A meta-analysis of cambium phenology and growth: linear and non-linear patterns in conifers of the northern hemisphere

Background and Aims Ongoing global warming has been implicated in shifting phenological patterns such as the timing and duration of the growing season across a wide variety of ecosystems. Linear models are routinely used to extrapolate these observed shifts in phenology into the future and to estimate changes in associated ecosystem properties such as net primary productivity. Yet, in nature, linear relationships may be special cases. Biological processes frequently follow more complex, non-linear patterns according to limiting factors that generate shifts and discontinuities, or contain thresholds beyond which responses change abruptly. This study investigates to what extent cambium phenology is associated with xylem growth and differentiation across conifer species of the northern hemisphere. Methods Xylem cell production is compared with the periods of cambial activity and cell differentiation assessed on a weekly time scale on histological sections of cambium and wood tissue collected from the stems of nine species in Canada and Europe over 1–9 years per site from 1998 to 2011. Key Results The dynamics of xylogenesis were surprisingly homogeneous among conifer species, although dispersions from the average were obviously observed. Within the range analysed, the relationships between the phenological timings were linear, with several slopes showing values close to or not statistically different from 1. The relationships between the phenological timings and cell production were distinctly non-linear, and involved an exponential pattern. Conclusions The trees adjust their phenological timings according to linear patterns. Thus, shifts of one phenological phase are associated with synchronous and comparable shifts of the successive phases. However, small increases in the duration of xylogenesis could correspond to a substantial increase in cell production. The findings suggest that the length of the growing season and the resulting amount of growth could respond differently to changes in environmental conditions.

From Broadstone to Zackenberg

Ecological networks are typically complex constructions of species and their interactions. During the last decade, the study of networks has moved from static to dynamic analyses, and has attained a deeper insight into their internal structure, heterogeneity, and temporal and spatial resolution. Here, we review, discuss and suggest research lines in the study of the spatio-temporal heterogeneity of networks and their hierarchical nature. We use case study data from two well-characterized model systems (the food web in Broadstone Stream in England and the pollination network at Zackenberg in Greenland), which are complemented with additional information from other studies. We focus upon eight topics: temporal dynamic space-for-time substitutions linkage constraints habitat borders network modularity individual-based networks invasions of networks and super networks that integrate different network types. Few studies have explicitly examined temporal change in networks, and we present examples that span from daily to decadal change: a common pattern that we see is a stable core surrounded by a group of dynamic, peripheral species, which, in pollinator networks enter the web via preferential linkage to the most generalist species. To some extent, temporal and spatial scales are interchangeable (i.e. networks exhibit ‘ergodicity’) and we explore how space-for-time substitutions can be used in the study of networks. Network structure is commonly constrained by phenological uncoupling (a temporal phenomenon), abundance, body size and population structure. Some potential links are never observed, that is they are ‘forbidden’ (fully constrained) or ‘missing’ (a sampling effect), and their absence can be just as ecologically significant as their presence. Spatial habitat borders can add heterogeneity to network structure, but their importance has rarely been studied: we explore how habitat generalization can be related to other resource dimensions. Many networks are hierarchically structured, with modules forming the basic building blocks, which can result in self-similarity. Scaling down from networks of species reveals another, finer-grained level of individual-based organization, the ecological consequences of which have yet to be fully explored. The few studies of individual-based ecological networks that are available suggest the potential for large intraspecific variance and, in the case of food webs, strong size-structuring. However, such data are still scarce and more studies are required to link individual-level and species-level networks. Invasions by alien species can be tracked by following the topological ‘career’ of the invader as it establishes itself within a network, with potentially important implications for conservation biology. Finally, by scaling up to a higher level of organization, it is possible to combine different network types (e.g. food webs and mutualistic networks) to form super networks, and this new approach has yet to be integrated into mainstream ecological research. We conclude by listing a set of research topics that we see as emerging candidates for ecological network studies in the near future.

The role of tree size in the leafing phenology of a seasonally dry tropical forest in Belize, Central America

Leafing phenology of two dry-forest sites on soils of different depth (S = shallow, D = deep) at Shipstern Reserve, Belize, were compared at the start of the rainy season (April-June 2000). Trees greater than or equal to 2.5 cm dbh were recorded weekly for 8 wk in three 0.04-ha plots per site. Ten species were analysed individually for their phenological patterns, of which the three most common were Bursera simaruba, Metopium brownei and Jatropha gaumeri. Trees were divided into those in the canopy (> 10 cm dbh) and the subcanopy (less than or equal to 10 cm dbh). Site S had larger trees on average than site D. The proportion of trees flushing leaves at any one time was generally higher in site S than in site D, for both canopy and subcanopy trees. Leaf flush started 2 wk earlier in site S than site D for subcanopy trees, but only 0.5 wk earlier for the canopy trees. Leaf flush duration was 1.5 wk longer in site S than site D. Large trees in the subcanopy flushed leaves earlier than small ones at both sites but in the canopy just at site D. Large trees flushed leaves earlier than small ones in three species and small trees flushed leaves more rapidly in two species. Bursera and Jatropha followed the general trends but Metopium, with larger trees in site D than site S, showed the converse with onset of flushing I wk earlier in site D than site S. Differences in response of the canopy and subcanopy trees on each site can be accounted for by the predominance of spring-flushing or stem-succulent species in site S and a tendency for evergreen species to occur in site D. Early flushing of relatively larger trees in site D most likely requires access to deeper soil water reserves but small and large trees utilize stored tree water in site S.

A 170 year spring phenology index of plants in eastern China

Extending phenological records into the past is essential for the understanding of past ecological change and evaluating the effects of climate change on ecosystems. A growing body of historical phenological information is now available for Europe, North America, and Asia. In East Asia, long-term phenological series are still relatively scarce. This study extracted plant phenological observations from old diaries in the period 1834–1962. A spring phenology index (SPI) for the modern period (1963–2009) was defined as the mean flowering time of three shrubs (first flowering of Amygdalus davidiana and Cercis chinensis, 50% of full flowering of Paeonia suffruticosa) according to the data availability. Applying calibrated transfer functions from the modern period to the historical data, we reconstructed a continuous SPI time series across eastern China from 1834 to 2009. In the recent 30 years, the SPI is 2.1–6.3 days earlier than during any other consecutive 30 year period before 1970. A moving linear trend analysis shows that the advancing trend of SPI over the past three decades reaches upward of 4.1 d/decade, which exceeds all previously observed trends in the past 30 year period. In addition, the SPI series correlates significantly with spring (February to April) temperatures in the study area, with an increase in spring temperature of 1°C inducing an earlier SPI by 3.1 days. These shifts of SPI provide important information regarding regional vegetation-climate relationships, and they are helpful to assess long term of climate change impacts on biophysical systems and biodiversity.

Spring-summer temperatures reconstructed for northern Switzerland and southwestern Germany from winter rye harvest dates, 1454–1970

This paper presents a unique 517-yr long documentary data-based reconstruction of spring-summer (MAMJJ) temperatures for northern Switzerland and southwestern Germany from 1454 to 1970. It is composed of 25 partial series of winter grain (secale cereale) harvest starting dates (WGHD) that are partly based on harvest related bookkeeping of institutions (hospitals, municipalities), partly on (early) phenological observations. The resulting main Basel WGHD series was homogenised with regard to dating style, data type and altitude. The calibration and verification approach was applied using the homogenous HISTALP temperature series from 1774–1824 for calibration (r = 0.78) and from 1920–1970 for verification (r = 0.75). The latter result even suffers from the weak data base available for 1870– 1950. Temperature reconstructions based on WGHD are more influenced by spring temperatures than those based on grape harvest dates (GHD), because rye in contrast to vines already begins to grow as soon as sunlight brings the plant to above freezing. The earliest and latest harvest dates were checked for consistency with narrative documentary weather reports. Comparisons with other European documentarybased GHD and WGHD temperature reconstructions generally reveal significant correlations decreasing with the distance from Switzerland. The new Basel WGHD series shows better skills in representing highly climate change sensitive variations of Swiss Alpine glaciers than available GHD series.

Eine tragfähige Datenbrücke – Phänologische Beobachtungen im Amtsblatt des Kantons Schaffhausen 1876–1950

The international standardisation of national meteorological networks in the late nineteenth century excluded biotic and abiotic observations from the objects to be henceforth published in the yearbooks. Skilled amateurs being in charge of three meteorological stations in Canton Schaffhausen (Switzerland) and their successors managed to continuously publish phenological observations gathered in the station environment alongside with meteorological data in the official gazette of this Canton from 1876 to 1950, i.e. up to the onset of phenological network observations in Switzerland. At least ten observations are available for 51 plant and animal phenological phases. Long series were assembled (N → = 30) for 14 plant phenological observations, among them for the first flowering of snowdrop (Galanthus nivalis), of hazel (Corylus avellana), of horse chestnut (Aesculus hippocastanum), of winter rye (Secale cereale) and of grape vine (Vitis vinifera) as well as the beginning of hay, winter rye and grape harvesting. Only the bare data were published without any metadata. The quality of 10 long series (N →=60) was checked by investigating the biographical and biological background of key observers and submitting their evidence to graphical (meteorological plausibility check of outliers) and statistical verification. The long term observers, mostly schoolteachers and high school professors, had a good knowledge of botany and the quality of their observations – disregarding obvious printing errors – is surprisingly good. A number of long series (seven) was completed with applicable data from the Swiss Phenological Network up to 2011. Besides anthropogenic shifts (beginning of hay and grape harvest) there is a contrast between a global warming-related earlier flowering of snowdrop and hazel and a later occurrence of grape vine flowering.

Are carbohydrate storage strategies of trees traceable by early–latewood carbon isotope differences?

Investigations of stable isotopes in early and latewood cellulose offer interesting insights to climate-driven adaptations of trees’ carbon storage strategy during different phenological phases.

Variation in life-history traits and their plasticities to elevational transplantation among seed families suggests potential for adaptative evolution of 15 tropical plant species to climate change

• Premise of the study: Because not all plant species will be able to move in response to global warming, adaptive evolution matters largely for plant persistence. As prerequisites for adaptive evolution, genetic variation in and selection on phenotypic traits are needed, but these aspects have not been studied in tropical species. We studied how plants respond to transplantation to different elevations on Mt. Kilimanjaro, Tanzania, and whether there is quantitative genetic (among-seed family) variation in and selection on life-history traits and their phenotypic plasticity to the different environments. • Methods: We reciprocally transplanted seed families of 15 common tropical, herbaceous species of the montane and savanna vegetation zone at Mt. Kilimanjaro to a watered experimental garden in the montane (1450 m) and in the savanna (880 m) zone at the mountain’s slope and measured performance, reproductive, and phenological traits. • Results: Plants generally performed worse in the savanna garden, indicating that the savanna climate was more stressful and thus that plants may suffer from future climate warming. We found significant quantitative genetic variation in all measured performance and reproductive traits in both gardens and for several measures of phenotypic plasticity in response to elevational transplantation. Moreover, we found positive selection on traits at low and intermediate trait values levelling to neutral or negative selection at high values. • Conclusions: We conclude that common plants at Mt. Kilimanjaro express quantitative genetic variation in fitness-relevant traits and in their plasticities, suggesting potential to adapt evolutionarily to future climate warming and increased temperature variability.

Genetic diversity and differentiation follow secondary succession in a multi-species study on woody plants from subtropical China

Aims: Species diversity and genetic diversity may be affected in parallel by similar environmental drivers. However, genetic diversity may also be affected independently by habitat characteristics. We aim at disentangling relationships between genetic diversity, species diversity and habitat characteristics of woody species in subtropical forest. Methods: We studied 11 dominant tree and shrub species in 27 plots in Gutianshan, China, and assessed their genetic diversity (Ar) and population differentiation (F’ST) with microsatellite markers. We tested if Ar and population specific F’ST were correlated to local species diversity and plot characteristics. Multi-model inference and model averaging were used to determine the relative importance of each predictor. Additionally we tested for isolation-by-distance and isolation-by-elevation by regressing pairwise F’ST against pairwise spatial and elevational distances. Important findings: Genetic diversity was not related to species diversity for any of the study species. Thus, our results do not support joint effects of habitat characteristics on these two levels of biodiversity. Instead, genetic diversity in two understory shrubs, Rhododendron simsii and Vaccinium carlesii, was affected by plot age with decreasing genetic diversity in successionally older plots. Population differentiation increased with plot age in Rhododendron simsii and Lithocarpus glaber. This shows that succession can reduce genetic diversity within, and increase genetic diversity between populations. Furthermore, we found four cases of isolation-by-distance and two cases of isolation-by-elevation. The former indicates inefficient pollen and seed dispersal by animals whereas the latter might be due to phenological asynchronies. These patterns indicate that succession can affect genetic diversity without parallel effects on species diversity and that gene flow in a continuous subtropical forest can be restricted even at a local scale.

Céspedes de estadios de fútbol : Mendoza (Argentina)

Se considera los estados de desarrollo de los céspedes de estadios de fútbol en Mendoza (Argentina). Desde un punto de vista fitosociológico se determinan dos grupos de plantas: Molinio-Arrhenatheretea R. Tx. 1937 (campos húmedos y pisoteados, con vegetación subnitrófila e higrófita) y Stellarietea mediae R. Tx. 1950 (vegetación arvense de los cultivos). Se indican las etapas dinámicas que se producen por sobrepisoteo y labores culturales. Del análisis se desprende que: 1. El sobrepisoteo y falta de cuidados culturales conducen a la pérdida de la cubierta vegetal y a la formación de peladeros sin vegetación. 2. La comunidad de Cynodon dactylon L., junto con elementos de la Molinio-Arrhenatheretea, es la más aceptada. Su cobertura representa el mejor estado del campo. 3. Adecuadas prácticas culturales asegurarían el mantenimiento y/o conservación de los céspedes favoreciendo la práctica del deporte y aportando belleza escénica.

Mapa bioclimático para las travesías de Mendoza (Argentina) basado en la fenología foliar

El objetivo fue generar un mapa bioclimático de la llanura de Mendoza que reflejara las diferencias climáticas expresadas por la actividad de la vegetación (fenología foliar) a escala regional. Se partió de la imagen digital del índice bioclimático de aridez P/ETP, generada en una etapa anterior a partir de una serie temporal de imágenes de índice verde (IVDN), y se recodificó en clases bioclimáticas. Se evaluó en cada clase la influencia antrópica y edáfica sobre las condiciones climáticas de aridez reflejadas por la vegetación. Se graficó la marcha fenológica anual media para cada bioclima a partir de una reconstrucción del IVDN. Las clases de clima húmedo y subhúmedo son de carácter edáfico debido al riego (oasis). Se proponen las clases: subdesértico (8,4%), árido inferior (15,3%), árido superior (24,2 %), semiárido inferior (25%) y semiárido superior (27,1%). Cada bioclima tiene una expresión vegetativa diferente en condiciones naturales. La marcha fenológica anual muestra que a mayor aridez menor es el contraste entre el IVDN mínimo y máximo, y que el momento de máxima cobertura vegetal varía de enero (semiárido) a abril (subdesértico). Esta propuesta permite extender y optimizar el conocimiento climático de las estaciones meteorológicas a través de toda la llanura mediante la expresión fenológica de la vegetación.

Modelización de la fenología del haba cv. Alameda

La predicción de la fenología del haba es un paso crítico para optimizar el manejo de los cultivos, así como para el desarrollo de modelos de cultivos. Este artículo examina la respuesta fenológica del cultivo de haba (Vicia faba L.) a diversos regímenes térmicos y fotoperíodicos evaluada mediante modelos lineales. Se realizaron diecisiete fechas de siembra durante tres años en Lugo, España (43°04’ N; 7°30’ W; altitud 480 m) en las que se hicieron observaciones fenológicas. El tiempo desde emergencia a floración se describe satisfactoriamente mediante un modelo fototérmico. Las tasas de desarrollo de siembra a emergencia, floración hasta la primera vaina y primera vaina hasta madurez fisiológica fueron modeladas en forma satisfactoria, utilizando solamente la temperatura como variable independiente. Los valores de temperaturas basales variaron entre 2,09 y 4,47°C, dependiendo del subperíodo fenológico. El fotoperíodo base resultó de 6,9 h mientras el fotoperíodo crítico fue de 16,2 h.

Evaluación de características de interés agronómico de damascos cv. Modesto

El cultivar de damascos Modesto es de introducción relativamente reciente en Mendoza (Argentina). El objetivo del trabajo fue evaluar las siguientes características de interés agronómico: la época de floración y la calidad de los frutos en cosecha y en postcosecha. El estudio se realizó en un monte comercial durante 2007 y 2008. Se registraron tres estados fenológicos: "D" (corola visible), "F" (flor abierta) y "H" (fruto cuajado). Se estableció el inicio de la floración, la plena floración y el fin de la misma. En los frutos las evaluaciones y mediciones se realizaron en el momento de la recolección y después del período de maduración de la fruta conservada en cámara frigorífica. Se determinó: color de fondo, peso, diámetro, firmeza de pulpa, contenido de sólidos solubles, pH, acidez titulable, presencia de hongos y desórdenes fisiológicos. Los resultados muestran que el cultivar Modesto florece en una época intermedia en la zona E de la provincia de Mendoza. La fecha de plena floración en los dos años del estudio fue el 8 de setiembre. En los estados de madurez evaluados los frutos reúnen los atributos de calidad demandados por el consumidor: alto CSS, color anaranjado en la madurez, buena firmeza de pulpa; sin embargo, el rápido, descenso de la firmeza durante la maduración en poscosecha obliga a una comercialización acelerada. Después del almacenamiento en cámara frigorífica durante treinta días y un período de maduración de dos días, la futa presentó el desorden harinosidad con una incidencia superior al 60%; esto señala que el cv. Modesto no puede almacenarse por un período tan prolongado.

Respuesta fotoperiódica en Origanum vulgare ssp. vulgare y ssp. hirtum Ietsw. : impacto sobre su desarrollo y crecimiento

La productividad del orégano (Origanum vulgare L.) está determinada por la conjunción entre cantidad de biomasa acumulada y contenido de aceite esencial hasta el momento de su cosecha. Numerosos autores han constatado que dicho contenido es máximo al momento de floración pero los procesos que determinan la ocurrencia de la misma son poco claros en esta especie. A través de la prolongación artificial del fotoperíodo, se evaluó la sensibilidad fotoperiódica de dos subespecies tradicionales de orégano (Compacto: Origanum vulgare ssp. vulgare y Criollo: Origanum vulgare ssp. hirtum Ietsw.) y su incidencia en el desarrollo, duración de fases fenológicas y en la dinámica de crecimiento. Se encontró que ambas responden al aumento del fotoperíodo reduciendo la longitud de su ciclo. Ante estas condiciones, las mismas difirieron en la magnitud de su respuesta, siendo la subespecie Criollo más sensible que Compacto. Esto sugiere que el umbral fotoperiódico de inducción a floración es menor en el orégano Criollo que en el orégano Compacto. El acortamiento de la fase de desarrollo vegetativo en ambas subespecies generó menor número de nudos y longitud de ramas finales (disminución más notoria en la subespecie Criollo). El fotoperíodo extendido generó un cambio en el modelo de crecimiento de la longitud de ramas de lineal a lineal con meseta o cuadrático.

Variabilidad en poblaciones de maíz nativo de la Mixteca Baja Oaxaqueña, México

La diversidad genética poblacional de maíz en México es muy dinámica y depende de factores biológicos, agroecológicos y socio-económicos, y necesidades familiares. En este trabajo se describió y clasificó la variabilidad morfológica de una colección de 60 muestras poblacionales de maíz, colectadas en 44 municipios de la Mixteca Baja Oaxaqueña (846 msnm a 1842 msnm). Las muestras se sembraron y cultivaron durante el ciclo primavera-verano de 2010, en Santo Domingo Tonala, Oaxaca, bajo un diseño de bloques completos al azar con cuatro repeticiones. Se evaluaron 19 caracteres morfológicos de planta, mazorca, grano y espiga (panoja), y se determinaron diferencias significativas entre poblacionales en estos caracteres. Los caracteres altura de planta y mazorca, días a floración masculina y femenina, y número de granos por hilera en la mazorca fueron determinantes para describir la variabilidad morfológica total. La variación morfológica y fenológica de las poblaciones de maíz se asocia con los patrones altitudinales y geográficos de donde proceden. Se determinaron seis grupos fenotípicos significativamente diferentes con características de mazorca, grano y planta semejantes a las descritas para las razas Celaya, Bolita, Pepitilla, Ancho, y ciertos complejos raciales entre Ancho, Mixteco, Celaya y Bolita.