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1.
Background and Aims: High elevation treelines occur worldwide at similar mean growing season temperatures. Does this result from direct impact of low temperature on growth or carbon metabolism, or does nutrient limitation, induced by low soil temperature, play a role? Similar treeline elevations at contrasting soil fertility argue against the latter, but the actual nutritional status of treeline trees (here addressed as foliage nitrogen concentration) has never been assessed systematically. Although needle nitrogen (N) concentration does not necessarily indicate growth limitation by N, the relative abundance of N would indicate obvious depletions at the treeline.

Methods: A central problem with any foliage nutrient assay is that the units for describing the element concentration are dependent on elevation themselves. Here we separate changes in N per unit tissue from changes in reference units.

Results: Needles of Pinus cembra and Picea abies in the Alps do not show elevational differences in N concentration per dry weight, water content, area or volume, thus, there is no N depletion near the elevational tree limit. Hence, nutrient supply is either unaltered, or growth is adjusted so that nutrient depletion in needle tissue does not occur.

Conclusions: Chronic N shortage at needle level is not an explanation for low tree vigour at the treeline.  相似文献   

2.
《Plant Ecology & Diversity》2013,6(3-4):365-375
Background: Temperature directly affects xylogenesis at high-elevation treelines. The low-temperature limitation of meristematic processes is thus key to understand treeline formation.

Aims: We aimed to experimentally test in situ the direct low-temperature effect on wood tissue formation at the alpine treeline.

Methods: We applied controlled Peltier-mediated cooling and warming (±3 K) to branch segments in Pinus uncinata at the treeline in the Swiss Alps. In addition, we studied xylogenesis in untreated trees during the growing season by sequential micro-coring.

Results: Micro-cores indicated that the cambial zone was fully developed by the time the cooling and warming treatment started, shortly after snowmelt. Presumably, because of this, experimental cooling of branches did not significantly reduce the number of cells produced per season. Warming extended the formation of early wood into the late season, and thus reduced the fraction of late wood.

Conclusions: We conclude that temperatures very early in the season determine the width of the cambial zone which, in turn, strongly controls the number of tracheids produced during the remaining growing season. Temperatures later in the season mainly determine the early wood to late wood ratio. These data provide an empirical basis for the mechanistic understanding of tree growth at the treeline in response to temperature.  相似文献   

3.
《Plant Ecology & Diversity》2013,6(3-4):329-342
Background: Climate, land-use and disturbance regimes are key drivers of treeline dynamics worldwide, but local and regional spatio-temporal patterns indicate that additional factors play an important role. Some studies suggest that shrub-tree interactions control tree seedling recruitment patterns across the treeline ecotone, but little is known about the generality of this interaction.

Aims: We established an experiment in a treeline ecotone in the central Pyrenees to investigate the role of such interactions and other environmental factors on tree seedling growth and survival. It was based on a similar experiment we completed recently in the subarctic Scandes. By comparing local phenomena between both experiments we assessed the generality of the findings across regions with different biogeographic histories and species characteristics.

Methods: We followed the survival and growth of transplanted Pinus uncinata seedlings during three growing seasons in a multi-factorial design (forest vs. treeline, +/– shrub removal, +/– temperature increase and +/– nutrient addition).

Results: There was better seedling growth at the treeline compared to the forest, and the presence of shrubs prevented seedling winter damage and herbivory. Both temperature and nutrient increase enhanced seedling performance.

Conclusions: Although some particular details were exclusive to the Pyrenees or to the Scandes, the similarities indicated general patterns and help to interpret the underlying mechanisms of treeline dynamics in both regions. This convergence of responses points to the potential of the development of a robust mechanistically founded predictive framework for scaling up shrub impacts on treeline dynamics to other regions.  相似文献   

4.
ABSTRACT

Background: Topoclimate can influence tree establishment within treeline ecotones. Yet much less is known about how regional topography, such as the Continental Divide, Rocky Mountains, mediates the role of climate in governing treeline dynamics.

Aims: To utilise the Continental Divide to test whether contrasts in growing-season moisture regimes to the west (summer-dry) and east (summer-wet) impact the spatio-temporal patterns of tree establishment and rates of treeline advance in the Northern Rocky Mountains.

Methods: We sampled trees at sites on north- and south-facing slopes, west and east of the Continental Divide. We used dendroecological techniques to reconstruct patterns of tree establishment. Age-structure data were quantitatively compared with climate to evaluate possible mechanistic linkages.

Results: Across all sites, 96% of trees established after 1950. There was a treeline advance (range = 39–140 m) accompanied by increases in tree density. Significantly more trees established during wet springs on both sides of the Divide.

Conclusions: Overall, snow duration in spring and autumn temperatures appear to influence patterns of tree recruitment at the treeline. Continued warming will likely amplify the role of autumn climate in regulating tree establishment throughout treeline ecotones in the Northern Rocky Mountains, particularly west of the Divide where summer-dry conditions persist.  相似文献   

5.
A world-wide study of high altitude treeline temperatures   总被引:43,自引:0,他引:43  
Aim At a coarse scale, the treelines of the world's mountains seem to follow a common isotherm, but the evidence for this has been indirect so far. Here we aim at underpinning this with facts. Location We present the results of a data‐logging campaign at 46 treeline sites between 68° N and 42° S. Methods We measured root‐zone temperatures with an hourly resolution over 1–3 years per site between 1996 and 2003. Results Disregarding taxon‐, landuse‐ or fire‐driven tree limits, high altitude climatic treelines are associated with a seasonal mean ground temperature of 6.7 °C (±0.8 SD; 2.2 K amplitude of means for different climatic zones), a surprisingly narrow range. Temperatures are higher (7–8 °C) in the temperate and Mediterranean zone treelines, and are lower in equatorial treelines (5–6 °C) and in the subarctic and boreal zone (6–7 °C). While air temperatures are higher than soil temperatures in warm periods, and are lower than soil temperatures in cold periods, daily means of air and soil temperature are almost the same at 6–7 °C, a physics driven coincidence with the global mean temperature at treeline. The length of the growing season, thermal extremes or thermal sums have no predictive value for treeline altitude on a global scale. Some Mediterranean (Fagus spp.) and temperate South Hemisphere treelines (Nothofagus spp.) and the native treeline in Hawaii (Metrosideros) are located at substantially higher isotherms and represent genus‐specific boundaries rather than boundaries of the life‐form tree. In seasonal climates, ground temperatures in winter (absolute minima) reflect local snow pack and seem uncritical. Main conclusions The data support the hypothesis of a common thermal threshold for forest growth at high elevation, but also reflect a moderate region and substantial taxonomic influence.  相似文献   

6.
Low temperatures are associated high‐altitude treelines, but the functional mechanism of treeline formation remains controversial. The relative contributions of carbon limitation (source activity) and growth limitation (sink activity) require more tests across taxa and regions. We examined temporal variations of mobile carbon supply in different tissues of Abies fargesii across treeline ecotones on north‐ and south‐facing slopes of the Qinling Mountains, China. Non‐structural carbohydrate (NSC) concentrations in tissues along the altitudinal gradient on both slopes changed significantly in the early and late growing season, but not in the mid‐growing season, indicating the season‐dependent carbon supply status. Late in the growing season on both slopes, trees at the upper limits had the highest NSC concentrations and total soluble sugars and lowest starch concentrations compared to trees at the lower elevations. NSC concentrations tended to increase in needles and branches throughout the growing season with increasing elevation on both slopes, but declined in roots and stems. NSC concentrations across sampling dates also indicated increases in needles and branches, and decreases in roots and stem with increasing elevation. Overall altitudinal trends of NSC in A. fargesii revealed no depletion of mobile carbon reserves at upper elevation limits, suggesting limitation of sink activity dominates tree life across treeline ecotones in both north‐ and south‐facing slopes. Carbon reserves in storage tissues (especially roots) in the late growing season might also play an important role in winter survival and early growth in spring at upper elevations on both slopes, which define the uppermost limit of A. fargesii.  相似文献   

7.

Key message

The treeline on Changbai Mountain controlled by low temperature and water stress, has not reached the position most commonly expected.

Abstract

Treeline pattern is an important consideration in exploring the general mechanisms controlling the response of treelines to climatic change. However, most of the present conclusions were derived from evergreen and/or conifer treeline, it is still not clear about the deciduous treeline. This study analyzed concentrations of non-structural carbohydrates (NSC) and their components (total soluble sugars and starch) in tree tissues of the deciduous species Erman’s birch (Betula ermanii) at four points along an elevational gradient ranging from 1,908–2,058 m a.s.l at the end of the growing season on Changbai Mountain in Northeast China. The mean 10-cm soil temperature of 8.2 °C under trees across the 129-day growing season at the treeline in this region was higher than that of the average threshold temperature found at treeline positions in the global and China’s climate studies. However, altitudinal trends of NSC concentrations increased significantly in all tissue types along the altitudinal gradients, revealing no depletion of carbon reserves at the treeline on Changbai Mountain. At the same time, the pronounced variation of δ13C in leaves and aged branches suggested that low temperature and water stress may simultaneously be operating at high altitudes to restrict the growth and NSC accumulation in trees above the treeline. In light of the above, we conclude that treeline formation on Changbai Mountain is no carbon depletion at the end of growing season, and most likely the result of sink limitation reflecting the combined effects of low temperature and water stress that determined the actual position of the treeline.  相似文献   

8.
Background: Recent work has shown little change in the position of the Smith fir treeline on the south-eastern Tibetan Plateau in response to global warming. However, the relationship between tree distribution patterns within the treeline ecotone and low responsiveness is unknown, and additional constraints than climate might be major drivers of these patterns (e.g. microsite availability for regeneration).

Aims: To characterise the spatial patterns of Smith fir alpine treelines and to infer the underlying processes driving their dynamics.

Methods: We investigated spatial patterns of Smith fir trees across two treeline ecotones in the Sygera Mountains, south-eastern Tibetan Plateau. The O(r)-ring statistic was used to analyse the univariate and bivariate spatial point patterns of three size classes (adults, juveniles and seedlings).

Results: Mature trees presented random spatial patterns. Clusters of juveniles and seedlings colonised areas not occupied by mature trees. Seedlings were clustered and established preferentially near juvenile firs, Rhododendron mats and over moss–lichen and organic matter substrates, indicating the importance of microsite availability for successful Smith fir recruitment.

Conclusions: Local factors such as microsite availability may play a major role in driving recent Smith fir treeline patterns and determine the lack of significant warming-induced upward shifts of these ecotones.  相似文献   

9.
Climate change is expected to increase woody vegetation abundance in the Arctic, yet the magnitude, spatial pattern and pathways of change remain uncertain. We compared historical orthophotos photos (1952 and 1979) with high-resolution satellite imagery (2015) to examine six decades of change in abundance of white spruce Picea glauca and tall shrubs (Salix spp., Alnus spp.) near the Agashashok River in northwest Alaska. We established ~3000 random points within our ~5500 ha study area for classification into nine cover types. To examine physiographic controls on tree abundance, we fit multinomial log-linear models with predictors derived from a digital elevation model and with arctic tundra, alpine tundra and ‘tree’ as levels of a categorical response variable. Between 1952 and 2015, points classified as arctic and alpine tundra decreased by 31% and 15%, respectively. Meanwhile, tall shrubs increased by 86%, trees mixed with tall shrubs increased by 385% and forest increased by 84%. Tundra with tall shrubs rarely transitioned to forest. The best multinomial model explained 71% of variation in cover and included elevation, slope and an interaction between slope and ‘northness’. Treeline was defined as the elevation where the probability of tree presence equaled that of tundra. Mean treeline elevation in 2015 was 202 m, corresponding with a June–August mean air temperature > 11°C, which is > 4°C warmer than the 6–7°C isotherm associated with global treeline elevations. Our results show dramatic increases in the abundance of trees and tall shrubs, question the universality of air temperature as a predictor of treeline elevation and suggest two mutually exclusive pathways of vegetation change, because tundra that gained tall shrubs rarely transitioned to forest. Conversion of tundra to tall shrubs and forest has important and potentially contrasting implications for carbon cycling, surface energy exchange and wildlife habitat in the Arctic.  相似文献   

10.
From 2001 to 2004 we experimentally warmed 40 large, naturally established, white spruce [Picea glauca (Moench) Voss] seedlings at alpine treeline in southwest Yukon, Canada, using passive open‐top chambers (OTCs) distributed equally between opposing north and south‐facing slopes. Our goal was to test the hypothesis that an increase in temperature consistent with global climate warming would elicit a positive growth response. OTCs increased growing season air temperatures by 1.8°C and annual growing degree‐days by one‐third. In response, warmed seedlings grew significantly taller and had higher photosynthetic rates compared with control seedlings. On the south aspect, soil temperatures averaged 1.0°C warmer and the snow‐free period was nearly 1 month longer. These seedlings grew longer branches and wider annual rings than seedlings on the north aspect, but had reduced Photosystem‐II efficiency and experienced higher winter needle mortality. The presence of OTCs tended to reduce winter dieback over the course of the experiment. These results indicate that climate warming will enhance vertical growth rates of young conifers, with implications for future changes to the structure and elevation of treeline contingent upon exposure‐related differences. Our results suggest that the growth of seedlings on north‐facing slopes is limited by low soil temperature in the presence of permafrost, while growth on south‐facing slopes appears limited by winter desiccation and cold‐induced photoinhibition.  相似文献   

11.
Background: Treeline ecotones represent environmental boundaries that fluctuate in space and time and thus induce changes in plant taxonomic and functional diversity.

Aims: To study changes through time in taxonomic and functional plant diversity patterns along the treeline ecotone.

Methods: In 2002, vegetation was sampled along a gradient from upper montane forest to the treeline–alpine transition in the South Ural Mountains, Russia. In 2014, vegetation was resampled and plant functional traits were collected. We studied spatial and temporal changes in plant species composition, functional composition and functional diversity.

Results: Species composition and diversity changed along the elevational gradient. The functional composition in height, leaf area, specific leaf area and leaf nitrogen content decreased with elevation, whereas functional composition of leaf carbon content increased. We found a temporal shift towards shorter plants with smaller leaves in treeline sites. Functional richness varied in several traits along the elevational gradient, while functional dispersion showed a trend towards increased functional dispersion in height, specific leaf area and leaf nitrogen in the treeline–tundra transition.

Conclusions: Tree encroachment across the treeline ecotone has resulted in a shift in plant species relative abundances and functional diversity, possibly affecting plant community assembly patterns.  相似文献   

12.
《Plant Ecology & Diversity》2013,6(3-4):383-392
Background: Arctic lichens and mosses are covered by snow for more than half the year and are generally considered as being dormant for most of this period. However, enhanced frequency of winter warming events due to climate change can cause increased disturbance of their protective subnivean environment.

Aim: To further understand cryptogamic responses to midwinter warming we compared the ecophysiological performance of one lichen and one moss species during a simulated warming event.

Methods: We measured photosynthesis and dark respiration in samples of the moss Hylocomium splendens and the lichen Peltigera aphthosa removed from under snow, and on natural refreezing after the warming event, which was simulated by using infrared heaters suspended above the ground.

Results: The moss exposed to light at +5 °C immediately after removal from their subnivean environment and from warmed plots showed positive net gas exchange within 332 s; the lichen required 1238 s. Photosynthesis and nitrogen fixation rates were equal to that, or higher than, during the preceding growing season. Upon refreezing after the event, moss photosynthesis declined considerably.

Conclusions: The moss, and to a lesser extent the lichen, may contribute to subnivean midwinter ecosystem respiration, and both are opportunistic, and can take advantage of warmer winter phases for photosynthesis and growth. This ought to be taken into account in vegetation change projections of cryptogam-rich ecosystems.  相似文献   

13.
Temperature is the most important factor affecting growth at high altitudes. As trees use much of the allocated carbon gained from photosynthesis to produce branches and stems, information on the timing and dynamics of secondary wood growth is crucial to assessing temperature thresholds for xylogenesis. We have carried out histological analyses to determine cambial activity and xylem cell differentiation in conifers growing at the treeline on the eastern Alps in two sites during 2002–2004 with the aim of linking the growth process with temperature and, consequently, of defining thresholds for xylogenesis. Cambial activity occurred from May to July–August and cell differentiation from May–June to September–October. The earliest start of radial enlargement was observed in stone pine in mid-May, while Norway spruce was the last species to begin tracheid differentiation. The duration of wood formation varied from 90 to 137 days, depending on year and site, with no difference between species. Longer durations were observed in trees on the south-facing site because of the earlier onset and later ending of cell production and differentiation. The threshold temperatures at which xylogenesis had a 0.5 probability of being active were calculated by logistic regressions. Xylogenesis was active when the mean daily air temperature was 5.6–8.5°C and mean stem temperature was 7.2–9°C. The similar thresholds among all trees suggested the existence of thermal limits in wood formation that correspond with temperatures of 6–8°C that are supposed to limit growth at the treeline. Different soil temperature thresholds between sites indicated that soil temperature may not be the main factor limiting xylogenesis. This study represents the first attempt to define a threshold through comparative assessment of xylem growth and tissue temperatures in stem meristems at high altitudes.  相似文献   

14.

Key message

The threshold minimum air temperature driving xylem growth of alpine  Rhododendron aganniphum is lower than that commonly observed at the treeline of conifers.

Abstract

Understanding how alpine shrubs grow and which environmental factors drive their biomass gain could help to functionally differentiate trees and shrubs. The cambium is the main meristem responsible for wood formation in trees and shrubs. Thus, a better knowledge of cambium growth dynamics in alpine shrubs would allow explaining why shrubs displace trees above the treeline. Here, we aim to investigate the timings and dynamics of xylogenesis and to identify the thermal thresholds controlling the onset of xylem growth of Rhododendron aganniphum, a tall shrub growing above the alpine treeline on the Tibetan Plateau. Timings of xylogenesis and radial growth rates were assessed from anatomical observations of the developing xylem during three growing seasons (2011, 2012, and 2013). The threshold temperature at which xylogenesis had a 0.5 probability of being active was calculated with logistic regressions. The onset of xylogenesis was observed between mid and late June, whereas the end of xylogenesis lasted from mid to late September. Overall, the duration of xylem growth lasted 88–101 days, and 94 % of the ring was formed from June to August. The threshold for the onset of xylem growth was observed at 2.0 ± 0.6 °C for the minimum air temperature, lower than that commonly observed for treeline conifers (ca. 6 °C). This low thermal threshold allows alpine shrubs to have a growing season long enough to complete xylem production and maturation during the warmest summer months. Our results suggest that the time required to complete xylogenesis is critical to understand why shrubs displace trees above the treeline.
  相似文献   

15.
Abstract

The resting metabolic rate (RMR) of Hemidactylus flaviviridis was measured at different temperatures from 20 to 35°C during winter and summer acclimatization. The mass exponent b values ranged between 0.67 and 0.72. Winter-acclimatized geckos of various body masses had significantly lower RMRs than summer-acclimatized geckos only at 20°C. It seems that low thermal sensitivity for summer–acclimatized group may facilitate activity during its active seasons, and high thermal sensitivity between 20 and 25°C for winter–acclimatized group may conserve energy during inactivity in winter.  相似文献   

16.
Abstract. Growth of coniferous trees at alpine treeline locations has traditionally been viewed as primarily temperature dependent. In this study, we use a physiologically mechanistic process model, ATE-BGC (Alpine Treeline Ecotone - BioGeochemical Cycles), to study the effects of a suite of environmental variables on the carbon balance of krummholz growth forms of subalpine Abies lasiocarpa under environmental conditions representative of treeline locations in Glacier National Park, Montana, USA. We perform a sensitivity analysis of ATE-BGC to determine the relative effects of temperature, insolation, snow, rain, soil depth, leaf area index, winter injury and elevation on krummholz carbon balance. Our results indicate that ATE-BGC carbon balance estimates are most highly influenced by temperature and winter injury. Leaf area index, soil depth, rainfall, insolation and snowfall follow temperature and winter injury in their effect on carbon balance. Due to the sensitivity of the model to moisture related variables, we propose that local irregularities in the location of the alpine treeline ecotone are the result of localized xeric conditions.  相似文献   

17.
Abstract

The day time periods of activity of Gazella dorcas dorcas L. in the Sudan are 06.00–10.00 and 14.00–18.00 hr in winter; 05.00–08.00 and 16.00–18.00 hr in summer when day temperatures may exceed 45° C. The main types of activity are described and observations made on periodicity in other types of behaviour viz: defaecation, chewing the cud, standing, lying, drinking, feeding, playing and courtship. These are all related to ambient temperature, time of day and season of year.  相似文献   

18.
A re-assessment of high elevation treeline positions and their explanation   总被引:67,自引:0,他引:67  
Christian Körner 《Oecologia》1998,115(4):445-459
In this review I first compile data for the worldwide position of climate-driven alpine treelines. Causes for treeline formation are then discussed with a global perspective. Available evidence suggests a combination of a general thermal boundary for tree growth, with regionally variable “modulatory” forces, including the presence of certain taxa. Much of the explanatory evidence found in the literature relates to these modulatory aspects at regional scales, whereas no good explanations emerged for the more fundamental global pattern related to temperature per se, on which this review is focused. I hypothesize that the life form “tree” is limited at treeline altitudes by the potential investment, rather than production, of assimilates (growth as such, rather than photosynthesis or the carbon balance, being limited). In shoots coupled to a cold atmosphere, meristem activity is suggested to be limited for much of the time, especially at night. By reducing soil heat flux during the growing season the forest canopy negatively affects root zone temperature. The lower threshold temperature for tissue growth and development appears to be higher than 3°C and lower than 10°C, possibly in the 5.5–7.5°C range, most commonly associated with seasonal means of air temperature at treeline positions. The physiological and developmental mechanisms responsible have yet to be analyzed. Root zone temperature, though largely unknown, is likely to be most critical. Received: 3 October 1997 / Accepted: 14 April 1998  相似文献   

19.
Infestation by rice and fruit bugs (Heteroptera) became a nationwide problem in Japan in the early 1970s. Nine rice bug species and three fruit bug species have been designated as economically important. Cropping restrictions for rice produced fallow paddy fields where various rice bugs reproduced and became abundant. Plautia crossota stali, Halyomorpha halys and Glaucias subpunctatus are dominant species of fruit bug that cause damage to a range of fruit crops. However, they require cones in order to complete their life cycle. Coniferous trees planted in the 1950s bore cones after 20 years. A dry and hot summer contributes to good masting the following year and good cone production, in turn, contributes to the abundance of fruit bugs in the third year. Thus, there is strong circumstantial evidence that land‐use changes were responsible for the abundance of both rice and fruit bugs during the last 30 years. Poleward range expansion was observed in Nezara viridula, Leptocorisa chinensis, G. subpunctatus and Paradasynus spinosus. A survey conducted to assess winter mortality revealed that every 1 °C rise in mean winter temperature resulted in a reduction of about 15% in winter mortality of N. viridula and H. halys in localities where the mean winter temperature ranges from 2 to 6 °C. In general, species with low developmental zero (T0) and small thermal constant (K) show the greatest increase in annual number of generations. Species with a high T0 for preoviposition period show the greatest increase in reproductive activity, while overwintering insects with a lower T0 tend to appear earlier in response to the elevation of temperature. Numbers of warnings issued by the prefectures on the occurrence of rice bugs and fruit bugs were correlated. Recent global warming operates in various ways, (e.g. by increasing annual number of generations, reproductive activity and food, to produce such correlation). There are no substantial bug problems in Korea where no significant land‐use changes have occurred. Rice bug outbreaks in Japan are predicted to become less frequent in the future, because there is no further scope for cropping restriction. Planting of coniferous trees has been continuously decreasing since 1970, but the area of coniferous forest is still almost the same. This suggests that the fruit bug problem will continue for the foreseeable future.  相似文献   

20.
Summary

In the southeast of Buenos Aires Province (Argentina), slugs cause reductions in crop yield by killing seeds and seedlings, by destroying stems and growing points, and by reducing leaf area. Deroceras reticulatum (Müller, 1774) was introduced into Argentina a long time ago, but the biology and ecology of this species in this country are still unknown. The aim of this study was to determine how temperature affects growth rate, reproduction and survival of D. reticulatum and also to assess the seasonal phenology of this species. The life cycle of D. reticulatum was investigated at 12°C, 20°C and at a temperature which alternated between 12°C for 16 h and 20°C for 8 h. The light/dark regime during the experiment was LD: 8/16 h. Slug populations were sampled from grassland located at the Experimental Station of INTA Balcarce. Eggs hatched after 16.3 to 39.8 days and fertility varied between 83.9% and 91.1%. The mean time from hatching to oviposition was longer and fecundity lower at 20°C than 12°C and 12/20°C (p <0.05). The net reproductive rate was 1.49 to 70.53 and the mean generation time varied between the 27 and 30 weeks. In natural conditions, eggs of D. reticulatum were found from early winter until the end of the spring, when soil temperature was 8–15°C and humidity was above 15%. These results indicate that D. reticulatum has one generation per year, univoltine phenology, with peak densities of slugs from the beginning of winter to the end of spring, which coincides with the sowing and emergence of winter and summer crops.  相似文献   

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