Coniferous Habitats Research Articles

Hunting behaviour of eastern coyotes in relation to vegetation cover, snow conditions, and hare distribution

:Coyotes (Canis latrans) have expanded their range to the north and east in North America during the last century. It has been suggested that vegetative cover reduces hunting efficiency of forest-dwelling coyotes, which could explain their lower performance compared to rural coyotes. Also, in their northern range, coyotes must periodically cope with deep and soft snow conditions for which they are not morphologically adapted. We snow-tracked coyotes in a forested landscape of southeastern Québec to test the hypothesis that coyotes modify their hunting strategy with respect to vegetative cover, snow condition, and hare distribution (Lepus americanus), the main prey species during winter in our study area. When snow sinking depth hampered mobility, coyotes used habitats with the lowest snow sinking depth (coniferous habitats) in a greater proportion than availability (47.0 versus 29.7%), despite dense lateral cover. As a result, coyotes chased hares over short distances (17 m on average). However, when snow conditions did not hamper mobility, use of coniferous habitats decreased to 18.5% in favour of open habitats, which were used more than availability (47.7 versus 32.5%), despite low hare abundance. Use of sites with little lateral cover and low sinking depth facilitated long chases by coyotes (126 m on average). Throughout winter, coyotes used sites characterized by less lateral cover and lower snow sinking depth than random sites. Our results suggest that coyotes selectively used habitat and hunting strategy to maximize their net energy budget throughout winter. Fragmentation of forest landscapes generates abundant openings and small refuges which may benefit coyotes to the detriment of native prey populations.

Performance of differential GPS collars in temperate mountain forest

To determine the performance of Global Positioning System (GPS) for habitat studies in free-ranging animals, we tested differential 6- and 8-channel GPS collars under six representative canopies and one open-field reference site in the ‘Parc national des Cévennes’, southern France. The proportion of successful locations decreased under taller trees and worsened with snow accumulation in mixed coniferous habitats. The mean location success of seven free-ranging red deer fitted with 6-channel GPS collars in the same study area increased with a shorter interval between location attempts and during the leaf-off period. Our data suggested that the differences in location success between leaf-on and leaf-off periods might reflect shifts in habitat use rather than a leaf effect under deciduous trees. To cite this article: G. Janeau et al., C. R. Biologies 327 (2004).

LAND USE AND LAND COVER CHANGE IN THE GREATER YELLOWSTONE ECOSYSTEM: 1975–1995

Shifts in the demographic and economic character of the Greater Yellowstone Ecosystem (GYE) are driving patterns of land cover and land use change in the region. Such changes may have important consequences for ecosystem functioning. The objective of this paper is to quantify the trajectories and rates of change in land cover and use across the GYE for the period 1975–1995 using satellite imagery. Spectral and geographic variables were used as inputs to classification tree regression analysis (CART) to find “rules” which defined land use and land cover classes on the landscape. The resulting CART functions were used to map land cover and land use across seven Landsat TM scenes for 1995. We then used a thresholding technique to identify locations that differed in spectral properties between the 1995 and 1985 time periods. These “changed” locations were classified using CART functions derived from spectral and geographic data from 1985. This was similarly done for the year 1975 based on Landsat MSS data. Differences between the 1975, 1985, and 1995 maps were considered change in land cover and use. We calibrated and tested the accuracy of our models using data acquired through manual interpretation of aerial photos. Elevation and vegetative indices derived from the remotely sensed satellite imagery explained the most variance in the land use and land cover classes (i.e., defined the “rules” most often). Overall accuracies from our study were good, ranging from 94% at the coarsest level of detail to 74% at the finest. The largest changes over the study period were the increases in burned, urban, and mixed conifer-herbaceous classes and decreases in woody deciduous, mixed woody deciduous–herbaceous, and conifer habitats. These changes have important implications for ecological function and biodiversity. The expansion of mixed conifer classes may increase fuel loads and enhance risk to the growing number of rural homes. The reduction of woody deciduous cover types is likely reducing population sizes for the numerous plant and animal species that specialize on this habitat type. Some of these species are also negatively influenced by the increase of rural homes in and near woody deciduous habitats. Corresponding Editor: I. C. Burke

Forest Landscape Patterns around Marbled Murrelet Nest Sites in the Oregon Coast Range

We analyzed forest composition and landscape preferences of marbled murrelets (Brachyramphus marmoratus). Using dawn surrreys and tree climbing, we found 41 murrelet nest sites on state and federal lands in the Oregon Coast Range Province in western Oregon between 1990 and 1998. Landscapes in 0.5- and 1.0-km-radius plots at these nest sites were compared to patterns in similar plots around a set of points randomly selected from stands of mature or old- growth trees on public lands in the same province. We found less open-sapling and hardwood forests in the plots at nest sites than at random sites for both plot sizes. Conversely, the pro- portion of pole-young conifer habitat was greater at nest sites than at random sites for both plot sizes. Landscape pattern analysis showed that the amount of edge-perimeter density, nest-patch perimeter, and high-contrast edge of nest patches was lower in the 1.0-km-radius plots at nest sites than at random sites. Our best multivariate logistic regression model indicated that greater amounts of pole-young and mature-old-growth forests, less edge (perimeter density and high- contrast edge at nest patches), and more cohesive nest-patch shape best distinguished murrelet nest sites from random sites. We hypothesize that murrelet nest-site selection at the landscape scale may be the result of an antipredator strategy to reduce predation risks on eggs and juvenile murrelets. Young (simple-structure) conifer stands adjacent to nesting areas may decrease pre- dation rates at murrelet nests. Land managers should consider limiting clearcut harvest units both adjacent to murrelet nest patches and within at least 1 km of murrelet nests, as clearcuts increase high-contrast edge in addition to increasing fragmentation levels.

Calcium shortage as a constraint on reproduction in great tits Parus major: a field experiment

Several bird species breeding in acidified habitats face serious problems caused by lowered availability of calcium‐rich foods. Moreover, recent studies have revealed that calcium shortage causes phenotypic variation in certain reproductive parameters of passerines even in non‐acidified habitats. However, to date, no clear evidence has been found of the limiting effect of calcium deficiency on the reproductive output of birds breeding in such habitats. In this paper, we present experimental evidence from a long‐term study that calcium shortage may indeed limit reproductive output of passerines that breed in non‐acidified habitats. We carried out calcium‐supplementation experiments with great tits Parus major breeding in coniferous and deciduous woodlands in Estonia. Calcium‐supplemented birds laid more eggs, tended larger broods and raised more fledglings with longer tarsi compared with control birds. Calcium shortage constrained reproduction of tits not only in spring, but also in mid‐summer, during second breeding attempts. Moreover, no significant differences were detected between deciduous and coniferous habitats with respect to the effect of calcium supplements on reproductive success.

Autoregressive moving average models of conifer crown profiles

A time-series autoregressive moving average (ARMA) approach was used to develop stochastic models of tree crown profiles for five conifer species of the Sierran mixed conifer habitat type. Models consisted of three components: (1) a polynomial trend: (2) and ARMA model; and (3) rand om error. A Bayesian information criterion was used to evaluate alternative models. It was found that 70% of the crown profiles could be modeled using first-order ARMA [AR(1) or MA(1)] models, and that an additional 25% could be modeled using a white noise model [(AR(0)]. When the coefficients of the ARMA models were statistically significant, the models proved to be both visually and statistically an improvement over the polynomial trend (a Euclideam model). A binary classification system was used to determine if model type was related to tree or stand characteristics. Using this classification we found that it was possible to relat the appropriate model type to forest tree size and forest stand density with acceptable accuracy.

DOES DENSITY REFLECT HABITAT QUALITY FOR NORTH AMERICAN RED SQUIRRELS DURING A SPRUCE-CONE FAILURE?

Predicting animal populations over time often is done using models of density–habitat relationships that assume animal density is a reflection of habitat quality. We explored whether this assumption was true for the North American red squirrel (Tamiasciurus hudsonicus), a species present at different densities in 3 conifer habitats: white spruce (Picea glauca; high resource quality, unstable availability), mixed conifer, and lodgepole pine (Pinus contorta; lower resource quality, stable availability). We documented density, body condition, survival, reproduction, and turnover in these 3 habitats during a failure of spruce-cone crop, when the potential for the relative importance of habitats to change was greatest. During this time squirrel densities in white spruce decreased by 66% to match with those found in pine and mixed-conifer forests. Red squirrels in spruce forests experienced lower survival and fewer females successfully weaned young, and juvenile production was lower. Adult and juvenile immigration was more important than local juvenile production in replacing squirrel mortality in spruce and mixed-conifer forests. Our results indicate that density does not always reflect habitat quality for red squirrels, and we question the historical high-quality rating of white spruce habitat for this species. Further, our findings suggest that movement of ostensibly highly territorial adults in late winter and early spring is an important mechanism in determining annual squirrel densities regardless of habitat type.

Densities of breeding birds and changes in vegetation in an Alaskan boreal forest following a massive disturbance by spruce beetles

We examined bird and plant communities among forest stands with different levels of spruce mortality following a large outbreak of spruce beetles (Dendroctonus rufipennis (Kirby)) in the Copper River Basin, Alaska. Spruce beetles avoided stands with black spruce (Picea mariana) and selectively killed larger diameter white spruce (Picea glauca), thereby altering forest structure and increasing the dominance of black spruce in the region. Alders (Alnus sp.) and crowberry (Empetrum nigrum) were more abundant in areas with heavy spruce mortality, possibly a response to the death of overstory spruce. Grasses and herbaceous plants did not proliferate as has been recorded following outbreaks in more coastal Alaskan forests. Two species closely tied to coniferous habitats, the tree-nesting Ruby-crowned Kinglet (Regulus calendula) and the red squirrel (Tamiasciurus hudsonicus), a major nest predator, were less abundant in forest stands with high spruce mortality than in low-mortality stands. Understory-nesting birds as a group were more abundant in forest stands with high levels of spruce mortality, although the response of individual bird species to tree mortality was variable. Birds breeding in stands with high spruce mortality likely benefited reproductively from lower squirrel densities and a greater abundance of shrubs to conceal nests from predators.

Densities of breeding birds and changes in vegetation in an Alaskan boreal forest following a massive disturbance by spruce beetles

We examined bird and plant communities among forest stands with different levels of spruce mortality following a large outbreak of spruce beetles (Dendroctonus rufipennis (Kirby)) in the Copper River Basin, Alaska. Spruce beetles avoided stands with black spruce (Picea mariana) and selectively killed larger diameter white spruce (Picea glauca), thereby altering forest structure and increasing the dominance of black spruce in the region. Alders (Alnus sp.) and crowberry (Empetrum nigrum) were more abundant in areas with heavy spruce mortality, possibly a response to the death of overstory spruce. Grasses and herbaceous plants did not proliferate as has been recorded following outbreaks in more coastal Alaskan forests. Two species closely tied to coniferous habitats, the tree-nesting Ruby-crowned Kinglet (Regulus calendula) and the red squirrel (Tamiasciurus hudsonicus), a major nest predator, were less abundant in forest stands with high spruce mortality than in low-mortality stands. Understory-nesting bir...

Body weights in grey and red squirrels: do seasonal weight increases occur in conifer woodland?

AbstractSeasonal body weight changes were investigated in red and grey squirrels in spruce‐dominated conifer plantations in the north of England. Annual seed food availability, particularly in the spruce plantations, varies markedly and is characterized by years with large cone crops (‘mast crops’) followed by several years of intermediate or low crops. The last mast crop in Kielder Forest District occurred in 1991 and we studied the two squirrel species when food availability was intermediate and low between 1992 and 1999. Neither red nor grey squirrels in the conifer plantations significantly increased their body weight for the autumn or winter. The responses of red and grey squirrels to the different foraging conditions in deciduous and conifer woodlands and the accumulation of seasonal fat stores may have to be seen in the context of habitat type and the pattern and predictability of seed availability. Our findings suggest that fat accumulation is likely to be less pronounced in conifer habitats where autumn and winter food supplies are more predictable, and manoeuvrability to feed on cones in the canopy is important.

Long‐term dynamics of pine and hardwood litter in contrasting environments: toward a global model of decomposition

SummaryWe analysed data on mass loss after five years of decomposition in the field from both fine root and leaf litters from two highly contrasting trees, Drypetes glauca, a tropical hardwood tree from Puerto Rico, and pine species from North America as part of the Long‐Term Intersite Decomposition Experiment (LIDET). LIDET is a reciprocal litterbag study involving the transplanting of litter from 27 species across 28 sites in North and Central America reflecting a wide variety of natural and managed ecosystems and climates, from Arctic tundra to tropical rainforest. After 5 years, estimated k‐values ranged from 0.032 to 3.734, lengths of Phase I (to 20% mass remaining) from 0.49 to 47.92 years, and fractional mass remaining from 0 to 0.81. Pine litter decomposed more slowly than Drypetes litter, supporting the notion of strong control of substrate quality over decomposition rates. Climate exerted strong and consistent effects on decomposition. Neither mean annual temperature or precipitation alone explained the global pattern of decomposition; variables including both moisture availability and temperature (i.e. actual evapotranspiration and DEFAC from the CENTURY model) were generally more robust than single variables. Across the LIDET range, decomposition of fine roots exhibited a Q10 of 2 and was more predictable than that of leaves, which had a higher Q10 and greater variability. Roots generally decomposed more slowly than leaves, regardless of genus, but the ratio of above‐ to belowground decomposition rates differed sharply across ecosystem types. Finally, Drypetes litter decomposed much more rapidly than pine litter in ‘broadleaved habitats’ than in ‘conifer habitats’, evidence for a ‘home‐field advantage’ for this litter. These results collectively suggest that relatively simple models can predict decomposition based on litter quality and regional climate, but that ecosystem‐specific problems may add complications.

The foraging behaviour and diet of red squirrels Sciurus vulgaris receiving supplemental feeding

The effect of ad libitum supplemental feeding upon the diet and foraging behaviour of red squirrels Sciurus vulgaris was studied in a coniferous habitat during May 1994 ‐ July 1996. Although supplemental feeding in the form of peanuts was widely available, the animals continued to exploit natural foods such as pine seed, buds and flowers. The degree to which pine seed was exploited was a function of pine cone availability. In contrast, peanut consumption was not correlated with peanut availability, but was determined by the abundance of pine cones. During late winter and spring when seed was becoming scarce, peanuts contributed 30–56% of the dietary items. In addition, increasing amounts of alternative ‘low energy’ foods such as buds and flowers were also exploited. These foods are calcium rich and their ingestion would compensate for the high phosphorus content of peanuts. The nutritional composition of peanuts appears to have restricted the degree to which they could be included in the diet. Nevertheless peanuts were an important dietary component, and seasonally contributed 25–181% of the animals' basic daily energy requirements. Supplemental feeding was associated with relatively large amounts of activity occurring on the ground (annual mean estimates were 47 and 50% of total foraging time). The animals continued to exhibit ‘scatter‐hoarding’ behaviour, with peanuts accounting for 90% of the identified cached items. This stored resource could satisfy 344 (1994/95) and 198 (1995/96) days of the animals' basic energy requirements.

Fragments are not islands: patch vs landscape perspectives on songbird presence and abundance in a harvested boreal forest

The boreal mixed‐wood forest of northern Alberta. Canada is characterized by a mosaic of deciduous and coniferous forest patches. Recently, the deciduous portion of the forest was allocated for industrial logging. Widespread habitat loss and fragmentation may negatively affect birds and other wildlife. Most research on the effects of habitat loss on bird abundance has focussed on the forest as a patch or island in a matrix of non‐habitat, but some species of songbird may use both the forest patch and the matrix. We hypothesized that some species of songbird might be able to compensate for a loss of deciduous forest by moving into other habitat types (termed “habitat compensation”). We report on a replicated field investigation in which we assessed the response of songbirds to commercial timber harvest by first examining their abundance within deciduous forest only, and then adding the clearcuts and coniferous forest in the surrounding areas to the analysis for a broader, landscape view of the system. Bird communities in deciduous and coniferous habitats had significant overlap in species composition: there was less overlap between forest and clearcuts. The shift from patch‐centred to landscape sampling altered our interpretation of over half of the most common species' responses to logging in at least one year, suggesting that habitat compensation may have been occurring. However, significant variation in responses of species was observed between the two study areas. Our past reliance on island biogeographic and other single habitat approaches may be inappropriate for this system, and we stress that a broad, landscape view is required to properly assess and interpret species' responses to habitat loss and fragmentation.

En jämförelse mellan fyra biotopers betydelse för fågelfaunan i mellersta Blekinge

The birds were counted in four different habitats (coniferous forest, deciduous forest and two types of mixed farmland with much pasture and meadows, one near the coast and the other one upland) during winter (November 1997–March 1998) and spring (late April through June 1998) in the province of Blekinge, southern Sweden. One point count route with 12 points was established in each habitat. Five complete counts were made in each season with a five minutes count at each point. All birds seen or heard were counted, but in this study I included only birds that actually used (keeping a territory, feeding, resting) in the specific habitat of each route. The coniferous habitat was poorer than the deciduous one, especially in winter. Six of the 12 points in the coniferous habitat had no deciduous trees and it was mainly these monocultures that were poor. The other points with some deciduous trees compared well with the deciduous habitat. The two pasture habitats were much richer than the forest habitats in winter. In spring there was only a small difference between the forest habitats and the upland pasture habitat, whereas the coastal pasture habitat was much richer, mainly because of a few coastal species. Vicinity to lakes, watercourses and marshes was a key factor contributing to high species diversity in both farmland habitats.

Diet of Fishers (Martes pennanti) at the Southernmost Extent of Their Range

Fishers ( Martes pennanti ) in the mountains of California's Sierra Nevada occur at the southwestern margin of their distribution and inhabit different forest types with different potential prey than elsewhere in their range. Two typical fisher prey, the snowshoe hare ( Lepus americanus ) and the porcupine ( Erethizon dorsatum ), are absent from our Sierra Nevada study area. We characterized the diet of fishers in the southern Sierra Nevada by analyzing the content of 201 feces (44 males: 157 females) collected either from trapped animals or from the rest sites of radio-collared animals. Mammals were the most frequent food item; however, unlike previous reports, reptiles (20.4% of feces) and insects (55.7%) were major components of the diet. We also sampled 24 feces for the presence of spores of hypogeous fungi (false truffles) and found that 91.7% had spores representing at least six fungal species. Diversity of the diet was indicated by the fact that remains of no single family of animal or plant group were found in >22% of feces. The fisher is reputed to be a habitat specialist in the late-seral mixed conifer-deciduous forests of the western United States. Perhaps it is for this reason that our data depict the species as a dietary generalist, for whom it may be necessary to forage on many of the animal, plant, and fungal species that occur in and near mature coniferous habitat.

The use of nest boxes by the Red Squirrel Sciurus vulgaris in a coniferous habitat

AbstractThis paper describes the results of a 4‐year study examining the use of nest boxes by Red Squirrels. Two different designs of nest box were erected in a conifer woodland which supported a high density of animals; spring population estimates were between 3.5 and 4.0 individuals per hectare. Boxes were used by adults, subadults, lactating females and their young. Breeding females more frequently used the larger of the two box designs, although this preference was not statistically significant. There was no difference in the mean sizes of litters found within the two box types. The mean size of litters present in boxes was 2.7 (n = 31, SE = 0.23) which was similar to the mean litter size of 2.5 (n = 8, SE = 0.50) found in dreys. Thus, nest boxes proved to be a valuable research tool for monitoring parturition dates, litter sizes, nestling sex ratios, and preweaning mortality.

An integrated geographic information system approach for modeling the suitability of conifer habitat in an alpine environment

Alpine periglacial environments within the forest-alpine tundra ecotone (FATE) may be among the first to reflect changes in habitat characteristics as a consequence of climatic change. Previous FATE studies used Integrated Geographic Information System (IGIS) techniques to collect and model biophysical data but lacked the necessary detail to model the micro-scale patterns and compositions of habitat within alpine periglacial environments. This paper describes several promising data collection, integration, and cartographic modeling techniques used in an IGIS approach to model alpine periglacial environments in Glacier National Park (GNP), Montana, USA. High-resolution (I X I m) multi-spectral remote sensing data and differentially corrected Global Positioning System (DGPS) data were integrated with other biophysical data using a raster-based IGIS approach. Biophysical factors, hypothesized to influence the pattern and composition of the FATE and the alpine tundra ecosystem, were derived from the high-resolution remote sensing data, in-situ GPS data, high-resolution models of digital elevation, and other thematic data using image processing techniques and cartographic modeling. Suitability models of conifer habitat were created using indices generated from the IGIS database. This IGIS approach identified suitable conifer habitat within the FATE and permitted the modeling of micro-scale periglacial features and alpine tundra communities that are absent from traditional approaches of landscape-scale (30 X 30 m) modeling.

Comparative demography of red squirrels (Sciurus vulgaris) and grey squirrels (Sciurus carolinensis) in deciduous and conifer woodland

The demography of red and grey squirrels was studied by live-trapping and radio-tagging at 14 deciduous and conifer sites in southern Britain and at eight conifer sites for one year in northern England. Densities and productivity correlated with tree seed crops for both squirrel species in deciduous and conifer habitats. Productivity was reduced by high density of full-grown squirrels relative to seed abundance. In oak-hazel woods, demography of grey squirrels correlated with abundance of acorns but not of hazel-nuts, whereas density and productivity of red squirrels correlated with hazel-nut abundance. Correlations of female density and productivity with pine-cone crops did not differ between red and grey squirrels. Predators ate many radio-tagged grey squirrels in conifers, and annual survival was only 50% compared with 80-82% for both species in other habitats. Grey squirrel populations in southern conifer sites were sustained by immigration, and at northern sites female density correlated with oak abundance within 500m. Failure to exploit acorn crops puts red squirrels at a competitive disadvantage in deciduous woodland. Red squirrels had higher survival than grey squirrels in conifers, which may give them an advantage in that habitat, but could also have been explained by a lack of predators on their island study site.

Comparative demography of red squirrels (Sciurus vulgaris) and grey squirrels (Sciurus carolinensis) in deciduous and conifer woodland

AbstractThe demography of red and grey squirrels was studied by live‐trapping and radio‐tagging at 14 deciduous and conifer sites in southern Britain and at eight conifer sites for one year in northern England. Densities and productivity correlated with tree seed crops for both squirrel species in deciduous and conifer habitats. Productivity was reduced by high density of full‐grown squirrels relative to seed abundance. In oak‐hazel woods, demography of grey squirrels correlated with abundance of acorns but not of hazel‐nuts, whereas density and productivity of red squirrels correlated with hazel‐nut abundance. Correlations of female density and productivity with pine‐cone crops did not differ between red and grey squirrels. Predators ate many radio‐tagged grey squirrels in conifers, and annual survival was only 50% compared with 80‐82% for both species in other habitats. Grey squirrel populations in southern conifer sites were sustained by immigration, and at northern sites female density correlated with oak abundance within 500m. Failure to exploit acorn crops puts red squirrels at a competitive disadvantage in deciduous woodland. Red squirrels had higher survival than grey squirrels in conifers, which may give them an advantage in that habitat, but could also have been explained by a lack of predators on their island study site.

Status of the Marbled Murrelet in the Inner North Coast Ranges of California

We sought to determine the presence or absence of marbled murrelets (Brachy- ramphus marmoratus) within the northern Inner North Coast Ranges of northwestern California. We conducted murrelet surveys and collected environmental data during 1995 and 1996 on na- tional forest lands that were south of the Klamath Mountains Section and within B. marmoratus management Zone 2 as designated by the Forest Ecosystem Management Assessment Team. Using a stratified random sampling design, we surveyed for murrelets within the 2 coniferous forest habitat types most likely to be used by murrelets: late mature and old-growth Douglas- fir (Pseudotsuga menziesii) and late mature and old-growth tanoak (Lithocarpus densiflora). We used the generalized binomial model to determine sample sizes and to estimate the power of survey results for a range of assumed levels of murrelet occurrence (p) and probabilities of de- tection (p'). In 1995 and 1996, we conducted 1424 surveys at 178 Douglas-fir sampling units, surveying 30.8% of the total amount of this stratum within the study area. In 1995, we con- ducted 760 surveys at 95 tanoak sampling units, surveying 58.6% of this stratum. We did not detect B. marmoratus in either habitat type. When we assumed a p of 0.03, and a p' of 0.10, the power of our surveys to detect murrelets was 0.95 and 0.81 for the Douglas-fir and tanoak hab- itat types, respectively. Given the high power associated with our results and the large pro- portion of the total potential habitat surveyed, we concluded that our study area was not within the current range of B. marmoratus. Most habitat characteristics, including corvid abundance, were similar to murrelet nesting habitat in other areas. The relatively high elevation and far- inland location, and the related characteristics of climate and habitat structure, may explain the apparent absence of B. marmoratus from our study area. Accordingly, we recommend that the Forest Ecosystem Management Assessment Team zone lines be adjusted so that the Inner North Coast Ranges are excluded from Zone 2.