横断山区的翠雀属植物:分类学和细胞学

翠雀属Delphinium L.是毛茛科中的一个大属，全世界有300－400 余种，广布于北半球温带地区。我国是世界上记载翠雀属植物种类最多的国家，尤以西南横断山区种类最为丰富。 通过文献查阅、标本室研究和野外考察，本文对我国特别是横断山区翠雀属植物的主要形态性状在居群间和居群内的变异式样进行了初步分析，认为根的形态、叶的分裂程度、植株尤其是花序轴和花梗被毛与否以及毛被类型、小苞片的形状和位置、退化雄蕊的颜色、上萼片的形状、心皮数目和种子形态特征是比较可靠的分类性状。但性状之间的相关性往往较小，即使是通常比较稳定的分类性状有时也表现出不同程度的变异，因此同一性状对于不同等级分类群的划分和在不同分类群中的分类价值常不一样。在具体的分类处理中，应具体情况具体分析，尽可能利用较多的分类学性状或性状组合。 根据上述对我国翠雀属植物主要形态性状变异式样的理解，本文主要对我国西南横断山区及其邻近一些地区的翠雀属植物进行了力所能及的分类修订：将42 种和22 变种降为异名，提出2 新组合(毛梗黑水翠雀D. potaninii var. mollipes　(W. T. Wang) Q. Yuan & Q. E. Yang 和腺毛康定翠雀D. tatsienense var. pseudomosoynense (W. T. Wang) Q. E. Yang & Q. Yuan)，澄清了直距翠雀D. tenii Lévl. 的名实问题和Munz 等学者对我国翠雀属植物标本的鉴定错误以及由此引起的我国有关翠雀属植物的地理分布的混乱; 另有两种( 硕片翠雀D. grandilimbum W. T. Wang & M. J. Warnock 和永宁翠雀D. yongningense W. T. Wang & M. J. Warnock)因未见到模式标本，故暂存疑。本文讨论到的我国翠雀属植物共有87 种、10 变种。 为了在染色体水平上了解横断山区翠雀属植物的物种形成和分化，同时增加对整个翠雀族tribe Delphineae Warming 植物染色体进化的理解，本文报道了我国主要采自该地区的乌头属Aconitum L. 30 种、1 变种共40 个居群和翠雀属 48 种、3 变种共87 个居群的染色体数目，分析了30 种、1 变种乌头属植物和 46 种、3 变种翠雀属植物的核型，发现46 种、3 变种翠雀属植物为二倍体(2n = 2x=16)，1 种(黄毛翠雀D. chrysotrichum Finet & Gagnep.)为四倍体(2n = 4x =32)，1种(螺距翠雀D. spirocentrum Hand.-Mazz.)有二倍体和四倍体两种细胞型，3 种(宽距翠雀D. beesianum W. W. Smith、裂瓣翠雀D. mosoynense Franch.和康定翠雀 D. tatsienense Franch.)的个别居群有B 染色体出现。这表明横断山区翠雀属植物的物种形成主要在二倍体水平上进行。可能由于核型直向选择（karyotypic orthoselection）的结果，我国翠雀属植物的核型就整体情况而言在属内相当一致，属内核型分化不如乌头属中明显，外部形态上显得最为原始的种类如短距翠雀 D. forrestii Diels 和毛翠雀D. trichophorum Franch.的核型与其他多年生种类的核型相比并无明显区别。尽管如此，翠雀属植物的核型在属内仍表现出一定程度的分化：在多年生种类中，大理翠雀群(D. taliense group)的种类的染色体内部不对称性(intrachromosomal asymmetry) 程度较高, 但染色体之间的不对称性 (interchromosomal asymmetry)程度偏低;一年生种类与多年生种类相比，我国唯一的一年生种还亮草D. anthriscifolium Hance 染色体之间不对称性最高，该种染色体内部不对称性程度在翠雀族中也偏高，因此就其核型的整体不对称性程度而言，在翠雀属乃至整个翠雀族中最高，而地中海分布的一年生种类的染色体内部不对称性明显增强, 但染色体之间的不对称性在翠雀族中最低。这种核型分化的系统学和生物地理学意义尚不清楚，以后需进一步研究。与乌头属相比，翠雀属植物的染色体通常较小，核型中st 染色体明显增多（可达6 对），核型的整体不对称性程度逐渐增强; 虽然乌头属牛扁亚属Aconitum subgen. Lycoctonum (DC.) Peterm.的二倍体种类也一般具有较多的st 染色体（4 对），但这些st 染色体比乌头亚属Aconitum L. subgen. Aconitum和露蕊乌头亚属Aconitum subgen. Gymnaconitum (Stapf) Rapaics 植物核型中相对应的m 或 sm 染色体以及翠雀属和飞燕草属Consolida (DC.) S. F. Gray 植物中相对应的st 或偶尔出现的sm 染色体要大。特别应当注意的是牛扁亚属中外部形态上显得最为原始的展喙乌头A. novoluridum Munz 的核型具有较多而且较大的sm 染色体，所以我们认为展喙乌头的核型保留了较多的原始性质，那些核型中具有较多st 染色体的牛扁亚属植物应当是该亚属中在核型上已比较特化的种类。这一结果不支持前人提出的乌头属植物的核型有从st 染色体进化到m 或sm 染色体的趋势的观点。就整个翠雀族而言，从现有资料看，其核型的进化趋势应当为：(1) 染色体从大进化到小, 从m 或sm 染色体进化到st 染色体;(2) 核型的整体不对称性程度逐渐增强，即核型的二型性从多年生的牛扁亚属植物到翠雀属植物的一年生的还亮草亚属Delphinium subgen. Delphinium 而愈趋明显。翠雀族植物核型的进化方向与整个毛茛科核型的进化方向看来是一致的，即都向核型不对称性增强的方向进化。

毛茛目的系统发育分析：分子和形态证据

本研究选取毛茛目99属，利用四个叶绿体和核基因或DNA片段，并结合形态性状，在属的水平上构建毛茛目的系统发育关系。分子数据用简约法和似然法分析，分子和形态性状联合数据用简约法分析。具体结果如下： 1) 毛茛目的单系性 毛茛目的单系性在所有的分析中都得到强支持。这也被形态性状所支持，包括大的S-型筛管分子、表皮层具蜡质的管、离生并折合状发生的心皮、两层珠被形成的珠孔、无托叶、轮状花被(花被在领春木科可能是次生性丢失)和2-3层细胞组成的内珠被等。 2) 科间系统发育关系 毛茛目由三个主要分支构成：领春木科、罂粟科和核心毛茛目。罂粟科是核心毛茛目的姊妹群，二者通常具复叶、单叶分裂或单叶掌状脉，雄蕊轮状排列以及胚珠无丹宁酸组织。核心毛茛目的单系性得到强支持，花3基数、雄蕊和花被片对生以及外部花被片具三或更多的微管束迹等是其共有衍征。木通科和星叶草科形成一支，细胞型胚乳是它们的共有衍征。防己科、小檗科和毛茛科形成一支，核型胚乳和具小檗碱支持该支的单系性。小檗科和毛茛科的姊妹群关系得到草本习性、具根状茎、V-型木质部和外珠被至少四层细胞等性状的支持。 3) 科下系统发育关系 a. 罂粟科 罂粟科分为两个单系支：狭义罂粟科形成一支，Pteridophyllm、角茴香属和狭义荷包牡丹科形成另一支。是否具乳汁、小苞片状的萼片是否包着其他花部器官、雄蕊数目和花粉外壁纹饰等性状支持这种划分。这两支分别被给予亚科的分类等级。Pteridophyllm是角茴香属的姊妹群也得到分辨，叶和雄蕊的性状支持这两属近缘。 b. 星叶草科 独叶草属和星叶草属形成一支得到强支持，二者组成星叶草科。二叉状分支的叶脉、条纹状的花粉纹饰、单珠被、薄珠心和无外种皮等性状是该科的共有衍征。 c. 木通科 木通科的单系性得到强支持。大血藤属是其他木通科的姊妹群，因其在心皮数目、心皮排列、胚珠数目、胚珠方向、染色体数目和植物化学等方面不同于其他类群，建议给予亚科的分类等级。此外，猫儿屎属和串果藤属均具有一些独特的性状，分别给予族的等级。 d. 防己科 古山龙属和Anamirta形成一支，意味着族Anamirteae sensu Diels应该被承认;广义的青牛胆族应包括天仙藤族;无胚乳的粉绿藤族嵌合在蝙蝠葛族之内。子叶和花柱残基对科内的划分是非常重要的。虽然传统上认为胚乳是科下划分最重要的性状，但它在防己科内应该是次生性丢失。 e. 小檗科 六个间断分布的属或属对被证实。单心皮瓶状发生是小檗科的共有衍征。小檗科分为三个主要分支：南天竹属(x = 10)和染色体基数为8的类群形成强支持的一支，这也得到胚珠数目(2-4)和圆锥花序等性状的支持;x = 7的类群形成一支，花瓣基部内侧具2枚腺体，花粉具平滑的外壁和未分化的外表层等是其共有衍征;x = 6的类群形成一支，被条纹状的花粉外壁和不连续的花粉外壁内层所支持。这三支分别被给予亚科的分类等级。 f. 毛茛科的单系性得到强支持。雄蕊多数和胚珠多数是毛茛科的共有衍征。毛茛科由五个分支组成：Glaucidium、Hydrastis、染色体基数为9的类群形成一支、染色体基数为7的类群形成一支、染色体基数为8的类群形成一支，分别给予亚科的分类等级。Glaucidium是所有其他毛茛科的姊妹群。染色体基数为8的类群的单系性得到极弱的支持，不含有或者含微量的苄基异喹啉生物碱及具R-型染色体似乎也支持该支为单系。星果草属应隶属于毛茛亚科而与黄连属和(/或)人字果属关系遥远。在唐松草亚科，具瘦果的唐松草属嵌合在具蓇葖果的类群之内，意味着依据果实类型划分该亚科是不合适的。美花草属和毛茛族及银莲花族形成一支，而与侧金盏花属关系遥远。美花草属在胚珠着生位置、珠被数目和植物化学等方面不同与毛茛族和银莲花族，因此我们建议给予美花草属族的分类等级。 根据分析结果，我们提出了毛茛目一个新的分类系统

毛茛科铁线莲属长瓣铁线莲组的分类学修订

毛茛科（Ranunculaceae ）铁线莲属(Clematis L.)长瓣铁线莲组(Sect. Atragene) 植物全世界约有14－18 种，我国约有6 种，广泛分布于北温带的高山上。该组植物由于外部形态变异大，种间或种下划分十分困难，目前已发表的种及种下分类群名称多达130 余个，名称使用较为混乱，组内系统关系尚存争议。本文通过文献考证、标本研究、野外观察及形态性状统计分析的综合研究，对长瓣铁线莲组进行了全面的分类学修订，结果如下： 1、性状分析 深入研究了1000 余号该组植物标本的形态性状变异式样、范围及其相关性，发现复叶类型、小叶大小、萼片毛被和质地、萼片脉突出与否、退化雄蕊长度等是较为稳定的分种性状。生长习性、萼片颜色、萼片边缘是否具短绒毛带、退化雄蕊形状等对种下划分具有重要的意义。 2、分类学处理 1）根据该组植物的萼片和退化雄蕊的特征和演化趋势，建立了5 个新系：Ser. 1. Occidentales、Ser. 2. Alpinae、Ser. 3. Koreanae 、Ser.4. Tomentosae 和Ser. 5. Macropetalae 。 2）对长期以来分类上争议很大，且分类十分困难的类群作出以下新的处理：把C. sibirica 和C. ochotensis 作为C. alpina 的亚种；把日本的C. fusijamana 和C. fauriei 降级为半钟铁线莲C. alpina ssp. ochotensis 的变种；把伊犁铁线莲C. iliensis 降级为西伯利亚铁线莲C. alpina ssp. sibirica 的变种；把C. chiisanensis 降级为朝鲜铁线莲C. koreana 的变种；把紫红花长瓣铁线莲C. macropetala var. punicoflora 和石生长瓣铁线莲C. macropetala var. rupestris 处理为长瓣铁线莲C. macropetala 的异名；把C. dianae 归并入白色长瓣铁线莲C. macropetala var. albiflora；支持长白铁线莲C. nobilis 作为一个独立的种。此外，还新指定了5 个名称的后选模式；纠正了前人对中国C. subtriternata 的鉴定错误。 3）确认该组植物有9 种2 亚种和9 变种（包括3 新等级），给出了各个分类群详细的文献引证、描述、生境、地理分布、标本引证和外部形态图，并编写了组下分系和分种检索表。认为亚洲东北部是本组植物分布的多样性中心。 3、形态性状演化趋势和各系的亲缘关系 探讨了该组植物形态性状演化的趋势和新建立的5 个系的亲缘关系，提出：退化雄蕊可能由外部雄蕊演化而来，并沿两条路线演化发展：其一是退化雄蕊伸长，与萼片近等长或长于萼片，顶端渐狭，呈披针形；其二是退化雄蕊不伸长，与雄蕊近等长，上部加宽，顶端由钝形、圆形发展至微凹，呈匙形。萼片演化的趋势可能由质地薄到质地厚，萼片脉由不凸出到3 条脉凸出。由此推测北美洲类群Ser. Occidentales 可能是本组的原始类群；Ser. Macropetalae 可能是退化雄蕊在第一条演化路线上最进化的类群；Ser. Alpinae 和Ser. Koreanae 之间亲缘关系较近；Ser. Tomentosae 的系统位置尚未确定。

烟草NtFAD2基因的沉默及功能研究

油酰磷脂酰胆碱去饱和酶（FAD2）是一种重要的植物脂肪酸去饱和酶，位于内质网中，催化油酸生成亚油酸。亚油酸等多不饱和脂肪酸含量过高的植物油脂不稳定，易氧化，不易贮藏，而且氧化产物对人体有害。因此油脂改良的一个重要目的是提高种子油脂的油酸含量，降低多不饱和脂肪酸含量。另外，关于FAD2的结构、功能和表达调控等问题目前还不清楚，有待深入研究。近年发展起来的RNAi技术可有效地沉默目标基因，研究植物FAD2基因沉默后膜脂和油脂的变化及温度对基因沉默的影响，有助于深入理解FAD2酶的功能、表达调控方式和有效地利用基因沉默技术改变膜脂组成、提高植物油脂品质。本研究的主要目的是，首先采用RNAi技术抑制烟草FAD2基因（NtFAD2）的表达以降低NtFAD2酶活性，得到膜脂不饱和度改变的烟草转基因株系。然后研究NtFAD2基因沉默后脂肪酸去饱和过程的变化及其机理，以及温度对NtFAD2基因沉默的影响。研究中首先用PCR方法克隆烟草NtFAD2基因，用其编码区的一个片段构建表达发卡RNA的沉默结构，用农杆菌介导的方法转化烟草，从第一代转基因植株中筛选出油酸含量高的突变体。然后对其中一个转基因株系S61进行脂肪酸分析，发现叶片总脂和质体外单脂PC和PE中的油酸含量大幅度增加。此外，NtFAD2基因沉默对叶片甘油脂的影响有多效性，即质体内的单脂油酸含量也有显著增加，有些单脂中软脂酸含量有所下降。烟草NtFAD2基因沉默后，在叶片和种子中油酸含量变化最大，说明NtFAD2基因在不同器官中的沉默效果不同：沉默结构对NtFAD2酶活性较高的器官抑制程度更大，内源基因没有表现均一的沉默效果。沉默植株的多种组织中NtFAD2基因的mRNA降解程度相似，说明油酸含量与NtFAD2转录本的丰度没有直接关系。 植物脂肪酸的不饱和程度受温度调控，因此研究FAD2基因沉默株系的油酸含量受温度影响情况及其调控规律对RNAi技术在油脂改良方面的实际应用具有十分重要的意义。研究结果表明，随着生长温度的下降，野生烟草叶片膜脂油酸含量降低，多不饱和脂肪酸含量随之增加;转基因烟草中油酸和多不饱和脂肪酸有相似的变化趋势，但是变化幅度远远大于野生烟草。低温下转基因烟草叶片油酸含量之所以大幅度下降，其中一个重要原因是被抑制的NtFAD2酶受低温的调节而活性升高。然而研究发现，与常温条件下相比，低温下转基因烟草NtFAD2基因的mRNA丰度不变，说明低温没有影响沉默结构对NtFAD2基因的mRNA的降解。低温环境下NtFAD2基因沉默植株的多不饱和脂肪酸含量有所回升可能是植物对低温的一种适应性响应。转基因烟草种子油脂的油酸含量也受温度影响：常温下表现高油酸性状，低温下油酸含量下降。因此种植这种高油酸品系时，需要考虑温度的影响。如果在适宜的地域和季节种植，避免转基因植株开花结实时遭遇持续低温，就可以利用转基因植株的遗传优势，生产富含油酸的优质植物油。

青藏高原矮嵩草草甸种子库的初步研究

The seed bank of Kobresia humilis meadow in Qing- Zang Plateau was preliminaryly studied and the relationships among seed bank , seed rain and above-ground vegetation were compared and analysed.The results showed that there were 23 species in the seed bank in both of green up and withering stage. In species composition of the seed bank ,Ranunculaceae and Leguminosae were dominant ,while the percentage of Cyperaceae and Gramineae were lower. The percentage of K. humilis seeds was relatively lower with 0. 97 % in green up stage and 1. 97 % in withering stage. The species composition of the seed bank correlated with that of the seed rain ( r = 0. 7505 , P < 0. 01) ,and there were 18 common species which was 78. 3 %of species of the seed bank. Only 41. 81 %of above-ground vegetation species presented in the seed bank. The seed numbers of the dominant plant species in vegetation were smaller , with lower percentage in the seed bank. Therefore , the dominant vegetation of K. humilis et al. mainly depended on vegetative reproduction to maintain and strengthen. From vegetation to the seed bank in green up stage ,the species diversity decreased by 9. 94 %. It implied that the seed bank was the source of vegetation regeneration , but also a mechanism of preserving species diversity of the plants.

东南极格罗夫山土壤研究与古环境分析

Three soil spots were found in Grove Mountains, east Antarctica during 1999-2000, when the Chinare 16th Antarctic expedition teams entered the inland Antarctica. The characteristics of soils in Grove Mountains are desert pavement coating the surface, abundant water soluble salt, negligible organ matter, and severe rubification and salinization, scarces of liquid water, partly with dry permafrost, corresponding with the soils of McMurdo, Transantarctic. The soils age of Grove Mountains is 0.5-3.5Ma. Podzolization and redoximorphism are the main features in coastal Wilks region, in addition, there is strong enrichment of organic matter in many soils of this region. The main soil processes of Fildes Peninsula of King George Island include the intense physical weathering, decalcification and weakly biochemical processes. Peat accumulation is the main processes in Arctic because of humid and cold environment.Based on synthesis of heavy minerals, particle size, quartz grain surface textures, as well as pollen in soils, the soils parent materials of Grove Mountains derived from alluvial sediment of the weathering bedrocks around soils, and formed during the warm period of Pliocene. The detailed information is followed .l)The results of heavy minerals particle size showed the parent minerals derived form the weathering bedrocks around soils. 2)The quartz sand surface textures include glacial crushing and abrasion such as abrasive conchoidal fractures and grain edges, abrasive subparallel linear fractures and angularity, subaqueous environments produce V-shaped and irregular impact pits, polished surface, and chemical textures, such as beehive solution pits, which showed the water is the main force during the sediment of the soil parent minerals. 3)The pollen consist of 40 plant species, of which at least 5 species including Ranunculaceae, Chenopodiaceae, Artemisia, Gramineae, Podocarpus belong to the Neogene vegetation except the species from the old continent. Compared with Neogene vegetation of Transantarctic Mountains, Antarctic, we concluded that they grow in warm Pliocene.

Triterpenoids from Pulsatilla chinensis

A new lupane type triterpenic acid, pulsatillic acid, and two new lupane type triterpenoid glycosides, pulsatilloside A and B, along with the known 23-hydroxybetulinic acid were isolated from the roots of Pulsatilla chinensis. Their structures were characterized as 3-oxo-23-hydroxy-lup-20(29)-en-28-oic acid, 3 beta, 23-dihydroxy-lup-20(29)-en-28-oic acid 3-O-alpha-L-arabinopyranoside and 3 beta, 23-dihydroxy-lup-20(29)-en-28-oic acid 28-O-beta-D-glucopyranosyl-(1-->6)-beta-D-glucopyranoside on the basis of hydrolysis and spectral evidence including two-dimensional relay HOHAHA, one-dimensional multiple relay COSY and ROESY NMR techniques. Pulsatillic acid exhibited cytotoxic activities against P-388, Lewis lung carcinoma and human large-cell lung carcinoma.

PATENSIN, A SAPONIN FROM PULSATILLA PATENS VAR MULTIFIDA

Patensin, a new triterpenoid glycoside, was isolated from the ethanolic extraction of the roots of Pulsatilla patens var. multifida. Its structure was established as hederagenin 3-O-beta-D-galactopyranosyl-(1-->2)-beta-D-glucopyranoside on the basis of hydrolysis and spectral evidence including 1D and 2D NMR techniques.

Modern pollen samples from alpine vegetation on the Tibetan Plateau.

1  A set of 316 modern surface pollen samples, sampling all the alpine vegetation types that occur on the Tibetan Plateau, has been compiled and analysed. Between 82 and 92% of the pollen present in these samples is derived from only 28 major taxa. These 28 taxa include examples of both tree (AP) and herb (NAP) pollen types. 2  Most of the modern surface pollen samples accurately reflect the composition of the modern vegetation in the sampling region. However, airborne dust-trap pollen samples do not provide a reliable assessment of the modern vegetation. Dust-trap samples contain much higher percentages of tree pollen than non-dust-trap samples, and many of the taxa present are exotic. In the extremely windy environments of the Tibetan Plateau, contamination of dust-trap samples by long-distance transport of exotic pollen is a serious problem. 3  The most characteristic vegetation types present on the Tibetan Plateau are alpine meadows, steppe and desert. Non-arboreal pollen (NAP) therefore dominates the pollen samples in most regions. Percentages of arboreal pollen (AP) are high in samples from the southern and eastern Tibetan Plateau, where alpine forests are an important component of the vegetation. The relative importance of forest and non-forest vegetation across the Plateau clearly follows climatic gradients: forests occur on the southern and eastern margins of the Plateau, supported by the penetration of moisture-bearing airmasses associated with the Indian and Pacific summer monsoons; open, treeless vegetation is dominant in the interior and northern margins of the Plateau, far from these moisture sources. 4  The different types of non-forest vegetation are characterized by different modern pollen assemblages. Thus, alpine deserts are characterized by high percentages of Chenopodiaceae and Artemisia, with Ephedra and Nitraria. Alpine meadows are characterized by high percentages of Cyperaceae and Artemisia, with Ranunculaceae and Polygonaceae. Alpine steppe is characterized by high abundances of Artemisia, with Compositae, Cruciferae and Chenopodiaceae. Although Artemisia is a common component of all non-forest vegetation types on the Tibetan Plateau, the presence of other taxa makes it possible to discriminate between the different vegetation types. 5  The good agreement between modern vegetation and modern surface pollen samples across the Tibetan Plateau provides a measure of the reliability of using pollen data to reconstruct past vegetation patterns in non-forested areas.

Pollen records and age determinations from 2 profiles at Lake Nikolay, Lena Delta

Radiocarbon-dated pollen, rhizopod, chironomid and total organic carbon (TOC) records from Nikolay Lake (73°20'N, 124°12'E) and a pollen record from a nearby peat sequence are used for a detailed environmental reconstruction of the Holocene in the Lena Delta area. Shrubby Alnus fruticosa and Betula exilis tundra existed during 10,300-4800 cal. yr BP and gradually disappeared after that time. Climate reconstructions based on the pollen and chironomid records suggest that the climate during ca. 10,300-9200 cal. yr BP was up to 2-3 °C warmer than the present day. Pollen-based reconstructions show that the climate was relatively warm during 9200-6000 cal. yr BP and rather unstable between ca. 5800-3700 cal. yr BP. Both the qualitative interpretation of pollen data and the results of quantitative reconstruction indicate that climate and vegetation became similar to modern-day conditions after ca. 3600 cal. yr BP. The chironomid-based temperature reconstruction suggests a relatively warm period between ca. 2300 and 1400 cal. yr BP, which corresponds to the slightly warmer climate conditions reconstructed from the pollen. Modern chironomid and rhizopod assemblages were established after ca. 1400 cal. yr BP.

Pollen record with radiocarbon datings from Faddeyevskiy Island, Laptev Sea

Pollen, plant macrofossil, loss-on-ignition and radiocarbon analyses of a 1.4-m section in thermokarst topography from Faddeyevskiy Island (75°20'N, 143°50'E, 30 m elevation) provides new information on Late Pleistocene interstadial environmental history of this high Arctic region. Conventional radiocarbon dates (25,700 ± 1000, 32,780 ± 500, 35,200 ± 650 yr BP) and two AMS dates (29,950 ± 660 and 42,990 ± 1280 yr BP) indicate that the deposits accumulated during the Kargian (Boutellier) interval. Numerous mammoth (Mammuthus primigenius) remains that have been collected in vicinity of the site in this study were radio-carbon dated to 36,700-18,500 yr BP. Rare bison (Bison priscus) bones were dated to 32,200 ± 600 and 33,100 ± 320 yr BP. Poaceae, Cyperaceae, and Artemisia pollen dominate the spectra with some Ranunculaceae, Caryophyllaceae, Rosaceae, and Asteraceae. The pollen spectra reflect steppe-like (tundra-steppe) vegetation, which was dominant on the exposed shelf of the Arctic Ocean. Numerous Carex macrofossils suggest that the summer climate was at least 2°C warmer than today. The productivity of the local vegetation during the Kargian interstadial was high enough to feed the population of grazing mammals.

Pollen record from Golkow, Poland

Janczyk-Kopikowa (1966): The series of the organic deposits, developed in the vicinity of Golkow near Warsaw as oil shales and peats, was laid down in a grough valley and now rests on the deposits of the Middle Polish Glaciation (Riss). The organic deposits are overlain by the fluviale deposits of the North Polish Glaciation (Würm). The locality Golkow occurs beyond the extent of the continental glacier of this glaciation. Polen analysis completed by microfloristic examinations allows to determine the age of the organic series that is thought to be Eemian. The pollen diagram from Golkow does not call in question the stratigraphical position of the deposits investigated mainly due to its characteristic features such as minimum content of coniferous trees in the climatic optimum - about 5%, high percentage of Corylus - 77.5% and well developed phase of hornbeam. It may be well compared with other Eemian diagrams from the area of Poland and reveals much similar features. The development of vegetation at Golkow has depended upon the prevailing climate. At first, the cool climate brings about the development of plants having small thermal requirements. Here belong thin, park-like forests with pine and birch (Pinus, Betula) accompanied by the heliophilic plants such as Hippohäe and Ephedra. Improvement of climate that becomes warm and humid provides for development of deciduous forests prevailing in the climatic optimum, of the interglacial. Decrease of temperature causes a repeated change in the type of forest. This latter changes into coniferous forest with prevailing spruce (Picea) and fir (Abies) at the beginning, and then with pine (Pinus) and birch (Betula). During the Eemian Interglacial, the development of plants at Golkow terminates with a new and long-lasting predominance of pine-birch forests. However, such a longevity may be apparent only. Apparent character of this phenomenon is proved by a fact that the pollen spectra of the warm climatic periods have found their reflex in the oil shale that increased considerably slower than the layers off feebly decomposed peat evidencing the existence of cool pine-birch forests from the decline of the Interglacial. The water basin, in which the polen grains were laid down from surrounding plants is characterized by a calm sedimentation as proved by the occurrence of the oil shale. An insignificant water flow left behind some thin sand laminae. The not too deep basin becomes shallower owing to the growing water vegetation, and marshy vegetation. The growing of the plants causes a complete shallowing of the basin and formation of peat bog in situ, as proved by the peat beds occurring in the section. ---- Gadomska (1966): In the vicinity of Golków a series of organic deposits occurs amounting to 6.5-9.3 m in thickness, and consisting of oil shales, lacustrine silts and sands, as well as peats and peaty silts. The organic deposits fill up an old, small, but fairly deep lake basin, probably of finger-lake origin. It may be seen to-day as a slight lowering of the relief, filled up with soaked ground, stretching from north to south. On the basis of palaeobotanical examinations the organic deposits considered are of Eemian Interglacial age (Z. Janczyk-Kopikowa, 1063). The lower part of the organic series consists of a compact oil shale horizon, the maximum thickness of which may attain up to 8 m. The oil shales contain particularly in their upper part, numerous intercalations of arenaceous silts, dark grey or black in colour, or of sands mainly of lacustrine provenance. At the top of the oil shales are found peats, up to 2.5 m in thickness, covered by black, humus silts with numerous plant remains. The Eemian Interglacial deposits are covered by a series of fluviatile sands belonging partly to the Baltic Glaciation (bottom part of the series), partly to the Holocene (top part of the series). The thickness of the sands is 0.5-3.7 m. Higher up, there are found the Holocene and present-day deposits developed as clayey alluvion, or arenaceous slide rocks, or arenaceous-silty soil.