Photosynthetic Pathways and Life Forms in Different Grassland Types from North China

Photosynthetic pathways (C₄, C₃, and CAM species) and plant life forms of three grassland types in North China were compared. Of the total 201 species, 144 species in 78 genera and 34 families had C₃ photosynthetic pathway, 56 species in 35 genera and 11 families had C₄ photosynthetic pathway, and 1 species had CAM photosynthetic pathway. The number of C₄ species in Songnen meadow was 70–80 % greater than that in Xilinguole steppe and Hunshandak desert grassland, but that for C₃ species did not differ significantly among the three grassland types. The number of therophytes in the Songnen meadow was relatively greater than that of the other two grassland types, but that of hemicryptophytes was lower. Thus the distribution of C₄ species and plant life form is probably related to precipitation.     

Photosynthetic Pathway Types of Forage Species Along Grazing Gradient from the Songnen Grassland,Northeastern China

Photosynthetic pathway types (C₃ and C₄ species) and their dynamics along grazing gradient were determined for 42 plant species in 30 genera and 13 families from the Songnen grassland, Northeastern China. Of the total, 10 species in 9 genera and 4 families had C₄ photosynthesis; 32 species in 21 genera and 12 families had C₃ photosynthesis. The proportion of C₄ species in total plants and C₄/C₃ increased with grazing intensity, and peaked in overgrazed plot. Most of the increased C₄ species (6 of 10) along the grazed gradient were annual grasses and halophytes. This indicated that the C₄ species had greater capacity to tolerate environmental stresses (e.g. drought and saline) caused by animal grazing in the Songnen grassland, Northeastern China.     

C<Subscript>3</Subscript> and C<Subscript>4</Subscript> photosynthetic pathways and life form types for native species from agro-forestry region,Northeastern China

Of the total 570 species, 194 species in 116 genera and 52 families were found with C₃ photosynthesis, 24 species in 17 genera and 6 families with C₄ photosynthesis, and 2 species in 1 genera and 1 family with CAM photosynthesis. 90 % of the total species can be found in Changbai Mountain flora, more a half (69 %) in North China flora, and about 1/3 in Mongolian flora and Xinan flora, respectively. The occurrence of C₄ species was not as common as that in adjacent grasslands and deserts, but relatively more than in the adjacent forests. Of the total 24 C₄ species, 63 % C₄ species (15 of 24) was found in Gramineae. Nine life form types can be found, reflecting the moist climate in the region, especially the occurrence of epiphyte and liana forms. Relatively more geophyte life form plants suggested the winter in the region was much colder than in grasslands. These indicated that both ecological studies and land management decisions must take into account plant photosynthetic pathway and life form patterns, for both of them are closely related to climatic changes and land use.     

The occurrence of C4 plants and their morphological functional types in the vegetation of Xinjiang, China

Floristic composition, morphological functional types and habitat distributions for C₄ species were studied in Xinjiang, North-western China. 89 species, in 9 families and 41 genera, were identified with C₄ photosynthesis. 48 % of these C₄ species were found in Monocotyledoneae, e.g. Cyperaceae (5 species), Gramineae (38 species), the other 52 % was in Dicotyledoneae, e.g. Chenopodiaceae (29 species), Amaranthaceae (7 species), and Polygonaceae (5 species). Compared with those in semi-arid grasslands in North China and tropical forests in South China, more plant families were found with the occurrence of C₄ plants in this arid region. Relatively higher annual species (63 %), shrubs (18 %), and Chenopodiaceae C₄ species (33 %) compositions were the primary characteristics for the C₄ species occurring in Xinjiang, and this was remarkably related with its arid environment. More Chenopodiaceae C₄ species occurring in the region suggested that this type of C₄ species may have higher capacity to fit the air and soil droughts. There was a strong relationship between C₄ occurrence and habitat distributions, more than half of the total 89 C₄ species were found in disturbed and cultivated lands and early stages of vegetation successions, indicating C₄ occurrence was not only related with climate changes, but also with land uses and vegetation dynamics.     

Photosynthetic Pathways and Life Form Types for Native Plant Species from Hulunbeier Rangelands,Inner Mongolia,North China

Photosynthetic pathway Types (C₃, C₄, and CAM) and life forms of native species from Hulunbeier rangelands, north China were studied. Of the total 258 species, 216 species in 132 genera and 42 families had C₃ photosynthetic pathway, including dominant herbs, e.g. Stipa baicalensis Roshev. and Leymus chinensis (Trin.) Tzvel., Filifolium sibiricum Kitam. and Arudinella hirta (Thunb.) Koidz. 38 species in 28 genera and 10 families were found with C₄ photosynthesis, and 4 species in 2 genera and 1 family had CAM photosynthetic pathway. The occurrence of C₄ species was common in Gramineae and Chenopodiaceae, and the two families were leading ones within C₄ plants. More than 52 % of the total 258 species were in H form, 21 % in Th form, 19 % in G form; the other life form Types, e.g. Ch, M, N, and HH, formed less than 3 %. 68 % of C₄ species were in Th form and 24 % in H form, indicating that these Types were the dominant life forms for C₄ species in the rangeland region. The occurrence of C₄ species was closely related with plant habitats, disturbed lands had the highest C₄ abundance (55 % of the total C₄ species), followed by grasslands and sandy soil, and forests had the lowest C₄ abundance (8 %). Hence the occurrence of C₄ species could be efficient indicator for rangeland dynamics in Hulunbeier rangelands.     

Plant functional types and their ecological responses to salinization in saline grasslands, Northeastern China

Photosynthetic pathways (C₃, C₄, and CAM) and morphological functional types (e.g. shrubs, high perennial grasses, short perennial graminaceous plants, annual grasses, annual forbs, perennial forbs, halophytes, and hydrophytes) were identified for the species from salinity grasslands in Northeastern China, using the data from both stable carbon isotope ratios (¹³C) and from the references published between 1993 and 2002. 150 species, in 99 genera and 37 families, were found with C₃ photosynthesis, and most of these species are dominants [e.g. Leymus chinensis (Trin.) Tzvel., Calamagrostis epigeios (L.), Suaeda corniculata (C.A. Mey.) Bunge]. 40 species in 25 genera and 8 families were identified with C₄ photosynthesis [e.g. Chloris virgata Sw., Aeluropus littoralis (Gouan) Parlat] and 1 species with CAM photosynthesis. Gramineae is the leading family with C₄ photosynthesis (27 species), Chenopodiaceae ranks the second (5 species). The significant increase of C₄ proportions with intense salinity suggested this type plant is remarkable response to the grassland salinization in the region. 191 species were classified into eight morphological functional types and the changes of most of these types (e.g. PEF, HAL, and HPG) were consistent with habitats and vegetation dynamics in the saline grassland. My findings suggest that the photosynthetic pathways, combined with morphological functional types, are efficient means for studying the linkage between species and ecosystems in this type of saline grassland in Northeastern China.This revised version was published online in March 2005 with corrections to the page numbers.     

基于结构方程模型分析森林草原带草本物种丰富度对景观因子的响应

受损景观是由不同比例的植被斑块组成的镶嵌体,阐明植被斑块的景观结构特征对物种多样性的影响,有助于提高受损景观物种多样性保护。在塞罕坝自然保护区选取38个天然植被斑块,其中包括12个草地、11个灌木林和15个天然次生林。根据光合作用的不同途径将草本物种划分为C_3和C_4功能群。选取斑块面积、形状指数、隔离度指数及每个斑块500 m缓冲区内的森林和草地比例作为景观因子。通过结构方程模型探讨C_3、C_4草本物种丰富度与景观因子的相互关系。斑块面积(2.18—74.06 hm~2)与C_3、C_4草本及总物种丰富度分布格局均具有显著的正相关关系(P0.05),且对C_3草本的影响最大;形状指数(1.06—3.11)、隔离度指数(33.51—327.65)对C_3、C_4草本及总物种丰富度影响不显著(P0.05);C_3、C_4草本及总物种丰富度与毗邻斑块草地比例(4.20%—64.95%)呈正相关,而与森林比例(35.05%—95.80%)呈负相关。研究区斑块面积和毗邻斑块植被构成是影响C_3、C_4草本植物的主要因素。在破碎化景观中保存面积大的天然植被斑块及提高毗邻斑块草地比例是保护关键C_3、C_4草本植物物种的有效途径。     

C4 Species and their Response to Large-Scale Longitudinal Climate Variables Along the Northeast China Transect (NECT)

Natural occurrence of C₄ species, life forms, and their longitudinal distribution patterns along the Northeast China Transect (NECT) were studied. Six vegetation regions experiencing similar irradiation regimes, but differing in longitude, precipitation, and altitude were selected along the NECT from 108 to 131 °E, around altitude of 43.5 °N. Seventy C₄ species were identified in 41 genera and 13 families. 84 % of the total C₄ species were found in four families: Gramineae (38 species), Chenopodiaceae (11 species), Cyperaceae (5 species), and Amaranthaceae (5 species). C₄ grasses make up 54 % of the total identified C₄ species along the NECT and form the leading C₄ family in meadow, steppe, and desert along the NECT. C₄ Chenopodiaceae species make up about 16 % of the C₄ species and become less important, particularly in the meadow and the eastern end of the NECT. 57 % of the total C₄ species are therophytes and 37 % are hemicryptophytes, which is consistent with floristic composition and land utilization. In general, the number of C₄ species decreased significantly from the west to the east or from dry to moist areas along the NECT, and was remarkably correlated with annual precipitation (r ²= 0.677) and aridity (r ²= 0.912), except for salinized meadow region. The proportion of C₄ species from all the six vegetation regions was considerably correlated with these two climatic parameters (r ²= 0.626 or 0.706, respectively). These findings suggest that the natural occurrence of C₄ species varies significantly along the large-scale longitudinal gradient of the NECT. The notable relationship of C₄ species number and proportion in the flora with variations in annual precipitation and aridity suggest that these two climatic parameters are the main factors controlling the longitudinal distribution patterns of C₄ species along the NECT.     

Photosynthetic Pathways,Life Forms,and Reproductive Types for Forage Species Along the Desertification Gradient on Hunshandake Desert,North China

Floristic compositions, life forms, reproductive types for forage species, and their responses to desertification in Hunshandake desert were studied. 164 species, in 30 families and 94 genera, were identified with C₃ (137 species), C₄ (25 species), and CAM (2 species) photosynthesis. Of the 25 C₄ species, 76 % were grasses and Chenopodiaceae species (hereafter chenopods). This suggests that the C₄ species mainly occurred in a few families in the desert region. The reduction of C₃ species and the increase of C₄ species with desertification indicated that C₄ species might have higher tolerance to environmental stresses (e.g. dry and poor soil). Relatively more hemicrytophyte and therophyte forms in the desert are related to the local temperate climate and vegetation dynamics. Relatively greater proportions of C₄/C₃ and clonal species/sexual species at mobile dune showed that the C₄ species and clonal species could make greater contribution to sand land restoration in the Hunshandake desert.     

Photosynthesis pathways,ecological characteristics,and the geographical distribution of the Cyperaceae in Japan

Summary The nature of the photosynthetic pathways of Cyperaceae found in Japan were investigated on the basis of Kranz anatomy, the CO₂ compensation concentration and previously reported data. Among 301 species (96% of all cyperaceous species recorded in the region), 58 species were classified as being C₄ plants. These C₄ species were scattered among the tribes Fimbristylideae, Lipocarpheae, Cypereae and Rhynchosporeae in the subfamily Cyperoideae. The genera Cyperus, Eleocharis and Rhynchospora included, in Japan, both C₃ and C₄ species within a single genus. Using these data, an analysis was made of the ecological characteristics and geographical distribution of the C₃ and C₄ species in Japan. Although cyperaceous species grow in markedly different environments, the majority were found in wet and aquatic areas (61%) or shaded areas, such as forest floors (20%). Most of the C₃ species were also hygrophytes (58%) and forest-living species (25%), and C₃ species growing in mesic and dry areas were relatively rare. The C₄ species inhabited wet and aquatic (75%), mesic (13%) and dry areas (6%) and showed marked ecological characteristics with respect to soil-moisture conditions, unlike other C₄ plants, although they were absent from shaded habitats. In order to determine the climatic factors that influence the relative floristic abundance of C₃ and C₄ members of the Cyperaceae in Japan, the ratios of number of C₄ species to the total number of members of Cyperaceae (C₄ percentage) in 16 representative locales were examined in terms of various climatic variables. There were strong positive correlations between the C₄ percentage and temperature. Among the C₃ groups of three subfamilies, there were different distributional trends for various temperature regimes. The C₃ subfamily Caricoideae increased its relative contribution to the cyperaceous flora with a decrease in mean annual temperature, while the C₃ subfamily Sclerioideae exhibited the opposite pattern. The C₃ group of the subfamily Cyperoideae did not show any marked change in pattern along temperature gradients, unlike the two other C₃ subfamilies, and seemed to be heterogeneous in terms of its response to temperature. The relationships between the C₄ biochemical subtypes and ecological characteristics are also discussed.     

The occurrence of C4 photosynthesis in Yunnan province, a tropical region in South-western China

Floristic composition, morphological functional types, and altitudinal distribution pattern for C₄ species were studied in Yunnan province, South-western China. 159 species, in 6 families and 60 genera, were identified with C₄ photosynthesis. 93 % of these C₄ species were found in Monocotyledoneae, e.g. Cyperaceae (18 species), Gramineae (129 species), and Commelinaceae (1 species), the other 7 % was in Dicotyledoneae, e.g. Amaranthaceae (5 species), Portulacaceae (4 species), and Chenopodiaceae (2 species). Hence C₄ plants mainly occurred in very few families in the tropical region. Compared with those in semi-arid grasslands and arid deserts in North China, more C₄ grasses and much less Chenopodiaceae C₄ species occurred in the tropical region. This indicates the physiological responses of C₄ plants from the two families are very different. Chenopodiaceae C₄ species may be more fit semi-arid and arid environments, while C₄ grasses are more fit the moist tropical conditions. There was a strong relationship between C₄ distribution and altitude in the tropical region. Altitudinal distribution pattern for C₄ species in the region was consistent with altitude, climate, and habitats.     

C4 Plants in the Vegetation of Tibet,China: Their Natural Occurrence and Altitude Distribution Pattern

Floristic composition, life forms for C₄ species, and the pattern of altitude distribution were studied on Tibetan Plateau. 79 species, in 7 families and 46 genera, were identified with C₄ photosynthesis. 95 % of these C₄ species belong to Gramineae (51 species), Cyperaceae (14 species), and Chenopodiaceae (10 species), indicating that C₄ plants mainly occur in very few families (7 of 204) on the Tibetan Plateau. High altitude distribution for all the Chenopodiaceae C₄ species (> 3 000 m above sea level) suggests that plants of this kind have large tolerance to cold, dryness, and strong ultraviolet radiation. Most Gramineae and Cyperaceae C₄ species occurrences are consistent with extensive distribution of steppes and meadows in the vast flat of the central Plateau (1 000–3 000 m a.s.l.). Relatively high amount of hemicryptophyte form plants (44 %) in the region indicates that the vegetation, especially grassland, meadows, and steppe, are in good condition. There is a strong relationship between numbers of C₄ species and altitude in the Tibetan Plateau. Occurrence of C₄ species is significantly less in both high and low altitude plateaux in Tibet. Altitude distribution pattern for C₄ species in the region is not only consistent with the altitude and climate, but also with the vegetation types in altitude gradient.     

Photosynthetic and morphological functional types for native species from mixed prairie in Southern Saskatchewan, Canada

Photosynthetic pathways (e.g. C₃, C₄) and morphological functional types (e.g. trees, shrubs, high perennial grasses, perennial forbs) were identified for the native species from the Saskatchewan mixed prairie, using the data from references published between 1950 and 2003. Of the total 219 identified species in 145 genera and 45 families, 208 species in 137 genera and 44 families were found with C₃ photosynthesis, and most of these species are dominants (e.g. Agropyron dasystachyum Hook. and Stipa spartea var. curtiseta Hitchc.). 11 species in 10 genera and 3 families were identified with C₄ photosynthesis (e.g. Atriplex argentea Nutt., Andropogon scoparius Michx., Boutelou gracilis Lag., Calamovilfa longifolia Hook.). The amount of total identified C₄ species in the region is much less than that from the South Dakota mixed prairie (27 species). Gramineae is the leading family with C₄ photosynthesis (8 species), Chenopodiaceae ranks the second (2 species). Relatively less forb types [50 % perennial forbs (PEF) and 12 % annual forbs (ANF)] and more graminoid types (25 %) composition suggested that the rangelands in the region are relatively stable. Lacking of the knowledge on the optimal traits for PFTs classification in the region, further studies (e.g. C₃ and C₄ plant identification and optimal trait selection) are needed to explore the relationships between PFTs and vegetation variations, as well as land-use and climate changes.     

Co-function of C3-and C4-photosynthetic pathways in C3, C4 and C3-C4 intermediate Flaveria species

The potential for C₄ photosynthesis was investigated in five C₃-C₄ intermediate species, one C₃ species, and one C₄ species in the genus Flaveria, using ¹⁴CO₂ pulse-¹²CO₂ chase techniques and quantum-yield measurements. All five intermediate species were capable of incorporating ¹⁴CO₂ into the C₄ acids malate and aspartate, following an 8-s pulse. The proportion of ¹⁴C label in these C₄ products ranged from 50–55% to 20–26% in the C₃-C₄ intermediates F. floridana Johnston and F. linearis Lag. respectively. All of the intermediate species incorporated as much, or more, ¹⁴CO₂ into aspartate as into malate. Generally, about 5–15% of the initial label in these species appeared as other organic acids. There was variation in the capacity for C₄ photosynthesis among the intermediate species based on the apparent rate of conversion of ¹⁴C label from the C₄ cycle to the C₃ cycle. In intermediate species such as F. pubescens Rydb., F. ramosissima Klatt., and F. floridana we observed a substantial decrease in label of C₄-cycle products and an increase in percentage label in C₃-cycle products during chase periods with ¹²CO₂, although the rate of change was slower than in the C₄ species, F. palmeri. In these C₃-C₄ intermediates both sucrose and fumarate were predominant products after a 20-min chase period. In the C₃-C₄ intermediates, F. anomala Robinson and f. linearis we observed no significant decrease in the label of C₄-cycle products during a 3-min chase period and a slow turnover during a 20-min chase, indicating a lower level of functional integration between the C₄ and C₃ cycles in these species, relative to the other intermediates. Although F. cronquistii Powell was previously identified as a C₃ species, 7–18% of the initial label was in malate+aspartate. However, only 40–50% of this label was in the C-4 position, indicating C₄-acid formation as secondary products of photosynthesis in F. cronquistii. In 21% O₂, the absorbed quantum yields for CO₂ uptake (in mol CO₂·[mol quanta]^-1) averaged 0.053 in F. cronquistii (C₃), 0.051 in F. trinervia (Spreng.) Mohr (C₄), 0.052 in F. ramosissima (C₃-C₄), 0.051 in F. anomala (C₃-C₄), 0.050 in F. linearis (C₃-C₄), 0.046 in F. floridana (C₃-C₄), and 0.044 in F. pubescens (C₃-C₄). In 2% O₂ an enhancement of the quantum yield was observed in all of the C₃-C₄ intermediate species, ranging from 21% in F. ramosissima to 43% in F. pubescens. In all intermediates the quantum yields in 2% O₂ were intermediate in value to the C₃ and C₄ species, indicating a co-function of the C₃ and C₄ cycles in CO₂ assimilation. The low quantum-yield values for F. pubescens and F. floridana in 21% O₂ presumably reflect an ineffcient transfer of carbon from the C₄ to the C₃ cycle. The response of the quantum yield to four increasing O₂ concentrations (2–35%) showed lower levels of O₂ inhibition in the C₃-C₄ intermediate F. ramosissima, relative to the C₃ species. This indicates that the co-function of the C₃ and C₄ cycles in this intermediate species leads to an increased CO₂ concentration at the site of ribulose-1,5-bisphosphate carboxylase/oxygenase and a concomitant decrease in the competitive inhibition by O₂.Abbreviations PEP phosphoenolpyruvate - PGA 3-phosphoglycerate - RuBP ribulose-1,5-bisphosphate     

The quantum yield for CO2 uptake in C3 and C4 grasses

The quantum yield for CO₂ uptake was measured in C₃ and C₄ monocot species from several different grassland habitats. When the quantum yield was measured in the presence of 21% O₂ and 340 cm³ m^-3 CO₂, values were very similar in C₃ monocots, C₃ dicots, and C₄ monocots (0.045–0.056 mole CO₂ · mole^-1 quanta absorbed). In the presence of 2% O₂ and 800 cm³ m^-3 CO₂, enhancements of the quantum yield values occurred for the C₃ plants (both monocots and dicots), but not for C₄ monocots. A dependence of the quantum yield on leaf temperature was observed in the C₃ grass, Agropyron smithii, but not in the C₄ grass, Bouteloua gracilis, in 21% O₂ and 340 cm³ m^-3 CO₂. At leaf temperatures between 22–25°C the quantum yield values were approximately equal in the two species.     

Increasing CO2 and plant-plant interactions: effects on natural vegetation

Plant species and functional groups of species show marked differences in photosynthesis and growth in relation to rising atmospheric CO₂ concentrations through the range of the 30 % increase of the recent past and the 100 % increase since the last glaciation. A large shift was found in the compositional mix of 26 species of C₃'s and 17 species of C₄'s grown from a native soil seed bank in a competitive mode along a CO₂ gradient that approximated the CO₂ increase of the past 150 years and before. The biomass of C₃'s increased from near zero to 50 % of the total while that of the C₄'s was reduced 25 % as CO₂ levels approached current ambient. The proposition that acclimation to rising CO₂ will largely negate the fertilization effect of higher CO₂ levels on C₃'s is not supported. No signs of photosynthetic acclimation were evident forAvena sativa, Prosopis glandulosa, andSchizachyrium scoparium plants grown in subambient CO₂. The effects of changing CO₂ levels on vegetation since the last glaciation are thought to have been at least as great, if not greater, than those which should be expected for a doubling of current CO₂ levels. Atmospheric CO₂ concentrations below 200 ppm are thought to have been instrumental in the rise of the C₄ grasslands of North America and other extensive C₄ grasslands and savannas of the world. Dramatic invasion of these areas by woody C₃ species are accompanying the historical increase in atmospheric CO₂ concentration now in progress.     

Purification and properties of glycolate oxidase from plants with different photosynthetic pathways: Distinctness of C4 enzyme from that of a C3 species and a C3–C4 intermediate

Glycolate oxidase (GO; EC 1.1.3.1) was purified from the leaves of three plant species:Amaranthus hypochondriacus L.(NAD-ME type C₄ dicot),Pisum sativum L. (C₃ species) andParthenium hysterophorus L. (C₃–C₄. intermediate). A flavin moiety was present in the enzyme from all the three species. The enzyme from the C₄ plant had a low specific activity, exhibited lower K_M for glycolate, and required a lower pH for maximal activity, compared to the C₃ enzyme. The enzyme from the C₄ species oxidized glyoxylate at <10% of the rate with glycolate, while the GO from the C₃ plant oxidized glyoxylate at a rate of about 35 to 40% of that with glycolate. The sensitivity of GO from C₄ plant to -hydroxypyridinemethane sulfonate, 2-hydroxy-3-butynoate and other inhibitors was less than that of the enzyme from C₃ source. The properties of GO fromParthenium hysterophorus, were similar to those of the enzyme fromPisum sativum. The characteristics of glycolate oxidase from leaves of a C₄ plant,Amaranthus hypochondriacus are different from those of the C₃ species or the C₃–C₄ intermediate.     

Reassimilation of carbon dioxide by Flaveria (Asteraceae) species representing different types of photosynthesis

The capability to reassimilate CO₂ originating from intracellular decarboxylating processes connected with the photorespiratory glycolate pathway and-or decarboxylation of C₄ acids during C₄ photosynthesis has been investigated with four species of the genus Flaveria (Asteraceae). The C₃-C₄ intermediate species F. pubescens and F. anomala reassimilated CO₂ much more efficiently than the C₃ species F. cronquistii and, with respect to this feature, behaved similarly to the C₄ species F. trinervia. Therefore, under atmospheric conditions the intermediate species photorespired with rates only between 10–20% of that measured with F. cronquistii. At low oxygen concentrations (1,5%) the reassimilation potential of F. anomala approached that of F. trinervia and was distinct from that found with F. pubescens. The data are discussed with respect to a possible sequence of events during evolution of C₄ photosynthesis. If compared with related data for C₃-C₄ intermediate species from other genera they support the hypothesis that, during evolution of C₄ photosynthesis, an efficient capacity for CO₂ reassimilation evolved prior to a CO₂-concentrating mechanism.Abbreviations C₃, C₄ assimilated CO₂ initially found in 3-phosphoglycerate (C₃) or malate and aspartate (C₄) - D reassimilation coefficient - R _n , R _t net, total CO₂ evolution as measured with 0.03 and 3% CO₂, respectively - RuBP ribulose-1,5-bisphosphate - TPS true photosynthesis     

Exotic plant species in a C4-dominated grassland: invasibility, disturbance, and community structure

We used data from a 15-year experiment in a C₄-dominated grassland to address the effects of community structure (i.e., plant species richness, dominance) and disturbance on invasibility, as measured by abundance and richness of exotic species. Our specific objectives were to assess the temporal and spatial patterns of exotic plant species in a native grassland in Kansas (USA) and to determine the factors that control exotic species abundance and richness (i.e., invasibility). Exotic species (90% C₃ plants) comprised approximately 10% of the flora, and their turnover was relatively high (30%) over the 15-year period. We found that disturbances significantly affected the abundance and richness of exotic species. In particular, long-term annually burned watersheds had lower cover of exotic species than unburned watersheds, and fire reduced exotic species richness by 80–90%. Exotic and native species richness were positively correlated across sites subjected to different fire (r = 0.72) and grazing (r = 0.67) treatments, and the number of exotic species was lowest on sites with the highest productivity of C₄ grasses (i.e., high dominance). These results provide strong evidence for the role of community structure, as affected by disturbance, in determining invasibility of this grassland. Moreover, a significant positive relationship between exotic and native species richness was observed within a disturbance regime (annually burned sites, r = 0.51; unburned sites, r = 0.59). Thus, invasibility of this C₄-dominated grassland can also be directly related to community structure independent of disturbance. Received: 9 February 1999 / Accepted: 12 May 1999