest that these specimens are the same
plant as the legume fossil from European Podocarpium podocarpum (A. Braun) Herendeen.
Key words Leguminosae, Podogonium Heer 1857, Podocarpium A. Braun ex Stizenberger
1851, Podocarpium podocarpum (A. Braun) Herendeen 1992, Shanwang flora, Miocene.
摘要 简要回顾了欧亚新近系广泛分布的豆科化石植物豆荚属Podogonium Heer 1857, nom. illeg.的命
名和分类。作者基本赞同Herendeen 1992年的观点, 即Podogonium Heer 1857是Podocarpium A. Braun ex
Stizenberger 1851一个晚出的异名。该属可能仅含有一个种——柄豆荚Podocarpium podocarpum (A.
Braun) Herendeen。同时, Podogonium Heer 1857也不宜作为保留名继续使用。作者重新调查了山东中新
世山旺植物群中过去被鉴定为Podogonium oehningense (Koenig) Kirchh.和P. knorrii (Braun) Heer的标本,
认为它们与欧洲发现的豆荚属应该为同一种植物, 即柄豆荚。
关键词 豆科; 豆荚属; 柄豆荚; 山旺植物群; 中新世
豆荚属Podogonium是Heer(1857)基于德国南部奥宁根(Oehningen)地区中新世地层发
现的豆科Leguminosae植物的花、果、种子以及叶化石而建立的, 以后广泛发现于欧亚新
近系, 尤其是中新世的植物群(Hantke, 1954; Bůžek, 1971; WGCPC, 1978; Herendeen,
1992a, b; Guo & Zhou, 1992; 刘裕生, 郑亚惠, 1995; Liu et al., 2001; Kovar-Eder et al.,
2004)。该属在我国的山东、山西、江苏、浙江、安徽、广西、青海、四川和云南等地的
新近系地层亦被广泛报道(Hu & Chaney, 1940; WGCPC, 1978; 郭双兴, 1980; 李浩敏,
植 物 分 类 学 报 44卷 198
1981; 李浩敏等, 1987; Guo & Zhou, 1992; 刘裕生, 郑亚惠, 1995; 孙博, 1999; 陶君容,
2000)。过去, 古植物学家对这种化石植物的分类意见并不一致, 在许多不同的属名(如水
盾草属Cabomba Aublet、黄檀属Dalbergia L.、皂荚属Gleditsia L.、似豆属Leguminosites
Bowerbank、Podogonium Heer和Podocarpium A. Braun ex Stizenberger等)之下对其进行了
描述(Stizenberger, 1851; Heer, 1857; Schimper, 1874; Chaney, 1933; Hantke, 1954; Bůžek,
1971; Gregor & Hantke, 1980; Herendeen, 1992a)。Herendeen(1992b)在详细讨论了豆荚属
的命名和分类后, 指出该属的正确名称应该是Podocarpium A. Braun ex Stizenberger 1851,
仅含有一个种, 即Podocarpium podocarpum (A. Braun) Herendeen。作者认为Herendeen对
豆荚属的分类处理是比较正确的, 但该属是否为单种属尚不能完全确定。我国近来有关
豆荚属的古植物学文献中并没有考虑Herendeen(1992b)的意见, 仍然沿用了过去的属名
Podogonium(刘裕生, 郑亚惠, 1995; 孙博, 1999; 陶君容, 2000), 属下种名也使用得比较
混乱。最近, 作者在研究山东临朐县山旺组(中中新世)的一些豆科植物化石时, 发现其中
的一些标本被命名为Podogonium oehningense (Koenig) Kirchh.和P. knorrii A. Braun。通过
仔细比较, 作者认为这些标本应归为一个种, 而且它们与欧洲发现的豆荚属为同一种植
物, 即柄豆荚(或单籽豆)Podocarpium podocarpum (A. Braun) Herendeen 1992。结合
Herendeen(1992b)的意见和Liu等(2001)的最新发现, 本文讨论了山东山旺植物群中豆荚
属的名实问题, 并重新描述了这些标本。
1 名实问题
欧洲的豆荚属最初根据采自德国南部奥宁根地区的果化石鉴定为睡莲科
Nymphaeaceae水盾草属的种Cabomba oehningense Koenig (Kirchheimer, 1957), Braun
(1851)将这种化石归入一个新属Podocarpium(Stizenberger, 1851), Koväts(1856)又将其归
入豆科的Copaifera L.属。Heer(1857)则认为这些化石与Copaifera不同, 后者的果柄较短,
且二者小叶叶脉相差甚远。此外, Heer(1857)也没有采纳Braun的新属Podocarpium, 由于
这个名字与罗汉松科Podocarpaceae罗汉松属Podocarpus L Hér. ex Pers.的拼写太相似了,
他就另建了一个新属Podogonium (Herendeen, 1992a, b)。根据命名优先律原则 ,
Podogonium是一个不合法的名字, 应该是Podocarpium的晚出异名, 但是, 一些作者考虑
到Podogonium在发表后已被使用了上百年, 建议或同意将它作为保留名(Bůžek, 1971;
李浩敏等, 1987)。然而, Herendeen(1992a, b)在详细讨论了豆荚属的命名和分类问题后发
现, 前面提及的Cabomba oehningense Koenig没有明确的发表日期, 因而不是一个有效发
表的名字。同时, Braun(1851)描述Podocarpium knorrii所依据的标本保存地点未知, 因此
也是一个不合法的名字。这样一来, 即使将Podogonium作为保留名继续使用, 其属下常
用的2个种名(即Podogonium oehningense和P. knorrii)也是不合法的, 因为用于描述它们
的模式标本都无从查证。因此, 保留Podogonium这个名字并不能使这些化石的命名保持
稳定。由于Gregor和Hantke(1980)在将Podogonium Heer归入现生皂荚属时, 他们从Heer
的原始材料中为Gleditsia knorrii指定了一个后选模式(lectotype)(即Podogonium knorrii
(Braun, 1851) Heer, 1857: pl. 135, fig. 21; 标本号: PL398=B803), 这个标本(为果化石)与
另外一个由Braun(1845)命名为Gleditschia podocarpa的标本实际上是相同的材料(注:
2期 王 祺: 山东中新世山旺植物群中豆荚属的名实问题 199
Gleditschia=Gleditsia), 但是根据皂荚属与豆荚属的果和叶结构特征的明显差异 ,
Herendeen(1992a)认为这个标本不属于皂荚属 , 而应归入豆荚属。因此 , Gleditschia
podocarpa Braun和Podocarpium knorrii Braun应该是同物异名。Heer(1857)特地将
Gleditschia podocarpa和Podocarpium knorrii都作为Podogonium knorrii的异名处理, 而描
述这些分类单位的材料都是来自同一产地, 即奥宁根地区。于是, Herendeen(1992b)重新
组合了一个名字Podocarpium podocarpum, 并提议将Gregor和Hantke(1980)给Gleditsia
knorrii的后选模式重新指定为Podocarpium的新模式(neotype)。这样既遵循了国际植物命
名法规中命名的优先律, 又确保了该植物化石的合格发表。
值得一提的是, Podocarpium (Benth.) Yang & Huang这个名称曾被用作豆科长柄山蚂
蝗属的属名(杨衔晋, 黄普华, 1979, 1995), 其模式为Podocarpium podocarpum (DC.) Yang
& Huang 1979。近来, Ohashi和Mill(2000)重新讨论了长柄山蚂蝗属的命名和分类, 指出
Podocarpium (Benth.) Yang & Huang与两个更早的名称(即Podocarpium A. Braun ex
Stizenberger 1851和Podocarpium Unger 1864)是异物同名, 于是他们为长柄山蚂蝗属建议
了一个新的属名Hylodesmum H. Ohashi & R. R. Mill。同时, Ohashi和Mill (2000)指出
Podocarpium podocarpum (A. Braun) Herendeen是Podocarpium podocarpum (DC.) Yang &
Huang一个晚出的双重异物同名(double-homonym), 考虑到后者为不合格发表的名称, 他
们认为可以保留前者, 但是提出保留Podocarpium Unger可能比 Podocarpium A. Braun ex
Stizenberger更好。然而, Podocarpium Unger已用于描述裸子植物罗汉松科的木材化石
(Herendeen, 1992a), 所以笔者认为最好还是保留Podocarpium A. Braun ex Stizenberger。
山东山旺植物群中豆荚属的标本包括叶和果(图1: A–D)以及相关的种子化石, 经常
被鉴定为Podogonium oehningense (Koenig) Kirchh.(WGCPC, 1978; 孙博 , 1999)和P.
knorrii (Braun) Heer (Hu & Chaney, 1940; 陶君容, 2000)。综上所述, 这些名字都是不合法
的, 应该被废弃。通过比较, 作者认为山旺植物群中豆荚属的标本应该为同一种植物, 它
们与欧洲发现的同属化石(Bůžek, 1971; Herendeen, 1992a)没有本质区别, 产出地层(距今
1500–1700万年)(李浩敏, 1981; Yang & Yang, 1994)也可以与欧洲同期地层横向比较。因
此, 山东山旺植物群中过去鉴定为Podogonium oehningense (Koenig) Kirchh.和P. knorrii
A. Braun的标本应该被归为一个种, 即柄豆荚(或单籽豆)Podocarpium podocarpum (A.
Braun) Herendeen 1992。需要说明的是, 中国其他产地报道(郭双兴, 1980; 李浩敏等,
1987; Guo & Zhou, 1992; 陶君容, 2000)的豆荚属标本也应归入这个种。
许多古植物学家都认为豆荚属化石同豆科植物有亲缘关系。Herendeen(1992a)基于果
和叶结构的比较形态学研究认为, 豆荚属可能是一个与豆科云实亚科Caesalpinioideae的
Detarieae或Amherstieae族中的几个属有密切亲缘关系的绝灭类群。然而, 基于相关的花粉
(Mohr & Gregor, 1984)和该属可能的花化石证据, Liu等(2001)则认为豆荚属作为一种绝灭
的豆科植物与云实亚科中现存属的差别较大, 它在豆科系统发育中可能处于靠近基部的位
置。

植 物 分 类 学 报 44卷 200
2 豆荚属的重新描述
豆荚属(豆科Leguminosae)
Podocarpium A. Braun ex Stizenberger 1851 in Ueber. Verstein. Groß. Baden: 90. 1851;
Herendeen in Taxon 41: 732. 1992; Y. S. Liu et al. in Palaeont. Abt. B 256: 162. 2001, non
Podocarpium Unger in Hochstetter et al., Paläont. Neu-Seeland: 13, pl. 5, fig. 1a–c. 1865.
Type: P. dactyloides Unger (post Nov. 1864, fossil Podocarpaceae), nec Podocarpium
(Benth.) Yang & Huang in Bull. Bot. Lab. North-East. Forest. Inst. 4 (4): 4. 1979; et in Fl.
Reip. Pop. Sin. 41: 47. 1995. nom. illeg. Type: Desmodium podocarpum DC.=Podocarpium
podocarpum (DC.) Yang & Yang.
Podogonium Heer in Tert. Fl. Schweiz, Fl. Tert. Helvet. 3 (6): 113. 1857. Type:
Podocarpium knorrii A. Braun ex Stizenberger, nom. illeg. (repository unknown).
柄豆荚(单籽豆) 图1: A–D
Podocarpium podocarpum (A. Braun) Herendeen in Taxon 41: 732. 1992; Y. S. Liu et
al. in Palaeont. Abt. B 256: 162. 2001; Kovar-Eder et al. in Ann. Naturhist. Mus. Wien 105 A:
74. 2004, non Podocarpium podocarpum (DC.) Yang & Huang in Bull. Bot. Lab. North-East.
Forest. Inst. 4 (4): 7. 1979; et in Fl. Reip. Pop. Sin. 41: 52, pl. 9, figs. 1–5. 1995, nom. illeg.
(=Hylodesmum podocarpum (DC.) H. Ohashi & R. R. Mill in Edinb. J. Bot. 57 (2): 181.
2000). ——Gleditschia podocarpa A. Braun in Neues Jahrb. Mineral. Geog. Geol.
Petr.-Kunde: 173. 1845. ——Dalbergia podocarpa (Braun) Unger in K. K. Akad. Wiss.
Denkschr. Math.-Nat. Cl. 2: 185, pl. 61, fig. 14. 1851.
Podocarpium knorrii Braun ex Stizenberger in Ueber. Verstein. Groβ. Baden: 90. 1851,
nom. illeg. ——Podogonium knorrii (Braun) Heer in Tert. Fl. Schweiz, Fl. Tert. Helvet. 3 (6):
114, pl. 134, figs. 22–26a, pl. 135, non fig. 19, pl. 136, figs. 1–9. 1857, nom. illeg.; Schimper
in Trait. Paléont. Vég.: 43, pl. 100, figs. 29–34. 1874; Hu & Chaney in Palaeont. Sin., N. S. A:
1, 76, pl. 50, figs. 4, 5, 7, 10, 11. 1940; Herendeen in Herendeen & Dilcher, Adv. Legum. Syst.
4 Fos. Rec.: 4, figs. 1–5, 12–20. 1992a; J. R. Tao in Evol. Late Cret.-Cenoz. Fl. China: 267,
figs. 18, 19. 2000; Gleditsia knorrii (Heer) Gregor in Gregor & Hantke, Feddes Repert. 91:
166, pl. 10, figs. 9–12, pl. 11, figs. 2–7, pl. 12, figs. 1–15. 1980, Mohr & Gregor in Paläont. Z.
58: 191. 1984.
Leguminosites climensis Chaney in Bull. Geol. Soc. China 12 (2): 134, pl. 1, fig. 7. 1933;
WGCPC in Cenoz. Pl. China, Fos. Pl. China 3: 113, pl. 90, fig. 2. 1978.
Podogonium latifolium Heer in Tert. Fl. Schweiz, Fl. Tert. Helvet. 3 (6): 116, pl. 136, figs.
10–21. 1857.
Podogonium constrictum Heer in Tert. Fl. Schweiz, Fl. Tert. Helvet. 3 (6): 118, pl. 136,
fig. 48. 1857.
Podogonium campylocarpum Heer in Tert. Fl. Schweiz, Fl. Tert. Helvet. 3 (6): 118, pl.
136, figs. 54, 55. 1857.
Podogonium obtusifolium Heer in Tert. Fl. Schweiz, Fl. Tert. Helvet. 3 (6): 118, pl. 134,
figs. 30–34. 1857.
Podogonium lyellianum Heer in Tert. Fl. Schweiz, Fl. Tert. Helvet. 3 (6): 117, pl. 136,
figs. 22–52. 1857; Hantke in Denkschr. Schweiz. Naturforsch. Gesellsch. 80: 73, pl. 12, figs.
11–15. 1954; Copaifera longestipata Kováts in Arbeiten Geol. Ges. Ungarn 1: 51, pl. 1, figs.
3, 4. 1856; Gleditsia lyelliana (Heer) Hantke in Gregor & Hantke, Feddes Repert. 91: 167.
1980.
Podogonium oehningense (Koenig) Kirchheimer in Laubgew. Braunk.: 412, pl. 20, fig.
90a, b, 1957; Bůžek in Rozpr. Ústr. Ústav. Geol. 36: 98, pl. 48, fig. 9, pl. 50, figs. 1–27, pl. 51,
figs. 1–12. 1971; WGCPC in Cenoz. Pl. China, Fos. Pl. China 3: 112, pl. 88, fig. 5, pl. 90, fig.
2, pl. 93, figs. 4, 5, 7, non pl. 88, fig. 7, pl. 89, fig. 7. 1978; S. X. Guo in Acta Palaeont. Sin.
19: 408, pl. 1, fig. 9. 1980; H. M. Li in Acta Palaeont. Sin. 26: 572, pl. 2, figs. 3, 3a, 4, 8, 8a,
2期 王 祺: 山东中新世山旺植物群中豆荚属的名实问题 201

图1 柄豆荚的叶和果 A. 一片不完整的偶数羽状复叶。标本号: 50483a。B. 一片完整的小叶。标本号: 52463。C.
一枚开裂的果。标本号: 51868。D. 一枚未开裂的果。标本号: 52913。标尺=1 cm。
Fig. 1. Leaf and fruit of Podocarpium podocarpum (A. Braun) Herendeen 1992. A, An incomplete paripinnate leaf.
Specimen number: 50483a. B, A complete leaflet. Specimen number: 52463. C, A dehiscent fruit. Specimen number:
51868. D, An indehiscent fruit. Specimen number: 52913. Scale bar=1 cm.
植 物 分 类 学 报 44卷 202
10–12, 15–19, 15a, 17a, 18a, 19a. 1987; B. Sun in Shanwang Pl. Fos. 42, 71, pl. 28, fig. 5.
1999.
Neotype: PL398=B803 (Podogonium knorrii (Braun, 1851) Heer in Tert. Fl. Schweiz, Fl.
Tert. Helvet. 3 (6): 114, pl. 135, fig. 21. 1857). Repository: Geological Institute, Eid-
genossische Technische Hochschule Zürich (ETH). Type locality: Oehningen, southern
Germany. Age: Miocene.Stratigraphic range and Geographic distribution: early Oligocene to
the Pliocene in Eurasia. Specimens examined herein: 50483a, 50483b, 50483c, 52463, 50508a,
50489a, 50489b, and 52058c (leaves); 2860, 52913, 51706, 52058a, 201a, 51868, 51989,
1569, 51721, and 50373 (fruits) (PE!). Locality: Shanwang Basin, Linqu County, Shandong
Province, China (36°54′ N, 118°21′ E). Age: middle Miocene.
Leaves paripinnate with at least 5 pairs of opposite, rarely subopposite leaflets, up to 7.5
cm long. Leaflets sessile to short-petioluled (petiolule ca. 0.5 mm long). Lamina lanceolate,
ovate to oblong-ovate, 2.0–4.4 cm long, 0.6–1.1 cm wide, margin entire, apex obtuse or acute,
rarely mucronate or emarginate, base asymmetrically rounded or slightly narrowed. Midvein
stout, usually straight, 1–2 prominent basal veins arising near midvein base at acute angles and
extending parallel to leaflet margin. Secondary veins at 30°–70° angles, numerous, over 12
pairs, pinnately brochidodromous, forked near leaflet margin, forming angular loops.
Intersecondary veins parallel to secondary veins. Tertiary veins fine, dense, forming irregular
network. Fruits tardily dehiscent, 1-seeded, stalked; stalk incompletely preserved, 2–4 cm
long, about 1 mm wide. Seed oblong, 12–15 mm long, 8–10 mm wide. Fruit valve with
indistinct venation, elliptical, 1.8–2.8 cm long, 0.8–1.1 cm wide, apex acute, base attenuate,
slightly oblique, margin not winged.
偶数羽状复叶, 对生(稀近对生), 至少5对, 长达7.5 cm。小叶无柄或具短柄, 约0.5
mm长。叶片呈披针形、卵形或长圆状卵形, 长2–4.4 cm, 宽0.6–1.1 cm, 全缘, 顶端钝或
急尖, 稀突尖或凹缺, 基部不对称圆形或稍狭。中脉较粗, 一般直, 有1–2条明显的基脉自
基部以锐角从中脉延伸靠近叶缘平行朝向叶顶。二次脉多对, 12对以上, 羽状弓形脉, 与
中脉的夹角呈30°–70°, 近叶缘内分叉, 环结。间二次脉与二次脉平行。三次脉细密, 彼
此构成不规则的脉网。荚果迟裂, 内仅含1粒种子, 果梗保存不完整, 2–4 cm长, 约1 mm
宽。种子长圆形, 12–15 mm长, 8–10 mm宽。果瓣椭圆形, 1.8–2.8 cm长, 0.8–1.1 cm宽, 表
面纹饰不明显。果顶部急尖, 基部渐狭, 微偏斜, 边缘无翅。
致谢 感谢德国Hans-Joachim Gregor、Johanna Kovar-Eder、洪堡大学自然历史博物馆
Barbara Mohr、美国威斯康星大学生物系刘裕生、中国科学院南京地质古生物研究所徐
洪河、中国科学院植物研究所朱相云和孙启高诸博士和陶君容教授以及北京华能电子仪
器厂李增新同志提供部分重要文献, 植物研究所李荣贵先生协助打印图片！
参 考 文 献
Braun A. 1845. Die Tertiär-flora von Oehningen. Neues Jahrbuch für Mineralogie, Geognosie, Geologie, und
Petrefakten-Kunde 1845: 164–173.
Braun A. 1851. Uebersicht der geognostischen und allgemeinen palaeontologischen Verhältnisse Badens. In:
Stizenberger E ed. Uebersicht der Versteinerungen des Großherzogthums Baden. Freiburg: Verlag der
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