四川重楼属植物2新记录变种

报道了四川重楼属植物2新分布变种,分别是宽瓣球药隔重楼(Paris fargesii var. latipetala H. Li et V. G. Soukup)和红果五指莲(Paris axialis H. Li var. rubra H. H. Zhou, K. Y. Wu et R. Tao)。其中红果五指莲为贵州特有类群,且为孤点分布。四川新分布的发现对于重楼属植物的系统分类和迁徙演化研究具有重要意义。     

5种沙地灌木对地下水埋深变化的响应

研究干旱、半干旱区植物功能性状对地形地貌分异引起的地下水埋深变化的响应,对于揭示我国西北地区植物应对地下水埋深增加的维持机制和开展植被生态恢复建设具有重要意义。以海流兔河流域典型草原生境的5种建群灌木沙柳（Salix psammophila）、乌柳（Salix cheilophila）、柠条（Caragana korshinskii）、油蒿（Artemisia ordosica）和杨柴（Hedysarum laeve）为研究对象,认识其在滩地、丘间-丘顶典型分布和地势抬升条件下,沙地灌木生物量分配、根系分布及其对地下水埋深变化的生理响应特征。研究结果表明：（1）沙柳为深根系植物,生理功能的维持不仅依赖浅层土壤水,而且吸取地下水;乌柳、柠条、油蒿和杨柴为浅根系植物,主要水源是浅层土壤水。（2）在滩地共生条件下,沙柳和乌柳黎明前小枝水势（ψ_pd）、正午小枝水势（ψ_m）、气孔导度（g_s）、最大光化学量子产量（F_v/F_m）、表观量子效率（Φ）、光饱和点净光合速率（P_s）及光补偿点光量子通量密度（I_c）无显著差异;油蒿和杨柴ψ_m、g_s与F_v/F_m存在显著差异。（3）由丘间-丘顶分布引起地下水埋深不同的条件下,丘间和丘顶沙柳ψ_pd、ψ_m、P_s和I_c差异显著,乌柳ψ_pd和g_s差异显著,柠条和油蒿仅ψ_m差异显著,这几种灌木其它生理性状在丘间和丘顶无显著差异。（4）沙柳生理性状与地下水埋深的相关性分析表明,随着地下水埋深的增加,g_s和P_s呈降低趋势,Φ呈升高趋势,I_c没有明显变化。综上所述,不同根系分布的灌木,对地下水埋深变化的响应存在明显差异。研究结果可以为干旱生态系统原生植被恢复的地下水资源管理提供参考。     

光枝勾儿茶内生真菌多样性及抑菌活性研究

为探索贵州苗药光枝勾儿茶内生真菌类群特征、分布部位及其抑菌活性,该研究采用传统方法对贵州省贵阳市和黔西市光枝勾儿茶内生真菌进行分离,并基于分子生物学及统计学对其分类地位进行鉴定及多样性评价,最后通过微量肉汤倍比稀释法筛选具有抑菌活性的菌株。结果表明:(1)从光枝勾儿茶中分离到191 株内生真菌,隶属于3 个门5 个纲10 个目15 个科19 个属,优势属为叶点霉属(Phyllosticta)、间座壳属(Diaporthe)、葡萄座腔菌属(Botryosphaeria)和刺盘孢属(Colletotrichum)。(2)黔西光枝勾儿茶内生真菌香农-维纳多样性指数(H''_Q=2.112)较贵阳(H''_G=1.801)高,索伦森相似性指数Cs_G-Q为0.923,不同组织香农-维纳多样性指数为茎(H''_S=2.004)>根(H''_R=1.764)>叶(H''_L=1.654)>果实(H''_F=1.473),茎和叶内生真菌的索伦森相似性最高(Cs_S-L=0.667)。(3)筛选出的21株内生真菌对供试菌大肠杆菌(Escherichia coli)、金黄色葡萄球菌(Staphylococcus aureus)和沙门氏菌(Salmonella enterica)具有抑菌效果,其中Diaporthe sp. QX4G6对大肠杆菌、金黄色葡萄球菌和沙门氏菌的最小抑菌浓度分别为12.5、6.25、12.5 mg·mL^-1,最小杀菌浓度分别为12.5、6.25、12.5 mg·mL^-1。以上研究结果揭示了光枝勾儿茶蕴藏丰富的内生真菌资源,不同地区及组织内生真菌类群组成有差异,多个分离菌株具有抗菌活性,为光枝勾儿茶内生真菌天然抗菌药物或药源研发奠定了基础。     

贵州稀有濒危种子植物物种多样性保护与利用的研究(附录)

全球性生物多样性保护与利用正逐步走向高质量发展,拥有丰富物种资源已成为夺取生物科技制高点的关键,因此,贵州稀有濒危种子植物多样性保护与利用的研究具有重要意义。在对贵州稀有濒危种子植物资源调查的基础上,选取具有重要应用价值的小黄花茶(Camellia luteoflora)、贵州红山茶(C. kweichowensis)等103种贵州稀有濒危植物,采用模式标本产地植物追溯、现存分布区及种群散布特征调查、物种濒危等级评估、混合采种迁地保育、生物资源测试分析与资源评价相结合的方法,开展了植物物种多样性编目及其迁地保护、种子生物学特性与种子散布及分布动态变化、种苗繁育与应用栽培试验等研究。结果表明:(1)调查评估的103种贵州稀有濒危种子植物都采取了就地保护措施,但迄今为止对濒危威胁程度减轻的效应不明显,抢救性迁地保护依然必要,其中极危(CR)23种、濒危(EN)30种、易危(VU)40种、近危(NT)7种、无危(LC)3种; 采用同一地理种源迁地保护有效种群P_n≥L_f·E_e·A_m的栽培方法,在贵州省植物园开展了迁地保护引种栽培试验。其中,国家重点保护野生植物33种、3 650株,引种栽培平均成活率为95.47%,贵州及毗邻川、滇、桂极少数地区特有分布的种子植物70种、11 010株,引种栽培平均成活率为95.80%,并且建成了活植物保育圃30 200 m²。(2)搜集文献大数据结合GIS信息追踪调查研究,发现了小黄花茶新分布区2.5 km²、长柱红山茶(C. longistyla)新分布区1.5 km²、美丽红山茶(C. delicata)新分布区1.0 km²、皱叶瘤果茶(C. rhytidophylla)新分布区6.0 km²、红花瘤果茶(C. rubituberculata)新分布区50.0 km²、贵州槭(Acer guizhouensis)新分布区0.3 km²; 新分布区种群增量对其减轻濒危程度的评估效果影响不明显。(3)选择了小黄花茶、贵州红山茶等20种具有贵州高原区域环境特色的经济植物开展了种苗繁育和应用栽培试验,建立种苗繁育基地13.4 hm²,繁育种苗73万余株,在林业绿化工程应用中示范栽培140.0 hm²(31万株),≥2年生苗木应用栽培存活率92.00%～98.00%,这为有关稀有濒危植物的科学研究及种质创新利用提供科学参考。     

黄土丘陵区生物结皮坡面产流产沙与其分布格局的关联

生物结皮在黄土丘陵区退耕地广泛发育，并在干扰等因素下形成不同分布格局。为明确生物结皮分布格局对坡面产流产沙的影响，采用模拟降雨试验研究了生物结皮条带状、棋盘状和随机3种分布格局对坡面产流产沙的影响，借助景观生态学方法，计算了生物结皮斑块的分布格局指数，解析了生物结皮分布格局与坡面产流产沙的关联。结果表明，（1）生物结皮分布格局显著影响坡面初始产流时间、径流率和土壤侵蚀速率。相同盖度条件下，生物结皮坡面的随机分布格局较带状和棋盘式分布格局显著降低了坡面径流量和产沙量，其径流率与土壤侵蚀速率分别是带状格局的71.7%、12.9%，是棋盘格局的89.6%、31.8%。（2）坡面径流率与生物结皮斑块最大斑块指数、斑块连结度呈极显著负相关，与分离度呈极显著正相关，与景观分裂指数呈显著正相关。（3）土壤侵蚀速率与生物结皮斑块最大斑块指数、斑块连结度呈显著负相关，与景观分裂指数、分离度呈显著正相关。（4）坡面径流率（Y₁）和土壤侵蚀速率（Y₂）与分离度（SPLIT，Splitting Index）的关系式为：Y₁=1.116+0.017SPLIT，R²：0.957；Y₂=22.767+17.936SPLIT，R²：0.801。（5）以上结果表明分布格局主要是通过影响生物结皮在坡面的分离度进而影响坡面产流产沙，随着生物结皮斑块分离度增大，生物结皮斑块破碎度增大，坡面产流产沙增多。从分布格局的角度解析了生物结皮对产流、产沙的影响，量化了生物结皮坡面径流率与侵蚀率与格局指数之间的关系，为量化认识生物结皮的水土保持功能提供了科学依据。     

云南48种木兰科野生植物资源遗传多样性研究

为了解云南省木兰科（Magnoliaceae）野生植物资源的遗传多样性,利用ISSR分子标记技术对48种木兰科野生植物资源进行研究。结果表明,10对引物共扩增出151条带,均为多态性条带,多态性条带百分率为100%。总的观测等位基因数（N_a）为2.000 0,有效等位基因数（N_e）为1.564 5,Nei’s基因多样性指数（H）0.337 9,Shannon’s信息指数（I）为0.510 1。总的基因多样性指数（H_t）为0.368 0,属间基因多样性指数（D_st）为0.251 9,占68.4%,基因分化系数（G_st）为0.684 0,基因流（N_m）为0.231 0。UPGMA聚类分析将48种木兰科植物划分为7个类群,各类群并非按照属聚在一起,而是不同属植物相间分布,长喙厚朴（Magnolia rostrata）、素黄含笑（Michelia flaviflora）和球花含笑（M.sphaerantha）可能为云南省木兰科植物中的原始种。48种木兰科野生植物总体具有较高的遗传多样性,但属间遗传变异较高,基因流较小,存在遗传漂变的风险,聚类结果与刘玉壶的分类系统存在分歧,这从分子水平为木兰科植物间的起源、进化与分类提供了重要依据。     

獐耳细辛属重瓣川鄂獐耳细辛在陕西省的新分布

该文报道了獐耳细辛属植物重瓣川鄂獐耳细辛(Hepatica henryi f. pleniflora Xiao D. Li et J. Q. Li)的新分布区——陕西省岚皋县南宫山国家森林公园及镇坪县化龙山国家森林公园,将该种的分布区向西北扩展到了陕西省的巴山地区,丰富了陕西省新增植物分布记录,为巴山地区的物种多样性研究提供了证据。     

根深决定不同个体大小梭梭对夏季干旱生理响应的差异

理解干旱与树木个体发育阶段或大小之间的相互作用关系是一个重要的优先研究方向,因为依赖于个体大小的特定死亡率分布格局对生态系统的结构和功能将产生重要的影响。以广泛分布于古尔班通古特沙漠南缘的主要建群种梭梭（Haloxylon ammodendron）为研究对象,通过测定越夏生长期间不同个体大小梭梭黎明前同化枝水势（Ψ_pd）、压力-容积（P-V）曲线、最大气孔导度（G_s）、叶绿素荧光（F_v/F_m）,调查不同个体大小梭梭根系分布特征和根深,将越夏生长期间土壤水分状况、植物生理性状与根深相结合,分析不同大小梭梭对干旱胁迫的响应差异。结果表明：（1）夏季干旱胁迫显著降低了梭梭幼苗生理性状的特征值：同成年植株相比,梭梭幼苗黎明前同化枝水势（Ψ_pd）、PSⅡ反应中心最大光化学效率（F_v/F_m）、饱和膨压渗透势（π₁₀₀）、膨压消失点渗透势（Ψ_TLP）显著降低;（2）梭梭生理性状与根深之间存在线性关系,根深决定不同大小梭梭的水分状况（Ψ_pd）进而主导了越夏生长期间的其他生理性状表达（F_v/F_m、π₁₀₀、Ψ_TLP）;（3）不同大小的梭梭对干旱胁迫的响应差异是植物根系分布特征和生境相互作用的结果,根深在梭梭应对夏季干旱胁迫中起重要作用。总之,梭梭幼苗根系发育不完全（浅根系）,只能利用降水带来的有限的浅层土壤水,对干旱胁迫响应敏感;而成年梭梭植株能最大限度的利用深层土壤水分,减少表层土壤干旱对其产生的不利影响。     

桂林喀斯特石山50种常见植物叶片光合特性

选取桂林喀斯特石山生境中常见的50种植物为研究对象,分别测定叶片单位面积最大净光合速率（A_area）、单位质量最大净光合速率（A_mass）、气孔导度（Gs）、水分利用效率（WUE）、胞间/环境CO₂浓度比值（Ci/Ca）和蒸腾速率（Tr）等光合特性指标,探讨不同物种光合特性的差异以及光合特性之间的内在联系,以此探究不同植物适应喀斯特石山生境所表现出的光合生理特性。结果表明,50种植物叶片A_area,A_mass,Gs,WUE,Ci/Ca和Tr的平均值分别为8.35 μmol m^-2 s^-1,110.98 nmol g^-1 s^-1,0.10 mol m^-2 s^-1,94.84 μmol/mol,0.57和2.37 mmol m^-2 s^-1;方差分析表明,不同物种之间在A_area,A_mass,Gs,WUE,Ci/Ca和Tr之间存在显著差异。Pearson相关性分析表明,表征50种常见植物叶片光合特性的6个指标相关性除Ci/Ca与A_area和A_mass,WUE与A_mass不一致外,其他指标两两之间相关性均表现为一致性,其中Gs与Ci/Ca呈极显著的正相关。主成分分析表明,在6个光合特性指标中,Gs和Ci/Ca可作为反映喀斯特石山植物适应生境的重要光合指标,主要表征对水分条件的敏感程度以及耐旱性强弱,同时反映了植物叶片光合速率大小,用于衡量植物对喀斯特生境的生理生态适应性。基于Gs和Ci/Ca进行聚类分析表明,50种植物划分为3类：即中等Gs较高Ci/Ca型,较低Gs较高Ci/Ca型和较低Gs,Ci/Ca型。本研究表明,喀斯特生境植物在生理生态方面所表现出的适应策略主要为对资源利用方式及抵御外界不利环境的适应策略,这为后续选择物种加速植被恢复演替进程提供了参考。     

内蒙古草原常见植物叶片δ13C和δ15N对环境因子的响应

在中国东北样带沿线的内蒙古草原地区采集了一些常见植物的叶片样品,并测定其δ¹³C和δ¹⁵N值,分析了其统计学特征以及对环境因子(年平均降雨量和温度)的响应模式。发现东北样带草原区同时存在C₃和C₄两种不同光合途径的植物,但是C₃植物占主导地位,C₄植物数量有限。C₃植物叶片δ¹³C随着年平均降雨量和年平均温度的升高而显著降低,反映了此区域C₃植物δ¹³C受控于降水量和温度。C₄植物的叶片δ¹³C值随着降雨量的增多而有轻微升高的趋势,但是C₄植物的叶片δ¹³C值对年平均温度的响应不敏感。不论对C₃植物还是C₄植物而言,叶片δ¹⁵N都随降雨量增加而显著降低,即干旱区的植物叶片δ¹⁵N大于湿润地区,这说明降水是影响植物叶片δ¹⁵N的一个重要因素。然而两者叶片δ¹⁵N对温度的响应不敏感。     

A new genus of Early Devonian plants with novel strobilar structures and vegetative appendages from the Posongchong Formation of Yunnan,China

A new plant, Dibracophyton acrovatum gen. et sp. nov., is described from the Lower Devonian (Pragian) Posongchong Formation of Wenshan District, southeastern Yunnan, China. The plant has creeping axes from which arise vegetative and fertile axes. The vegetative axes helically bear lateral dichotomous appendages with curved or round tips. The fertile axes possess terminal strobili with numerous fertile units arranged in irregular helices. Each fertile unit consists of a stalked long-elliptical sporangium, with dehiscence into two equal valves, and two discrete long-ovate bracts covering sporangium from above–below directions. A new genus and species is thus established based on these characters and is temporarily regarded as incertae sedis of Tracheophyta, although it perhaps closes to the barinophytes in affinity. Detailed comparisons with other plants having a similar architecture, i.e., sporangia closely associated with modified vegetative structures, are made. The discovery of D. acrovatum further enriches the composition of the Posongchong flora and demonstrates great morphological disparity of the Early Devonian vascular plants.     

A New Lower Devonian Genus from Yunnan,with Notes on the Origgin of Leaf

A new genus and species, Eophyllophyton bellum, is described from the Posongchong Formation (Siegenian) of the Lower Devonian in Wenshan cfistrict, Yunnan, China This plant bears lateral branching laminate structures interpreted as leaves, which are believed to derive from lateral axial branching systems In addition similarly organised sporophylls are considered homologous to these leaves. In the possession of these vegetative leaves and sporophylls Eophyllophyton, is significantly more advanced than any other Siegenian taxon.     

早泥盆世始叶蕨（新属）及叶子起源的讨论

本文报道了滇东南文山地区下泥盆统坡松冲组（西根阶）地层中一陆生植物新属,种－纤细始叶蕨（Eophyllophyton bellum gen.et sp.nov),植物体矮小,多次二叉分枝,远端枝二列互生长出次一级二叉分枝为主的末级枝,枝系起源叶顶生成侧生,多次分叉,缩合,扁化,蹼化并部分并合,生殖单位可下垂,由二枚顶端内卷的孢子叶组成,孢子囊圆至肾形,沿远,端加厚带开裂,成列侧生在孢子叶相对面的叶轴上,上述特征表明新植物远比同期的莱尼类,三枝蕨类和工蕨类复杂和进化,并显示出和工蕨类的某些亲缘关系,文内还探讨了枝系起源叶（大型叶）的发生和演化。     

A new microphyllous plant from the Lower Devonian of Yunnan Province, China

Sterile and fertile shoots of a microphyllous vascular plant are described from the Lower Devonian Posongchong Formation (? upper Siegenian), Yunnan Province, southern China and placed in the new taxon Halleophyton zhichangense gen. et sp. nov. The sterile axes are characterized by conspicuous, contiguous swollen rhomboidal to hexagonal leaf bases and relatively broad spatulate laminae. The relief of the leaf bases is not so pronounced in the fertile areas, which bear numerous cauline sporangia with two equal valves. In vegetative characters the new fossils, although smaller and more fragmentary, resemble Drepanophycus gaspianus but in the absence of information on the fertile parts of the latter and because of the controversy surrounding the concept of the genus Drepanophycus, we decided to erect a new genus. Its relationship to the lycophytes sensu stricto is equivocal as exact details of the relationship of sporangium with leaf and sporangium vascularization remain unknown.     

EARLY VERTEBRATE FOSSILS FROM THE EARLY DEVONIAN OF ZHAOTONG DISTRICT, NORTHEASTERN YUNNAN

Astothegaleaspids,thePosongchongFormationofZhaotongDistrictismostsimilartothePingyipoFormationofJiangyou,Sichuan.BothLungmenshanaspisandSanqiaspisareexclusivelyfoundinthesetworegions,indicatingthepresenceofbiogeographicconnection.PaleontologicStudiesSubclassGaleaspidaTarlo,1967orderHuananaspiformesJanvier,l975FamiIySanqiaspididaeLiu,l975GenusSanqiasPisLiu,l975TyPespeciesSanqiasPisrostrataLiul975.ReferredspeciesSanqiasPiszhaotongensisLiul97iEmendeddiagnosisCephalicshieldnarrow,rou…     

A new species of Estinnophyton from the Lower Devonian Posongchong Formation, Yunnan, China; its phylogenetic and palaeophytogeographical significance

A new species of early vascular land plant, Estinnophyton yunnanense , is described from the Lower Devonian (upper Pragian) Posongchong Formation at Zhichang village, Gumu Town, Wenshan District, Yunnan Province, China. This plant possesses aerial stems with isotomous branching. The leaves are slender, once- or twice-bifurcated, and arranged in low spirals or pseudowhorls. Fertile leaves are morphologically identical to the vegetative ones, clustered on the axis, forming a loose strobilus-like structure. There are three to six fertile leaves in each gyre. Two ovoid-elongate sporangia are singly attached to the adaxial surface each subtended by a recurved stalk, either at the base of each segment in a once-bifurcate fertile leaf or below the points of successive divisions in a twice-bifurcate fertile leaf. Estinnophyton yunnanense differs from two other species described previously from Europe, E .  gracile and E .  wahnbachense , mainly in possessing two single stalked sporangia on each fertile leaf rather than two pairs of sporangia, as well as in the dimensions of its leaves. In view of the complex taxonomy and nomenclature surrounding the type material of Estinnophyton , we considered possible emendation of the generic diagnosis based on the new Chinese material. Following comparison with typical members of the Protolepidodendrales and other taxa, Estinnophyton is considered to have had a closer affinity with primitive sphenopsids than the protolepidodendralean lycopsids. A comparison between Chinese and European floras containing Estinnophyton indicates that South China and Western Europe had floristic elements in common although they were in different phytogeographical units during the Early Devonian epoch.  © 2004 The Linnean Society of London, Botanical Journal of the Linnean Society , 2004, 146 , 201–216.     

Stachyophyton gen. Nov. Discovers from Lower Devonian of Yunnan and Its Significance

This paper describes a new plant—Stachyophyton yunnanense gen. et sp. nov., from Posongchong Formation (Siegenian) of the Lower Devonian in Wenshan district, Yunnan, China. This plant shows some similar organization in fertile region and sterile parts in comparison with other Devonian genera (Protolepidodendron, Colpodexylon, Leclercqia, Enigmophyton, Krithodeophyton). It may be that the plant represents an intermediate stage in evolution between Enigmophyton and Krithodeophyton, and shows perhaps a certain affinity to lycopods. According to the salient features of Stachyophyton, it can not be, at present, placed in any group known in vascular plant. It can be assigned to definite position of taxon basing on much fossils record and further studies.     

Diversity dynamics of Zosterophyllopsida

The Zosterophyllopsida were major contributors to the diversification of early land plants. We present the first detailed analysis of the diversity dynamics of these plants from an updated database of all currently recognized zosterophyllopsid species. A set of quantitative methods classically used in palaeodiversity studies was applied to two data sets. The first one, ‘Zosterophyllopsida sensu stricto’, corresponds to the clade identified by Hao & Xue (The Early Devonian Posongchong Flora of Yunnan. (2013), Science Press). In the second, called ‘Zosterophyllopsida sensu lato’, barinophytalean‐type plants and taxa for which zosterophyllopsid affinities are suspected are added. The number of localities is used to explore sampling bias. Results show that sampling effect is minimal for the Early Devonian. For this time interval, both data sets record consistent patterns of changes suggesting that, whatever their affinities, all taxa included in the Zosterophyllopsida sensu lato show similar evolutionary trends. The diversity dynamics of zosterophyllopsids are characterized by a radiation during the Lochkovian, maximal values in the Pragian and a decline starting in the Emsian. The proportion of zosterophyllalean taxa with terminal sporangia is high until the Late Lochkovian when gosslingialean taxa without terminal sporangia evolved. During the Middle and Late Devonian, when diversity patterns are strongly affected by sampling, zosterophyllopsid diversity is low and characterized by a high proportion of barinophytacean and gosslingialean taxa, the latter becoming extinct in the Early Frasnian.     

Notes on a fertile rhyniophytoid from the Lower Devonian of Guangxi,southwestern China

A fertile rhyniophytoid is discovered from the Lower Devonian of Guangxi, southwestern China, which is characterised by small size, dichotomous branchings and terminal sporangia. This plant is similar to the genus Aberlemnia Gonez et Gerrienne and also, is comparable to some mesofossil morphotypes of early land plants from the Early Devonian (Lochkovian) Old Red Sandstone floras. This study adds new data to the generally zosterophyll-dominated Early Devonian floras of South China and sheds some lights on the palaeophytogeography of rhyniophytoids.     

A NEW SPECIES OF ZOSTEROPHYLLUM FROM THE EARLY DEVONIAN OF NEW BRUNSWICK

A new species of Zosterophyllum, Z. divaricatum Gensel, is described from the late Early Devonian (Emsian) of northern New Brunswick, Canada. It is a Platyzosterophyllum type, consisting of slender sometimes bifurcating axes with laterally borne sporangia oriented to one side of the axis. The species is distinctive in that axes bifurcate within fertile regions and in sporangium shape and attachment. Aspects of the morphology of axis and sporangium cuticle, tracheids, and spores are presented and considered in relation to comparable features in other Zosterophyllum species. Associated vegetative axes exhibiting H- and K-branching patterns and also cuticular features similar to the fertile specimens are described and it is suggested that they may represent parts of the same plant. Zosterophyllum divaricatum is most similar to Z. llanoveranum, Z. fertile, and Z. spectabile, and also resembles Rebuchia ovata to some extent. Z. divaricatum offers considerable information on variation within one species concerning sporangium shape, attachment, and distribution and expands the known diversity of Platyzosterophyllum types.