Impacts of changing climate and land use on vegetation dynamics in a Mediterranean ecosystem: insights from paleoecology and dynamic modeling

Forests near the Mediterranean coast have been shaped by millennia of human disturbance. Consequently, ecological studies relying on modern observations or historical records may have difficulty assessing natural vegetation dynamics under current and future climate. We combined a sedimentary pollen record from Lago di Massacciucoli, Tuscany, Italy with simulations from the LandClim dynamic vegetation model to determine what vegetation preceded intense human disturbance, how past changes in vegetation relate to fire and browsing, and the potential of an extinct vegetation type under present climate. We simulated vegetation dynamics near Lago di Massaciucoli for the last 7,000 years using a local chironomid-inferred temperature reconstruction with combinations of three fire regimes (small infrequent, large infrequent, small frequent) and three browsing intensities (no browsing, light browsing, and moderate browsing), and compared model output to pollen data. Simulations with low disturbance support pollen-inferred evidence for a mixed forest dominated by Quercus ilex (a Mediterranean species) and Abies alba (a montane species). Whereas pollen data record the collapse of A. alba after 6000 cal yr bp, simulated populations expanded with declining summer temperatures during the late Holocene. Simulations with increased fire and browsing are consistent with evidence for expansion by deciduous species after A. alba collapsed. According to our combined paleo-environmental and modeling evidence, mixed Q. ilex and A. alba forests remain possible with current climate and limited disturbance, and provide a viable management objective for ecosystems near the Mediterranean coast and in regions that are expected to experience a mediterranean-type climate in the future.

Evidence for past variations in methane availability in a Siberian thermokarst lake based on δ13C of chitinous invertebrate remains

Understanding past methane dynamics in arctic wetlands and lakes is crucial for estimating future methane release. Methane fluxes from lake ecosystems have increasingly been studied, yet only few reconstructions of past methane emissions from lakes are available. In this study, we develop an approach to assess changes in methane availability in lakes based on δ13C of chitinous invertebrate remains and apply this to a sediment record from a Siberian thermokarst lake. Diffusive methane fluxes from the surface of ten newly sampled Siberian lakes and seven previously studied Swedish lakes were compared to taxon-specific δ13C values of invertebrate remains from lake surface sediments to investigate whether these invertebrates assimilated 13C-depleted carbon typical for methane. Remains of chironomid larvae of the tribe Orthocladiinae that, in the study lakes, mainly assimilate plant-derived carbon had higher δ13C than other invertebrate groups. δ13C of other invertebrates such as several chironomid groups (Chironomus, Chironomini, Tanytarsini, and Tanypodinae), cladocerans (Daphnia), and ostracods were generally lower. δ13C of Chironomini and Daphnia, and to a lesser extent Tanytarsini was variable in the lakes and lower at sites with higher diffusive methane fluxes. δ13C of Chironomini, Tanytarsini, and Daphnia were correlated significantly with diffusive methane flux in the combined Siberian and Swedish dataset (r = −0.72, p = 0.001, r = −0.53, p = 0.03, and r = −0.81, p < 0.001, respectively), suggesting that δ13C in these invertebrates was affected by methane availability. In a second step, we measured δ13C of invertebrate remains from a sediment record of Lake S1, a shallow thermokarst lake in northeast Siberia. In this record, covering the past ca 1000 years, δ13C of taxa most sensitive to methane availability (Chironomini, Tanytarsini, and Daphnia) was lowest in sediments deposited from ca AD 1250 to ca AD 1500, and after AD 1970, coinciding with warmer climate as indicated by an independent local temperature record. As a consequence the offset in δ13C between methane-sensitive taxa and bulk organic matter was higher in these sections than in other parts of the core. In contrast, δ13C of other invertebrate taxa did not show this trend. Our results suggest higher methane availability in the study lake during warmer periods and that thermokarst lakes can respond dynamically in their methane output to changing environmental conditions.

Climatic and environmental changes during the Weichselian Lateglacial Interstadial in the Weerterbos region, the Netherlands

Lake sediment records from the Weerterbos region, in the southern Netherlands, were studied to reconstruct summer temperature and environmental changes during the Weichselian Lateglacial Interstadial. A sediment core obtained from a small lacustrine basin was analysed for multiple proxies, including lithological changes, oxygen isotopes of bulk carbonates, pollen and chironomids. It was found that the oxygen isotope record differed strongly from the other proxies. Based on a comparison with three additional lake sediment records from the same region, it emerged that the oxygen isotope records were strongly affected by local environmental conditions, impeding the distinction of a regional palaeoclimate signal. The chironomid-inferred July air temperature reconstruction produced inferred interstadial temperatures ranging between ∼15° and 18°C, largely consistent with previously published results from the northern part of the Netherlands. A temporary regressive phase in the pollen record, which can be tentatively correlated with the Older Dryas, preceded the expansion of birch woodland. Despite differences between the four pollen records from the Weerterbos region, a comparable regressive vegetation phase that was possibly the result of a shift to drier conditions could be discerned in all of the profiles. In addition, a temporary temperature decline of ∼1.5°C was inferred from the chironomid record during this regressive phase. The multi-proxy approach used here enabled a direct comparison of inferred changes in temperature, vegetation and environmental conditions at an individual site, while the multi-site approach provided insight into the factors influencing the pollen and isotope records from these small-scale depressions.

New data on the Lateglacial period of SW Europe: a high resolution multiproxy record from Laguna de la Roya (NW Iberia)

High-resolution multiproxy analyses were performed on a 128 cm section of organic sediments accumulated in a small mountain lake in NW Iberia (Laguna de la Roya, 1608 m asl). The pollen stratigraphy together with radiocarbon dating provided the basis for a chronology ranging from 15,600 to 10,500 cal yr BP. Chironomid-inferred July air temperatures suggest a temperature range from 7 to 13 degrees C, also evidencing two well-established cold periods which may be equivalent to the INTIMATE stages GS-2a and GS-1. Furthermore, a number of short cold events (with summer temperatures dropping about 0.5-1 degrees C) appear intercalated within the Lateglacial Interstadial (possibly equivalent to the INTIMATE cold events GI-1d, GI-1c2 and GI-1b) and the early Holocene (possibly equivalent to the 11.2 k event). The temperature variations predicted by our reconstruction allow explaining the changes in local conditions and productivity of the lake inferred from the biological record of the same sediment core. Furthermore, they also agree with the local and regional vegetation dynamics, and the main oscillations deduced for the vegetation belts. Based on its chronology our multiproxy record indicates a similar temperature development in NW Iberia as inferred by the Greenland delta O-18 record, the marine deep-sea records off the Atlantic Iberian Margin, and other chironomid-based Lateglacial temperature reconstructions from Europe. Nevertheless, the impact of most of the less intense Lateglacial/early Holocene cold events in NW Iberia was most probably limited to very sensitive sites that were very close to ecotonal situations. Particularly, our new pollen record indicates that they were represented as three minor environmental crises occurring during the Lateglacial Interestadial in this area. The Older Dryas event (in our usage corresponding to the Aegelsee Oscillation in Central Europe and event GI-1d in central Greenland) has previously been described in this region, but its age and duration (ca 14,250-14050 cal yr BP) is now better constrained. The two subsequent stages, La Roya I (ca 13,600-13,400 cal yr BP) and La Roya II (ca 13,300-12,900 cal yr BP) have been described for first time in NW Iberia. (C) 2013 Elsevier Ltd. All rights reserved.

A model-data comparison of Holocene timberline changes in the Swiss Alps reveals past and future drivers of mountain forest dynamics

Mountain vegetation is strongly affected by temperature and is expected to shift upwards with climate change. Dynamic vegetation models are often used to assess the impact of climate on vegetation and model output can be compared with paleobotanical data as a reality check. Recent paleoecological studies have revealed regional variation in the upward shift of timberlines in the Northern and Central European Alps in response to rapid warming at the Younger Dryas/Preboreal transition ca. 11700years ago, probably caused by a climatic gradient across the Alps. This contrasts with previous studies that successfully simulated the early Holocene afforestation in the (warmer) Central Alps with a chironomid-inferred temperature reconstruction from the (colder) Northern Alps. We use LandClim, a dynamic landscape vegetation model to simulate mountain forests under different temperature, soil and precipitation scenarios around Iffigsee (2065m a.s.l.) a lake in the Northwestern Swiss Alps, and compare the model output with the paleobotanical records. The model clearly overestimates the upward shift of timberline in a climate scenario that applies chironomid-inferred July-temperature anomalies to all months. However, forest establishment at 9800 cal. BP at Iffigsee is successfully simulated with lower moisture availability and monthly temperatures corrected for stronger seasonality during the early Holocene. The model-data comparison reveals a contraction in the realized niche of Abies alba due to the prominent role of anthropogenic disturbance after ca. 5000 cal. BP, which has important implications for species distribution models (SDMs) that rely on equilibrium with climate and niche stability. Under future climate projections, LandClim indicates a rapid upward shift of mountain vegetation belts by ca. 500m and treeline positions of ca. 2500m a.s.l. by the end of this century. Resulting biodiversity losses in the alpine vegetation belt might be mitigated with low-impact pastoralism to preserve species-rich alpine meadows.

Taxon-specific δ13C analysis of chitinous invertebrate remains in sediments from Strandsjön, Sweden

Taxon-specific stable carbon isotope (δ13C) analysis of chitinous remains of invertebrates can provide valuable information about the carbon sources used by invertebrates living in specific habitats of lake ecosystems (for example, sediments, water column, or aquatic vegetation). This is complementary to δ13C of sedimentary organic matter (SOM), which provides an integrated signal of organic matter produced in a lake and its catchment, and of diagenetic processes within sediments. In a sediment record from Strandsjön (Sweden) covering the past circa 140 years, we analyzed SOM geochemistry (δ13C, C:Natomic, organic carbon content) and δ13C of chitinous invertebrate remains in order to examine whether taxon-specific δ13C records could be developed for different invertebrate groups and whether these analyses provide insights into past changes of organic carbon sources for lacustrine invertebrates available in benthic and planktonic compartments of the lake. Invertebrate taxa included benthic chironomids (Chironomus, Chironomini excluding Chironomus, Tanytarsini, and Tanypodinae), filter-feeders on suspended particulate organic matter (Daphnia, Plumatella and Cristatella mucedo), and Rhabdocoela. δ13C of chironomid remains indicated periodic availability of 13C-depleted carbon sources in the benthic environment of the lake as δ13C values of the different chironomid taxa fluctuated simultaneously between -34.7 and -30.5‰ (VPDB). Daphnia and Bryozoa showed parallel changes in their δ13C values which did not coincide with variations in δ13C of chironomids, though, and a 2-3‰ decrease since circa AD 1960. The decrease in δ13C of Daphnia and Bryozoa could indicate a decrease in phytoplankton δ13C as a result of lower lake productivity, which is in accordance with historical information about the lake that suggests a shift to less eutrophic conditions after AD 1960. In contrast, Rhabdocoela cocoons were characterized by relatively high δ13C values (-30.4 to -28.2‰) that did not show a strong temporal trend, which could be related to the predatory feeding mode and wide prey spectrum of this organism group. The taxon-specific δ13C analyses of invertebrate remains indicated that different carbon sources were available for the benthic chironomid larvae than for the filter-feeding Daphnia and bryozoans. Our results therefore demonstrate that taxon-specific analysis of δ13C of organic invertebrate remains can provide complementary information to measurements on bulk SOM and that δ13C of invertebrate remains may allow the reconstruction of past changes in carbon sources and their δ13C in different habitats of lake ecosystems.

Validation of climate model-inferred regional temperature change for late-glacial Europe

Comparisons of climate model hindcasts with independent proxy data are essential for assessing model performance in non-analogue situations. However, standardized palaeoclimate data sets for assessing the spatial pattern of past climatic change across continents are lacking for some of the most dynamic episodes of Earth’s recent past. Here we present a new chironomid-based palaeotemperature dataset designed to assess climate model hindcasts of regional summer temperature change in Europe during the late-glacial and early Holocene. Latitudinal and longitudinal patterns of inferred temperature change are in excellent agreement with simulations by the ECHAM-4 model, implying that atmospheric general circulation models like ECHAM-4 can successfully predict regionally diverging temperature trends in Europe, even when conditions differ significantly from present. However, ECHAM-4 infers larger amplitudes of change and higher temperatures during warm phases than our palaeotemperature estimates, suggesting that this and similar models may overestimate past and potentially also future summer temperature changes in Europe.

Late-Holocene climate variability and ecosystem responses in Alaska inferred from high-resolution multiproxy sediment analyses at Grizzly Lake

The late-Holocene shift from Picea glauca (white spruce) to Picea mariana (black spruce) forests marked the establishment of modern boreal forests in Alaska. To understand the patterns and drivers of this vegetational change and the associated late-Holocene environmental dynamics, we analyzed radiocarbon-dated sediments from Grizzly Lake for chironomids, diatoms, pollen, macrofossils, charcoal, element composition, particle size, and magnetic properties for the period 4100–1800 cal BP. Chironomid assemblages reveal two episodes of decreased July temperature, at ca. 3300–3150 (ca −1 °C) and 2900–2550 cal BP (ca −2 °C). These episodes coincided with climate change elsewhere in the Northern Hemisphere, atmospheric reorganization, and low solar activity. Diatom-inferred lake levels dropped by ca. 5 m at 3200 cal BP, suggesting dry conditions during the period 3200–1800 cal BP. P. glauca declined and P. mariana expanded at ca. 3200 cal BP; this vegetational change was linked to diatom-inferred low lake levels and thus decreased moisture availability. Forest cover declined at 3300–3100, 2800–2500 and 2300–2100 cal BP and soil erosion as inferred from increased values of Al, K, Si, Ti, and Ca intensified, when solar irradiance was low. Plant taxa adapted to disturbance and cold climate (e.g. Alnus viridis, shrub Betula, Epilobium) expanded during these periods of reduced forest cover. This open vegetation type was associated with high fire activity that peaked at 2800 cal BP, when climatic conditions were particularly cold and dry. Forest recovery lagged behind subsequent climate warming (≤+3 °C) by ca. 75–225 years. Our multiproxy data set suggests that P. glauca was dominant under warm-moist climatic conditions, whereas P. mariana prevailed under cold-dry and warm-dry conditions. This pattern implies that climatic warming, as anticipated for this century, may promote P. glauca expansions, if moisture availability will be sufficiently high, while P. mariana may expand under dry conditions, possibly exacerbating climate impacts on the fire regime.

The sediment record of the past 200 years in a Swiss high-alpine lake: Hagelseewli (2339 ma.s.l.)

Sediment cores spanning the last two centuries were taken in Hagelseewli, a high-elevation lake in the Swiss Alps. Contiguous 0.5 cm samples were analysed for biological remains, including diatoms, chironomids, cladocera, chrysophyte cysts, and fossil pigments. In addition, sedimentological and geochemical variables such as loss-on-ignition, total carbon, nitrogen, sulphur, grain-size and magnetic mineralogy were determined. The results of these analyses were compared to a long instrumental air temperature record that was adapted to the elevation of Hagelseewli by applying mean monthly lapse rates. During much of the time, the lake is in the shadow of a high cliff to the south, so that the lake is ice-covered during much of the year and thus decoupled from climatic forcing. Lake biology is therefore influenced more by the duration of ice-cover than by direct temperature effects during the short open-water season. Long periods of ice-cover result in anoxic water conditions and dissolution of authigenic calcites, leading to carbonate-free sediments. The diversity of chironomid and cladoceran assemblages is extremely low, whereas that of diatom and chrysophyte cyst assemblages is much higher. Weak correlations were observed between the diatom and chrysophyte cyst assemblages on the one hand and summer or autumn air temperatures on the other, but the proportion of variance explained is low, so that air temperature alone cannot account for the degree of variation observed in the paleolimnological record. Analyses of mineral magnetic parameters, spheroidal carbonaceous particles and lead suggest that atmospheric pollution has had a significant effect on the sediments of Hagelseewli, but little effect on the water quality as reflected in the lake biota.

Evidence for cooler European summers during periods of changing meltwater flux to the North Atlantic

We analyzed fossil chironomids (nonbiting midges) and pollen in two lake-sediment records to reconstruct and quantify Holocene summer-temperature fluctuations in the European Alps. Chironomid and pollen records indicate five centennial-scale cooling episodes during the early- and mid-Holocene. The strongest temperature declines of ≈1°C are inferred at ≈10,700–10,500 and 8,200–7,600 calibrated 14C years B.P., whereas other temperature fluctuations are of smaller amplitude. Two forcing mechanisms have been presented recently to explain centennial-scale climate variability in Europe during the early- and mid-Holocene, both involving changes in Atlantic thermohaline circulation. In the first mechanism, changes in meltwater flux from the North American continent to the North Atlantic are responsible for changes in the Atlantic thermohaline circulation, thereby affecting circum-Atlantic climate. In the second mechanism, solar variability is the cause of Holocene climatic fluctuations, possibly triggering changes in Atlantic thermohaline overturning. Within their dating uncertainty, the two major cooling periods in the European Alps are coeval with substantial changes in the routing of North American freshwater runoff to the North Atlantic, whereas quantitatively, our climatic reconstructions show a poor agreement with available records of past solar activity. Thus, our results suggest that, during the early- and mid-Holocene, freshwater-induced Atlantic circulation changes had stronger influence on Alpine summer temperatures than solar variability and that Holocene thermohaline circulation reductions have led to summer-temperature declines of up to 1°C in central Europe.

Effect of low count sums on quantitative environmental reconstructions: an example using subfossil chironomids

The concentrations of chironomid remains in lake sediments are very variable and, therefore, chironomid stratigraphies often include samples with a low number of counts. Thus, the effect of low count sums on reconstructed temperatures is an important issue when applying chironomid‐temperature inference models. Using an existing data set, we simulated low count sums by randomly picking subsets of head capsules from surface‐sediment samples with a high number of specimens. Subsequently, a chironomid‐temperature inference model was used to assess how the inferred temperatures are affected by low counts. The simulations indicate that the variability of inferred temperatures increases progressively with decreasing count sums. At counts below 50 specimens, a further reduction in count sum can cause a disproportionate increase in the variation of inferred temperatures, whereas at higher count sums the inferences are more stable. Furthermore, low count samples may consistently infer too low or too high temperatures and, therefore, produce a systematic error in a reconstruction. Smoothing reconstructed temperatures downcore is proposed as a possible way to compensate for the high variability due to low count sums. By combining adjacent samples in a stratigraphy, to produce samples of a more reliable size, it is possible to assess if low counts cause a systematic error in inferred temperatures.

Sedimentological and biostratigraphical analyses of short sediment cores from Hagelseewli (2339 m a.s.l.) in the Swiss Alps

Several short sediment cores of between 35 and 40 cm from Hagelseewli, a small, remote lake in the Swiss Alps at an elevation of 2339 m a.s.l. were correlated according to their organic matter content. The sediments are characterized by organic silts and show in their uppermost part a surprisingly high amount of organic matter (30-35%). Synchronous changes, occurring in pollen from snow-bed vegetation, the alga Pediastrum, chironomids, and grain-size composition, point to a climatic change interpreted as cooler or shorter summers that led to prolonged ice-cover on the lake. According to palynological results the sediments date back to at least the early 15th century A.D., with the cooling phase encompassing the period between late 16th and the mid-19th century thus coinciding with the Little Ice Age. Low concentrations of both chironomid head capsules and cladoceran remains in combination with results from fossil pigment analyses point to longer periods of bottom-water anoxia as a result of long-lasting ice-cover that prevented mixing of the water column. According to our results aquatic biota in Hagelseewli are mainly indirectly influenced by climate change. The duration of ice-cover on the lake controls the mixing of the water column as well as light-availability for phytoplankton blooms.

Modeling postglacial vegetation dynamics of temperate forests on the Olympic Peninsula (WA, USA) with special regard to snowpack

Past and future forest composition and distribution in temperate mountain ranges is strongly influenced by temperature and snowpack. We used LANDCLIM, a spatially explicit, dynamic vegetation model, to simulate forest dynamics for the last 16,000 years and compared the simulation results to pollen and macrofossil records at five sites on the Olympic Peninsula (Washington, USA). To address the hydrological effects of climate-driven variations in snowpack on simulated forest dynamics, we added a simple snow accumulation-and-melt module to the vegetation model and compared simulations with and without the module. LANDCLIM produced realistic present-day species composition with respect to elevation and precipitation gradients. Over the last 16,000 years, simulations driven by transient climate data from an atmosphere-ocean general circulation model (AOGCM) and by a chironomid-based temperature reconstruction captured Late-glacial to Late Holocene transitions in forest communities. Overall, the reconstruction-driven vegetation simulations matched observed vegetation changes better than the AOGCM-driven simulations. This study also indicates that forest composition is very sensitive to snowpack-mediated changes in soil moisture. Simulations without the snow module showed a strong effect of snowpack on key bioclimatic variables and species composition at higher elevations. A projected upward shift of the snow line and a decrease in snowpack might lead to drastic changes in mountain forests composition and even a shift to dry meadows due to insufficient moisture availability in shallow alpine soils.