One Sentence Summary: Empirical evidence from grasslands around the world demonstrates a humped-back relationship between plant species richness and biomass at the 1 m 2 plot scale.Abstract: One of the central problems of ecology is the prediction of species diversity. The humped-back model (HBM) suggests that plant diversity is highest at intermediate levels of productivity; at low productivity few species can tolerate the environmental stresses and at high productivity a small number of highly competitive species dominate. A recent study claims to have comprehensively refuted the HBM. Here we show, using the largest, most geographically diverse dataset ever compiled and specifically built for testing this model that if the conditions are met, namely a wide range in biomass at the 1 m 2 plot level and the inclusion of plant litter, the relationship between plant biomass and species richness is hump shaped, supporting the HBM. Our findings shed new light on the prediction of plant diversity in grasslands, which is crucial for supporting management practices for effective conservation of biodiversity. 4Main Text: The relationship between plant diversity and productivity is a topic of intense debate (1-6). The HBM states that plant species richness peaks at intermediate productivity, taking above-ground biomass as a proxy for annual net primary productivity (ANPP) (7-9). This diversity peak is driven by two opposing processes; in unproductive and disturbed ecosystems where there is low plant biomass, species richness is limited by either stress, such as insufficient water and mineral nutrients, or high levels of disturbance-induced removal of biomass, which few species are able to tolerate. In contrast, in the low disturbance and productive conditions that generate high plant biomass it is competitive exclusion by a small number of highly competitive species that is hypothesized to constrain species richness (7-9). Other mechanisms proposed to explain the unimodal relationship between species richness and productivity include disturbance (10), evolutionary history and dispersal limitation (11,12), and density limitation affected by plant size (13).Different case studies have supported or rejected the HBM, and three separate meta-analyses reached different conclusions (14). This inconsistency may indicate a lack of generality of the HBM, or it may reflect a sensitivity to study characteristics including the type(s) of plant communities considered, the taxonomic scope, the length of the gradient sampled, the spatial grain and extent of analyses (14,15), and the particular measure of net primary productivity (16). Although others would argue (6), we maintain that the question remains whether the HBM serves as a useful and general model for grassland ecosystem theory and management. 5 We quantified the form and strength of the richness-productivity relationship using novel data from a globally-coordinated (17), distributed, scale-standardized and consistently designed survey, in which plant richness and biomass were m...

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RESUMO -Variáveis morfogênicas e estruturais foram quantificadas a partir da técnica de "perfilhos marcados" no período de agosto a outubro de 1999 numa pastagem de azevém anual, mantida em quatro diferentes alturas (5, 10, 15 e 20 cm) através de lotação contínua em carga variável com ovinos. O delineamento experimental foi o de blocos completos casualizados com três repetições. Utilizaram-se 50 perfilhos marcados por unidade experimental, distribuídos ao longo de transectas. As observações foram efetuadas em dois períodos: no primeiro, da entrada dos animais até a obtenção das alturas pretendidas, não se observaram diferenças entre os tratamentos. No segundo (fase de estabilização das alturas pretendidas), a taxa de elongação foliar, o comprimento da lâmina verde total por perfilho e o tamanho das folhas apresentaram resposta positiva frente a maiores alturas de manejo, enquanto o tempo de elongação da lâmina foliar diminui com a altura. As variáveis taxa de surgimento de folhas, tempo de vida das folhas, densidade populacional de perfilhos e número de folhas vivas por perfilho não foram afetadas pelos tratamentos.Palavras-chave: perfilhos marcados, altura, azevém, pastejo, ovinos Morphogenetic and Structural Traits of Ryegrass (Lolium multiflorum Lam.) Managed under Different Sward HeightsABSTRACT -Morphogenetic and structural variables were obtained from measurements performed on marked tillers and were quantified during August to October 1999 in a ryegrass pasture grazed by sheep under continuous stocking and variable stocking rate in order to maintain the sward at four different heights (5, 10, 15 and 20 cm). The experimental design was a randomized complete block with three replicates. Fifty marked tillers per experimental unit were assessed and distributed along transects lines. Sward measurements were performed in two periods: the first one, from the start of grazing until the point where the intended grazing heights were established, with no differences among treatments. In the second (after achieving the intended grazing heights), leaf elongation rate, total green leaf lamina length per tiller and leaf size showed a positive response to increasing sward heights, while the time to leaf lamina elongation decreased. The variables leaf appearance rate and leaf life span, tiller population density and number of live leaves per tiller were not affected by treatments.

RESUMO -Em pastagem de azevém anual sob lotação contínua com ovinos, quantificaram-se os fluxos de biomassa no período de 02 a 20 de outubro de 1999. Os tratamentos consistiram de quatro alturas (5, 10, 15 e 20 cm) de manejo, mantidas pelo uso de lotação variável. O delineamento experimental utilizado foi o de blocos casualizados com três repetições. Os fluxos foram determinados por meio de variáveis morfogênicas e estruturais obtidas a partir de medições feitas em afilhos marcados. No período de estabilização das alturas pretendidas, diferenças significativas na estrutura da pastagem resultaram em alterações nos fluxos de biomassa. Os fluxos de crescimento e de consumo apresentaram resposta quadrática às alturas de manejo, com valores máximos entre 10 e 15 cm, enquanto o fluxo de senescência aumentou linearmente com a altura. Máximos ganhos médios diários e ganhos por hectare foram alcançados nesse intervalo, indicando que essa faixa é apropriada para se manejar a pastagem. O período de avaliação coincidiu com a fase pós-indução floral, determinando diminuição na massa de lâminas e balanço negativo no fluxo de biomassa para todos os tratamentos.Palavras-chave: afilhos marcados, consumo, crescimento, morfogênese, ovinos, senescência Biomass Flows in Italian Ryegrass Pastures (Lolium multiflorum Lam.) Managed under Different Sward HeightsABSTRACT -In an italian ryegrass pasture under continuous stocking with sheep, biomass flows were estimated from 2 to 20 October 1999. Treatments corresponded to four different sward heights (5, 10, 15 and 20 cm) maintained by variable stocking. The experimental design was the randomized complete block design with three replicates. The fluxes were defined by morphogenetic and structural variables obtained from measurements performed on marked tillers. During the period in which the intended sward heights were established, significant effects in sward structure resulted in alterations on the biomass fluxes. The growth and intake fluxes showed a quadratic response with maximum values between 10 and 15 cm grazing heights, while the senescence flux increased linearly with sward height. Maximum daily live weight gain and animal yield per hectare were obtained within this interval, indicating this interval being the most appropriate to pasture management. The evaluation period was coincident with the floral induction phase, so the leaf lamina mass was progressively smaller and biomass fluxes showed a negative balance for all treatments.Key Words: intake, growth, marked tillers, morphogenesis, senescence, sheep R. Bras. Zootec., v.33, n.3, p.529-537 IntroduçãoO conhecimento do processo de morfogênese (dinâmica de geração e expansão da forma da planta no espaço) possibilita modelar o fluxo de biomassa foliar em nível de um afilho individual, através do crescimento, morte e colheita dos diferentes componentes vegetais (Davies, 1993). As taxas de alongamento foliar e de surgimento de folhas e o tempo de vida das folhas constituem os fatores morfogênicos do afilho que determinam o ritmo de...

Moderate grazing intensity is considered the basic requirement to enhance ecosystem function in grasslands. Yet, deterioration by overgrazing is common in many biomes, including Campos grasslands in South America. Understanding how grazing management can lead to recovery of ecosystem function is essential to design and implement effective strategies for sustainable use of this resource. In a long‐term field experiment carried out in Southern Brazil, we studied the effects of temporal grazing exclusions (spring or fall) at moderate and severe livestock grazing intensities (maintained by adjusting contrasting forage allowances) on the species richness, botanical composition, forage mass, sward height, and photosynthetic active radiation intercepted. The experiment was arranged in a completely randomized design with three replications of grazing exclusions, applied simultaneously at moderate and severe grazing intensities. Moderate grazing intensity showed a bimodal structure of shorter and taller canopies, and high species richness. Severe grazing created a shorter and homogeneous sward structure characterized by less standing biomass and species loss. In response to grazing exclusions, sward height, standing biomass, and light interception recovered almost to the levels of moderate grazing. Further, within 2 years grass species richness increased and botanical composition changed toward grasses with erect habit prevailing in moderate grazing intensity. Our study confirms that (1) moderate grazing intensities allow the coexistence of high number of species and (2) spring grazing exclusions of long‐term overgrazed grasslands can lead to a quick start to recover the grass species richness, primary productivity, and species composition like that prevailing in well‐managed grasslands.

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This work aimed to evaluate the following hypotheses: 1) the daily grazing time (GT) and 2) forage searching are more associated with the sward structure than with the levels of daily forage allowance (FA). To this end we proposed a model that was tested through an analysis of the sward structure, grazing time, and displacement in grazing by heifers on the natural grassland of the Pampa Biome (southern Brazil), which has been managed by FA levels since 1986. For three seasons, between January 2009 and February 2010, we evaluated the effect of FA on the main descriptors of the sward structure (herbage mass, sward height, and tussocks frequency) and the effect of these on the GT, displacement rate (DR), and daily displacement (D) in grazing. The data were analyzed with the use of regression and descriptive analyses from three-dimensional contour graphs with the data of the sward structure and GT. The DR was not associated with the FA levels or sward structure; however, the DR presented a positive linear relationship with the D and GT. The incremental change in the GT was accompanied by an increase in the D. Lastly, independently of the level of the FA and season evaluated, the lower values of GT were always associated with the following structural configuration: forage mass between 1 400 and 2 200 kg DM Á ha À1 , sward height between 9 and 13 cm, and tussock levels not exceeding 35%. Outside these limits, a penalty occurred in the GT and displacement patterns of the heifers. We found evidence that a better understanding of the cause-effect relationships between the sward structure and the ingestive behavior of the animals demonstrates the possibility of increasing the performance of domestic herbivores with important economic and ecological consequences. Resumen El objetivo del estudio fue evaluar las siguientes hipó tesis: (i) si el tiempo de pastoreo diario (TP) y (ii) la búsqueda de forraje está n má s estrechamente relacionados a la estructura del pasto que a los niveles diarios de oferta de forraje (OF). Con este fin, propusimos un modelo que se puso a prueba en base al aná lisis de la estructura del pasto, el tiempo de pastoreo y el desplazamiento en pastoreo en terneras sobre un pastizal natural del Bioma Pampa (sur de Brasil) que, desde 1986, se ha manejado con distintos niveles de OF. En tresépocas, entre Ene/2009 y Feb/2010, se evaluó el efecto de la OF sobre los principales descriptores de la estructura del pasto (biomasa de forraje, altura y frecuencia de matas) y el efecto deéstos sobre el tiempo de pastoreo (TP), la tasa de desplazamiento (TD) y el desplazamiento diario (D). Los datos fueron analizados mediante regresió n y por aná lisis descriptivos a partir de grá ficos de contorno tridimensionales en base a los datos de estructura del pasto y TP. La TD no tuvo relació n con OF ni con la estructura del pasto, pero mostró una relació n lineal positiva con D. Incrementos en TP estuvieron asociados a incrementos en D. El estudio demostró la importancia de la estructura del pasto al constatar que, indep...