1 JULY 2014 SWANN ET AL.

4857

Remote Vegetation Feedbacks and the Mid-Holocene Green Sahara

ABIGAIL L. S. SWANN
Department of Atmospheric Sciences, University of Washington, Seattle, Washington

INEZ Y. FUNG
Department of Earth and Planetary Science, University of California, Berkeley, Berkeley, California

YUWEI LIU AND JOHN C. H. CHIANG

Department of Geography, University of California, Berkeley, Berkeley, California

(Manuscript received 12 November 2013, in final form 26 February 2014)

ABSTRACT

In the mid-Holocene, the climate of northern Africa was characterized by wetter conditions than present, as
evidenced by higher paleolake levels and pollen assemblages of savannah vegetation suggesting a wetter,
greener Sahara. Previous modeling studies have struggled to simulate sufficient amounts of precipitation
when considering orbital forcing alone, with limited improvement from considering the effects of local
grasslands. Here it is proposed that remote forcing from expanded forest cover in Eurasia relative to today is
capable of shifting the intertropical convergence zone northward, resulting in an enhancement in pre-
cipitation over northern Africa approximately 6000 years ago greater than that resulting from orbital forcing
and local vegetation alone. It is demonstrated that the remote and local forcing of atmospheric circulation by
vegetation can lead to different dynamical patterns with consequences for precipitation across the globe.
These ecoclimate teleconnections represent the linkages between the land surface in different regions of the
globe and by inference show that proxy records of plant cover represent not only the response of vegetation to
local climate but also that vegetation’s influence on global climate patterns.

1. Introduction in the past. We suggest that the influence of vegetation on

atmospheric circulation patterns can be used to paint
Paleo-proxy records based on pollen assemblages are
a broader view of global climate conditions. Here we
commonly used to indicate local climate conditions of
show that the remote and local forcing of atmospheric
the past based on the environmental constraints of the
circulation by vegetation leads to different dynamical
pollen producing plants. Given a sufficient network of
circulation patterns with consequences for precipitation
plant cover information, a picture of climate emerges.
across the globe. Additionally, by considering ecoclimate
This view is necessarily one-sided, with the assumption
teleconnections, knowledge of vegetation cover in one
being that climate determines plant cover. The observed
region in the past can lead to additional, testable hy-
plant cover itself, while in energy balance with local
pothesis about climate in other parts of the globe.
climate, may be forcing climate both locally and remotely
through ecoclimate teleconnections (i.e., by changing
a. The ‘‘Green Sahara’’ problem
large-scale thermal and energy gradients and hence at-
mospheric circulation). As the paleo-proxy records do As an example for how vegetation cover can be used
not provide complete coverage in the spatial domain, we to understand climate patterns beyond the region they
do not have a full picture of climate globally at any time cover, we will consider here what is called the Green
Sahara problem. Paleo-proxy records indicate that
North African climate was more humid during the mid-
Corresponding author address: Abigail Swann, University of
Washington, Atmospheric Sciences, Box 351640, Seattle, WA
Holocene, approximately 6000 years ago, compared
98102-1640. to today (McIntosh and McIntosh 1981; Ritchie et al.
E-mail: aswann@u.washington.edu 1985; Petit-Maire and Guo 1996). Evidence for wetter

DOI: 10.1175/JCLI-D-13-00690.1

Ó 2014 American Meteorological Society

4858 JOURNAL OF CLIMATE
conditions across what is now the Sahara desert in-
cludes higher paleolake levels (Street and Grove 1979;
Street-Perrott and Perrott 1993) and, of particular in-
terest to this study, pollen records indicate that the
Sahara desert supported vegetation during the mid-
Holocene (Hoelzmann et al. 1998; Joussaume et al.
1999) with Sahel-like vegetation as far north as at least
238N (Jolly et al. 1998b).
There are two broad categories of mechanisms that
have been invoked to explain how the Sahara was ca-
pable of maintaining sufficient precipitation for vege-
tation in the mid-Holocene. First, the orbital forcing in
the mid-Holocene was such that Earth was slightly
closer to the sun in the Northern Hemisphere summer
and farther in the winter compared to the present day
(Berger 1978). This leads to an increase in the season-
ality of temperature in the Northern Hemisphere with
warmer summers and cooler winters as well as a farther
northward push of the Hadley circulation during boreal
summer (Kutzbach 1981; Braconnot et al. 2007a).
Second, the Charney hypothesis (Charney et al. 1975;
Charney 1975) suggests that the contrast in surface al-
bedo between a bright desert and a dark ocean surface
maintains an east–west circulation cycle with updraft
over the ocean and enhanced downwelling of dry air
over the desert. If the Sahara were instead vegetated, the
surface would be darker, thus reducing the land–ocean
albedo contrast and subsequently the zonal circulation.
If the Sahara were somehow vegetated, it might main-
tain its own precipitation by decreasing the downwelling
of dry air relative to the desert state. While Charney was
not referring directly to the mid-Holocene green Sahara,
his general hypothesis (i.e., local surface albedo controls
circulation and precipitation) is central to many argu-
ments describing how what is currently the Sahara des-
ert at one time supported green vegetation.
As we have noted, over the Sahara and present-day
Saudi Arabia, green vegetation was more widespread in
the mid-Holocene relative to today (e.g., Hoelzmann
et al. 1998). In addition to a greener Sahara, vegetation
distributions inferred from pollen records show that
deciduous forests were widespread in the midlatitudes,
especially in Europe (Prentice et al. 1998). Relative to
the present day, the distribution of grasslands in Eurasia
was reduced, with deciduous forest in its place during
the mid-Holocene (Tarasov et al. 1998). In this study we
perform a series of experiments to test the climate im-
pact of these two pattern changes in vegetation cover
implied by proxy record–based estimates from the mid-
Holocene.
A significant body of literature primarily based on
modeling techniques has attempted to incorporate and
evaluate the effects of orbital forcing, local vegetation
VOLUME 27
changes, and changes in ocean surface temperatures to
explain the difference in vegetation distribution be-
tween the mid-Holocene and today in the Sahara/Sahel
region of Africa (e.g., Joussaume et al. 1995; Kutzbach
et al. 1996; Claussen and Gayler 1997; Brovkin et al.
1998; Claussen et al. 1999; Braconnot et al. 2000; Bonfils
et al. 2001; Levis et al. 2004; Otto-Bliesner et al. 2006).
Studies have also focused on this region during the
present day, finding that local vegetation enhances
rainfall variability through feedbacks from both evapo-
transpiration and surface albedo (Zeng et al. 1999).
A northward shift of the intertropical convergence
zone (ITCZ) is observed in model experiments of the
mid-Holocene due simply to the change in orbital forc-
ing (e.g., Kutzbach 1981). Orbital forcing is insufficient
to produce precipitation comparable to estimates from
proxy records in many studies; however, simulation of
precipitation in the mid-Holocene over North Africa is
improved by the inclusion of either interactively cou-
pled vegetation or offline forcing from vegetation (e.g.,
Claussen et al. 1999). Most importantly, previous work
has shown that local decreases in surface albedo prop-
erties (e.g., Claussen and Gayler 1997; Bonfils et al. 2001;
Levis et al. 2004) are required to reduce the high surface
albedo–driven downwelling identified by Charney.
The experiments from the Paleoclimate Modelling
Intercomparison Project phase 2 (PMIP2) and phase 5
of the Coupled Model Intercomparison Project (CMIP5)
(Braconnot et al. 2007a,b; Harrison et al. 2013) show
some shift in the ITCZ with changes in orbital forcing
and, in some cases, dynamic representation of vegeta-
tion distributions predicted from climate. However, all
of the models from both archives, including models with
both static and dynamic vegetation, underestimate the
relatively increased precipitation in Afro-Asian mon-
soon precipitation in the mid-Holocene (Harrison et al.
2013).
In this study we investigate a new mechanism for el-
evated levels of precipitation over North Africa in the
mid-Holocene. We propose that an expansion of Eur-
asian forest relative to today would cause a northward
shift in the Hadley circulation and alter precipitation
patterns in the tropics and subtropics. This mechanism
has been identified in the present day by Swann et al.
(2012); however, during the mid-Holocene, this mech-
anism can act in combination with both orbital forcing
and local changes in vegetation to increase precipitation
over North Africa.
b. Remote forcing of precipitation over Africa
It has long been observed that the location of tropical
precipitation is highly correlated with sea surface tem-
peratures in the tropics (e.g., Bjerknes 1969; Manabe
1 JULY 2014 SWANN ET AL.
et al. 1974; Cornejo-Garrido and Stone 1977; Liebmann
and Hartmann 1982; Gill and Rasmussen 1983). At low
latitudes, nearby ocean surface temperature is thus a big
driver of local precipitation. There is, however, emerg-
ing evidence in the literature for a midlatitude control
on the tropical Hadley circulation (e.g., Chiang and Bitz
2005; Broccoli et al. 2006; Frierson and Hwang 2012;
Chiang and Friedman 2012; Hwang et al. 2013), specif-
ically with regards to shifting the latitudinal position of
the ITCZ.
Several examples of different types of forcing in the
midlatitudes have been shown to influence the Hadley
circulation and shift the ITCZ. Theoretical work with
an interactive ocean indicates that this tropical response
is driven by increased cross-equatorial atmospheric energy
transport requirements, requiring an altered Hadley
circulation (Kang et al. 2008, 2009). Consistent with this,
analysis of nine future climate simulations with slab
oceans found that all models shift their ITCZ toward the
hemisphere with an increase in energy sources (Frierson
and Hwang 2012). The relationship between inter-
hemispheric energy difference and the location of the
ITCZ has been shown in coupled models as well (e.g.,
Broccoli et al. 2006). This mechanism has been invoked
to explain a southward ITCZ shift during the Last
Glacial Maximum (Chiang and Bitz 2005), the circula-
tion response to anthropogenic aerosol forcing of the
Northern Hemisphere in recent decades (Rotstayn et al.
2000; Yoshimori and Broccoli 2008; Hwang et al. 2013),
and increases in midlatitude forest cover (Swann et al.
2012). It has also been postulated that this mechanism
could be responsible for an abrupt climate change ob-
served at 14.6 kya (1 kya 5 103 yr ago; Broecker and
Putnam 2013).
Numerous studies have investigated the link between
vegetation cover and local climate across the globe (i.e.,
Kleidon et al. 2000; Bala et al. 2007; Bathiany et al.
2010). Several studies have also investigated the in-
fluence that vegetation might have on ‘‘remote’’ loca-
tions through changes in atmospheric circulation (e.g.,
Notaro and Liu 2007; Chen et al. 2012; Ma et al. 2013)
and links have been found between changes in tropical
forest cover and large-scale climate patterns (Kabat
et al. 2004; Avissar and Werth 2005). As mentioned
above, there is an extensive literature on the influence
of local vegetation on the climate of the mid-Holocene
over Africa (e.g., Claussen 2009). Reale and Dirmeyer
(2000) also find a displacement of precipitation over
Africa related to land use change in the Mediterranean
region during the Roman classical period, a few thou-
sand years following the time period of this study.
Swann et al. (2012) was the first to demonstrate a link
between midlatitude forest cover and the location of the
4859
ITCZ. In their experiments, an increase in midlatitude
forest cover in the Northern Hemisphere causes more
solar energy to be absorbed at the surface because of the
relatively lower albedo of forest versus grass. The ex-
panded forest cover caused the Northern Hemisphere to
warm as insufficient soil moisture limited the ability of
the surface to shed energy through latent heat flux, in-
stead balancing the surface energy budget through an
increase in sensible heat flux and subsequent increase in
surface temperature. Later studies using a different cli-
mate model have also shown links between future ex-
pected vegetation distributions and the location of the
ITCZ and the South Asian monsoon (Falloon et al. 2012;
McCarthy et al. 2012). We propose here that the in-
fluence of midlatitude forests found in Swann et al.
(2012) holds in the mid-Holocene as well, and that the
expansion of forests in Eurasia is able to enhance a shift
in the ITCZ and cross-equatorial energy transport in
addition to any changes induced by orbital forcing.
2. Methods
To investigate the role of vegetation changes over
Eurasia and the Sahara, we use the National Center for
Atmospheric Research (NCAR) Community Atmo-
sphere and Land Models with a carbon cycle and slab
ocean (CAM3.5-CLM3.5-CASA0 ) (Gent et al. 2010;
Oleson et al. 2008; Chen et al. 2010). All simulations
were also tested with an earlier version of the atmo-
spheric model (CAM3.0-CLM3.5-CASA0 ) with qualita-
tively similar results. CAM3.5 uses an updated convective
scheme compared with CAM3.0 as described in Chen
et al. (2010).
Vegetation in the model interacts with climate
through changing stomatal conductance, leaf area, and
biomass and hence surface albedo, transpiration, and
roughness. The leaf and stem area of vegetation is in-
teractive with climate while the fraction of each grid cell
assigned to each plant functional type is fixed for a given
simulation. The interactive leaf area of plants in this
model can feed back on climate and changes in pro-
ductivity are explicitly calculated. The slab ocean model
has interactive sea surface temperatures as other studies
have shown that thermodynamic interaction between
the ocean and atmosphere is necessary for allowing
shifts in large-scale circulation and precipitation. The
slab ocean model also has thermodynamic sea ice.
The version of the model used in this study has been
evaluated against observations by Gent et al. (2010) and
is found to have improvements over previous versions,
particularly in ENSO behavior and monsoon dynamics
(Chen et al. 2010). A closely related land model to that
used for this study has been extensively tested against
4860 JOURNAL OF CLIMATE VOLUME 27

observations as part of the Carbon-Land Model In-

tercomparison Project (Randerson et al. 2009), which
created observational benchmarks against which to
compare the performance of land surface–carbon cycle
models. CAM-CASA0 performs reasonably against ob-
servations with the most obvious deficiencies being too
little carbon uptake during the growing season in
northern biomes and net primary productivity (NPP)
estimates that are overall slightly too high. The hy-
draulic aspects of the land model have additionally been
tested against FLUXNET data (St€ ockli et al. 2008).
Using this model setup we performed two control
simulations and three experiments. In the first experi-
ment, all the midlatitude area in Eurasia (158W–1658E)
between 308 and 608N currently occupied by either C3
grasses or agriculture is converted to broadleaf deciduous
forests [Eurasian vegetation (EV); see also Fig. 1b]. This
particular conversion in Eurasia (from grass and crops
to forest) is used to simulate the previously forested
regions of Eurasia currently occupied by grasslands and
agricultural regions. In the second experiment, bare
ground over the Sahara desert and Saudi Arabia is
converted to grassland [Sahara vegetation (SV); see also
Fig. 1a], similar to projections of mid-Holocene vege- FIG. 1. Changes to vegetation cover: Patterns of vegetation
change imposed in model experiments. (a) Map of new grasses
tation cover (Hoelzmann et al. 1998). In the third ex- added over the Sahara and Saudi Arabia for SV experiment in
periment these two vegetation changes are imposed percent of grid cell converted. New grasses were added in a pattern
simultaneously [Sahara and Eurasia vegetation (SEV)]. similar to Hoelzmann et al. (1998). (b) Map of new deciduous
In all experiments, the remaining vegetation is left un- forests added over Eurasia for EV experiment in percent of grid
modified, as are glaciers and lakes. These experiments cell converted. New forests were added on agricultural lands and
C3 grassland in Europe and Asia between 308 and 608N. The pat-
are performed for the orbital conditions of the mid- terns of vegetation change from (a) and (b) are combined for
Holocene (MH). The two control runs use present day a third experiment (SEV).
vegetation cover and represent the orbital conditions of
the mid-Holocene (CMH), and the present-day orbital
conditions [preindustrial (CPI)]. CO2 levels are held angle, plant type, snow cover, the moisture content and
fixed at preindustrial CO2 levels of 280 ppm. A list of soil type of bare ground, and other land cover types
control simulations and experiments can be found in (lakes and ice). As a general order of magnitude, the
Table 1. albedo of a flat leaf in visible light is 0.1 for broadleaf
The base value of optical properties for each plant deciduous trees and 0.11 for both C3 and C4 grasses.
type is specified in the model, but realized surface al- Bare soil albedo varies between 0.08 and 0.36 depend-
bedo values in any simulation are interactively calcu- ing on the soil color class and the soil moisture relative
lated as a function of dynamic leaf area, stem area, leaf to saturation.

TABLE 1. Summary of simulations. Experiments and control runs shown in this paper are summarized here with the abbreviation,
a short description, the vegetation cover imposed, orbital conditions, and CO2 concentrations used in each simulation. The vegetation
patterns imposed are shown in Figs. 1a and 1b. Present-day orbital conditions are set to the year 1950, and mid-Holocene orbital conditions
are set to 6000 yr before present.

Abbreviation Description Vegetation Orbital conditions CO2 (ppm)

CPI Preindustrial control Present day Present day 280
CMH Mid-Holocene control Present day Mid-Holocene 280
SVMH Sahara grassland Grass in the Sahara Mid-Holocene 280
EVMH European forests Forests in Europe Mid-Holocene 280
SEVMH European forests and Forests in Europe and Mid-Holocene 280
Sahara grassland grass in the Sahara
1 JULY 2014 SWANN ET AL. 4861

The model simulations are integrated for 60 yr,

and the results presented are averages of the last 50 yr.
The spinup time of 10 years is sufficient to bring cli-
mate variables (temperature, precipitation, sea ice
cover, snow cover, soil moisture, etc.) into equilibrium.
CO2 is held fixed at 280 ppm for the entire 60 yr in all
simulations. The spatial resolution of the model uses
T42 spectral truncation, which corresponds to approx-
imately 2.88 by 2.88 grid cells, and there are 10 soil
layers.
Resulting climate fields are then analyzed by com-
paring experiments with controls and experiments with
one another. Anomalies are calculated as the differ-
ence between an experiment (SV, EV, or SEV) and
present-day vegetation (CPI or CMH). We focus in
particular on two comparisons. The first is between the
combined experiment [Saharan grass, Eurasian forests,
and orbital forcing (SEVMH)] and the preindustrial
control (CPI) where a change in variable f is repre-
sented by Df 5 fSEVMH 2 fCPI. The second isolates
only the additional added effect of expanded Eurasian
forests during the mid-Holocene where we compare
SEVMH with SVMH so that Df 5 fSEVMH 2 fSVMH.
Statistical significance for Df is determined using a
Student’s t test and significance is reported as p values,
where a p value of 0.05 indicates that we reject the null
hypothesis that the anomaly is zero with 95% confi-
dence. In general we use a significance threshold of
95% but report actual p values and indicate higher
FIG. 2. Changes to temperature: Change in annually averaged
significance where possible. We estimate a lagged au-
temperature (K) for (a) the mid-Holocene control compared with
tocorrelation of 2 years or less for all variables and have the preindustrial control (CMH–CPI) showing the effect of orbital
used a conservative estimate of 25 degrees of freedom forcing; (b) the SEVMH simulation compared with preindustrial
for each 50-yr period. control (SEVMH–CPI) showing the combined effect of orbital
The northward energy flux by the atmosphere in watts forcing, grassland in the Sahara, and expanded forests in Europe;
and (c) the SEVMH experiment compared with the SVMH ex-
per meter squared at each latitude u is calculated as
periment (SEVMH–SVMH) showing the isolated additive effect of
ðu ð 2p expanded European forests. A zonal average is shown to the right
Fu 5 RTOA a2 cosu dl du, (1) of each map. Values that do not pass a significance test at 95%
2p/2 0 confidence have been omitted.

where l is longitude, a is the radius of Earth in meters,

and RTOA is the net radiation at the top of the atmosphere
in watts per meter squared [Eq. (2.21) from Hartmann
(1994)]. Cloud contribution to net radiation at the top of
the atmosphere is measured as the difference between
clear-sky and all-sky net radiation at the top of the at-
mosphere. Surface albedo contribution to radiation at the
top of the atmosphere is measured as the change between
two experiments in shortwave flux at the top of the at-
mosphere under clear-sky conditions.
A global radiative kernel was used to characterize the
differences in energy fluxes at the top of the atmosphere
between model experiments following the form (Soden
et al. 2008)
›RTOA
dRTOA ’ å dXi , (2)
i ›Xi
where dRTOA is the change in top-of-atmosphere energy
flux, and terms inside the summation designate the
change of RTOA due to individual component Xi, where
Xi represents surface temperature, atmospheric tem-
perature, water vapor, surface albedo, or clouds. The
radiative kernel for each component represents the
change in top-of-atmosphere energy flux due to a unit
change in that component and is represented by the
differential ›RTOA/›Xi. Kernels for water vapor and
clouds are further split into shortwave and longwave
4862 JOURNAL OF CLIMATE VOLUME 27

FIG. 3. Changes to temperature for SEVMH–CMH: Change in temperature (K) for each of four seasons for the SEVMH simulation
compared with the mid-Holocene control (SEVMH–CMH) showing the combined effect of grassland in the Sahara and expanded forests
in Europe during the mid-Holocene. The four seasons are (a) DJF, (b) MAM, (c) JJA, and (d) SON. Values that do not pass a significance
test at 95% confidence have been omitted.

contributions. We use radiative kernels calculated in
CAM3 (Shell et al. 2008; Soden et al. 2008) for all
components except for clouds. Top-of-atmosphere en-
ergy flux change due to clouds is calculated as the dif-
ference between the clear-sky and all-sky fluxes at the
top of the atmosphere, with changes in cloud radiative
effects caused solely by noncloud components removed
(Shell et al. 2008). The kernels used here were devel-
oped for CAM3; however, we find it reasonable to use
them for CAM3.5 as the radiative transfer algorithms in
the two versions of the model are the same (Chen et al.
2010) and it has been shown that even across models the
difference of kernels is small (Soden et al. 2008). Kernel
calculations compare each experiment with the control
simulation of the same orbital forcing, so for these cal-
culations we compare SEVMH with CMH rather than
CPI as in the rest of the paper.
a. Impact of orbital forcing, Saharan grasslands,
and Eurasian forests
An expansion of forests in Eurasia and grasslands in
the Sahara during the mid-Holocene relative to today
has several remote impacts beyond the region with di-
rectly modified vegetation. These include impacts to
temperature and precipitation patterns, changes in plant
productivity, and changes in atmospheric dynamics and
energy transport. We first present results showing changes
in climate patterns, and second discuss changes in energy
transport and dynamics, which, in some cases, are the
proximate cause for changes in climate patterns.
1) TEMPERATURE AND PRECIPITATION
Compared with preindustrial climate, temperatures
during the mid-Holocene were generally colder in
lower latitudes and higher near the poles (Fig. 2a). This
is expected from the orbital forcing differences be-
tween approximately 6000 years ago and the present
day that led to an increase in the seasonality of tem-
perature in the Northern Hemisphere with warmer
summers and cooler winters (Kutzbach 1981; Braconnot
et al. 2007a). The combined experiment (SEVMH)
shows some aspects of the same low-latitude cooling
pattern driven mostly by orbital forcing (Figs. 2a,b),
but Eurasian forests have the additional effect of in-
creasing temperatures over the Northern Hemisphere
continents (Fig. 2c). Forests were added only to Eura-
sia in the SEVMH experiment; however, temperatures
over North America are also increased (Fig. 2c). The
spatial pattern of annual mean temperature change
driven by vegetation cover change is consistent across
seasons for SEVMH relative to the mid-Holocene con-
trol, with pronounced cooling over new grasslands in the
1 JULY 2014 SWANN ET AL.
FIG. 4. As in Fig. 2, but for changes to precipitation (mm day21).
Sahara and warming over Northern Hemisphere conti-
nental regions (Fig. 3).
Precipitation patterns change in the mid-Holocene
are due to orbital forcing but are further enhanced by
changes in vegetation distributions (Fig. 4). Large
changes in precipitation are prominent in tropical re-
gions associated with a northward shift of the ITCZ. The
increased seasonality in temperature in the Northern
Hemisphere during the mid-Holocene led to summers
that were warmer than the present day. For the com-
bined experiment (SEVMH), these warm summers lead
to a shift in the precipitation weighted location of the
ITCZ (Lintner et al. 2004) of 0.68 northward in August
(SEVMH–CPI, p value 5 0.04). The shift in the ITCZ
can also be seen visually as drying bands to the south of
wetting bands (Fig. 4). The shift in ITCZ location is
consistent with changes in atmospheric dynamics and
energy transport as discussed in section 3 below.
The spatial pattern of precipitation change driven by
vegetation cover change is relatively consistent across
seasons for SEVMH relative to the mid-Holocene con-
trol (Fig. 5) with a few notable variations. The shift in the
4863
ITCZ over the Atlantic migrates north and south with
the sun, with the most pronounced drying over northern
South America in June–August (JJA; Fig. 5c), and
drying in the south of Brazil in March–May (MAM) and
September–November (SON; Figs. 5b,d). This is con-
sistent with times when the ITCZ is shifted northward
due to the addition of vegetation causing drying. Over
the Sahara, wetter conditions are seen in three of four
seasons, with local winter showing no significant change
[December–February (DJF); Fig. 5a].
The additional contribution of midlatitude Eurasian
forests to precipitation patterns is subtle, but most no-
tably shows a modest, yet statistically significant, in-
crease in precipitation over North Africa (Fig. 4c) and
coincident increase in plant productivity (see Fig. 8d).
The precipitation increases over North Africa by an
average of 26.3% (8 mm yr21) in the domain 158–308N
and 08–308E (box shown in Fig. 6c) due to the addition
of Eurasian forests compared to the Saharan grass
only experiment (SEVMH–SVMH; Fig. 7). This sug-
gests that additional Eurasian forest cover relative to
today enhances precipitation over the green Sahara,
acting in an additive manner beyond other forcing.
Precipitation decreases in the storm track off of Asia in
the mid-Holocene are also diminished by forests in
Eurasia, resulting in little change near the coast com-
pared to the preindustrial (Fig. 4c). Precipitation changes
little over the local areas where forests were added in
Eurasia.
In the mid-Holocene, proxy records for plant avail-
able moisture and estimates of precipitation minus
evaporation (P 2 E) from lake level data indicate that
it was generally wetter over much of Africa (e.g.,
Street-Perrott and Perrott 1993; Hoelzmann et al. 1998;
Joussaume et al. 1999). This magnitude of precipitation
increase in our model simulations (SEVMH or SVMH)
when averaged across the basin is lower than predictions
of the excess precipitation needed to maintain steppe
vegetation observed to inhabit the region in the mid-
Holocene (Jolly et al. 1998a,b; Joussaume et al. 1999),
although local increases are sufficiently large (Fig. 6b).
As noted above, the additional influence of forests in
Eurasia is to further increase precipitation over Africa
beyond the change driven by orbital forcing and local
grass in the Sahara (Fig. 6c).
2) PLANT PRODUCTIVITY AND TELECONNECTIONS
The net carbon uptake by plants, or NPP, is lower in
the mid-Holocene relative to present day outside of
areas where plant cover was purposefully added (Fig. 8).
The reduced NPP is driven by a combination of de-
creases in temperature in the spring and fall seasons,
decreases in growing season length, and decreases in
4864 JOURNAL OF CLIMATE
FIG. 5. As in Fig. 3, but for changes to precipitation (mm day21) for SEVMH–CMH. Note that the scale on the color bar is different from
that in Fig. 4.
regional precipitation patterns with associated changes
in soil moisture.
Productivity increases in the areas where vegetation
has been directly added in each experiment including
increases in NPP over Eurasia (EVMH; Figs. 8b and 1b)
and the Sahara (SVMH; Figs. 8c and 1a). No change
in productivity is observed over the Sahara in in either
CMH or EVMH as there are no plants designated
over the Sahara desert in the present-day vegetation
map. We might expect to find that the combined ex-
periment with both grass in the Sahara and forests in
Eurasia (SEVMH) to be a near-linear combination of its
component experiments, particularly in locations where
vegetation was modified, yet we find that Eurasian for-
ests have the additional effect of making the grasslands
in the Sahara even more productive—consistent with
the increase in precipitation over North Africa (Figs. 6c
and 8d). That is to say that there is an additional feed-
back from Eurasian forests that enhances precipitation
over the Sahara, and this feedback operates through
shifts in atmospheric dynamics as discussed below.
We find decreases in productivity in eastern North
America in response to adding forests in Eurasia (Fig.
8b). The teleconnection to North America is also acti-
vated by adding grass in the Sahara (Fig. 8c), but the
additional effect of all forcing together shows that
Eurasian forests further exacerbate the decrease in NPP
VOLUME 27
(Fig. 8d). Proxy records including lake levels, aeolian
deposits, and estimates of precipitation minus evapora-
tion and aridity all show drying across North America at
this time (Viau and Gajewski 2001; Forman et al. 2001;
Harrison and Prentice 2003) consistent with both the
effects of orbital forcing alone and the enhanced de-
crease in productivity associated with ecoclimate tele-
connections from forest cover in Eurasia or grass in the
Sahara.
Expanding forests in Eurasia also decreases pro-
ductivity over the cerrado regions of Brazil (Fig. 8b),
while adding grass in the Sahara causes decreases in
NPP primarily in southern Brazil (Fig. 8c). Estimates of
precipitation in the mid-Holocene over the Amazon are
sparse. Proxy observations suggest drier conditions over
the southern flank of the Amazon in South America
(Prado et al. 2013), with wetter conditions over the
Andes (Liu et al. 2004). Charcoal records show a more
mixed signal over South America (Power et al. 2008).
Ecosystem reconstructions from pollen data suggest
a shift to vegetation cover consistent with drier condi-
tions relative to today (Mayle and Power 2008) and the
results from our simulations. This decrease in NPP is
enhanced in our simulations, particularly over western
Brazil, by expanding forest cover in Eurasia (Fig. 8d).
Orbitally driven changes in productivity over Brazil are
small (Fig. 8a).
1 JULY 2014 SWANN ET AL.
FIG. 6. Changes to precipitation over Africa: Change in annually
averaged precipitation (mm day21) for the (a) the mid-Holocene
control compared with the preindustrial control (CMH–CPI)
showing the effect of orbital forcing, (b) the SEVMH simulation
compared with preindustrial control (SEVMH–CPI) showing the
combined effect of orbital forcing, grassland in the Sahara, and
expanded forests in Europe, and (c) the SEVMH experiment
compared with the SVMH experiment (SEVMH–SVMH) showing
the isolated additive effect of expanded European forests. The box
drawn in (c) corresponds to the domain 158–308N, 08–308E and
represents the averaging region used in Fig. 7. Values that do not
pass a significance test at 95% confidence have been omitted.
3) ATMOSPHERIC CIRCULATION AND DYNAMICS
An expansion of forest in Eurasia and grassland in the
Sahara result in a major northward shift in the location
of precipitation in the tropics during the mid-Holocene
(Fig. 4b). We hypothesize that an increase in energy
absorbed by a darker vegetated surface creates an en-
ergy imbalance between the two hemispheres, leading to
a northward shift in the Hadley circulation and de-
creased northward flux of energy across the equator to
restore energy balance (Fig. 9; see also Chiang and
Friedman 2012). A change in energy flux by the
4865
FIG. 7. Changes to precipitation over Africa: Change in precip-
itation (mm day21) averaged over the domain 158–308N, 08–308E is
shown for the annual mean and four seasons; DJF, MAM, JJA, and
SON. Each time average is shown for CMH–CPI showing the effect
of orbital forcing (red), EVMH–CPI showing the effect of expanded
forest cover in Eurasia (green), SVMH–CPI showing the effect of
local grassland in the Sahara (brown), SEVMH–CPI showing the
combined effect of orbital forcing, grassland in the Sahara, and ex-
panded forests in Europe (blue), and SEVMH–SVMH showing the
isolated additive effect of expanded European forests (cyan). The
averaging domain is represented by the box in Fig. 6c.
atmosphere across the equator then indicates that the
shift in circulation has occurred. The anomalous energy
flux could occur in the ocean as well, but heat flux by the
ocean is held fixed in these simulations.
Orbital forcing alone leads to an anomalous north-
ward energy flux across the equator in the annual mean.
Adding vegetation in the Northern Hemisphere, either
in Eurasia alone or the Sahara alone, leads to statisti-
cally insignificant changes in cross-equatorial energy
flux. However, increasing forest cover in Eurasia com-
bined with grass cover in the Sahara perturbs the annual
mean energy balance enough to shift the Hadley circu-
lation southward (Fig. 9). Changes in vegetation cover
increase the solar energy absorbed by the surface as the
albedo decreases both directly with the addition of
vegetation, and indirectly through the relative loss of sea
ice at high northern latitudes due to relative increases in
temperature. The surface albedo change results in an
average of 4.5 W m22 decrease in net top-of-atmosphere
energy flux north of 308N for the SEVMH experiment
(2.4% change from CPI, p value , 0.001). This energy
imbalance, with a deficit in the Southern Hemisphere,
then requires a northward shift in the Hadley circulation
and a corresponding shift in the ITCZ, which is man-
ifested in the change in precipitation patterns (Fig. 4b).
Orbital forcing alone does not lead to an annual mean
shift in cross equatorial energy flux; however, the addi-
tion of forests in Eurasia on top of other forcing
(SEVMH) does significantly increase the southward
cross equatorial energy flux. The combination of all
forcing together (SEVMH) decreases the northward
4866 JOURNAL OF CLIMATE VOLUME 27

FIG. 8. Changes to NPP: Change in net primary production (gC m22 yr21) for (a) CMH–CPI isolating the effects of
orbital forcing, (b) EVMH–CMH isolating the effects of expanded forests in Eurasia, (c) SVMH–CMH isolating the
effects of grass in the Sahara, and (d) SEVMH–SVMH isolating the additive effect of expanded Eurasian forests.
Areas where more than 10% of each grid cell was modified as in Fig. 1 are outlined in magenta, although note that this
pattern of vegetation change is fully implemented only in (d). Note that (b) has modifications to land cover only in
Eurasia and (c) only in Africa and Saudi Arabia. Values that do not pass a significance test at 95% confidence have
been omitted.

cross equatorial energy flux by more than twice as much
as EVMH, reducing the northward flux from 1.8 PW
in CPI by approximately 0.054 PW—a change of 3%
(Fig. 10, Table 2). In other words, the effect of Eurasian
afforestation combined with Saharan grasslands on
cross-equatorial energy flux is larger than the linear
combination as the combined effect includes feedbacks.
We next quantify the factors that are driving changes
in the interhemispheric energy difference. Much of the
additional accumulated energy in the Northern Hemi-
sphere in our experiments is coming from a decrease in
surface albedo. Calculations using a global radiative
kernel allow us to separate the contribution to the top-
of-atmosphere net radiative balance between the hemi-
spheres from clouds, water vapor, temperature, and
albedo. We find that adding Saharan grasslands leads
to less than 0.5 W m22 energy difference between the
hemispheres (SVMH–CMH), primarily from noncloud
feedbacks (94.5%) such as surface albedo and enhanced
longwave forcing from water vapor (Fig. 11). Expanding
Eurasian forests more than doubles the interhemispheric
energy difference to 1.1 W m22 (SEVMH–CMH) with
the additional feedback coming about one-third (33.2%)
from cloud feedbacks and two-thirds (66.8%) from non-
cloud feedbacks (Fig. 11). The larger interhemispheric
energy difference in the combined experiment is con-
sistent with SEVMH having the largest change in
differential energy absorption between the hemi-
spheres (from a decrease in surface albedo due to
forest and grassland expansion) and as well as the
largest change in both cross-equatorial energy flux and
precipitation shift (Figs. 4b and 10). The latitudinal
profile of energy flux (Fig. 11) shows that grass in the
Sahara leads to a locally increased energy flux from
noncloud processes, while Eurasian forests increase the
energy flux at local latitudes primarily through cloud
processes.
b. Summary
The research presented here illustrates the concept of
ecoclimate teleconnections using the Green Sahara as
an example. We discuss how distant vegetation changes
can influence both the local climate in a given location,
and our plant-derived proxy record of that local climate.
We have shown three main consequences of increases in
Eurasian forest area during the mid-Holocene relative
to today. First, the addition of forest cover in Eurasia
and grasses in the Sahara leads to a decrease in surface
albedo and consequent increase in energy absorbed into
the Northern Hemisphere. This additional energy leads
to an imbalance between the hemispheres that persists
throughout the year. Second, this energy imbalance
leads to a northward shift in the Hadley circulation in
order to transport more energy across the equator and
1 JULY 2014 SWANN ET AL.
FIG. 9. Forest expansion shifts the Hadley circulation: The dia-
grams describe the hypothesized mechanism by which the ecocli-
mate teleconnection acts in this study. (b) The forest expansion
case has an increase of midlatitude forests relative to (a) the control
vegetation, with trees replacing areas occupied by grass in the
present day. The relatively darker expanded forest absorbs addi-
tional solar energy (Eabsorbed) leading to an imbalance in energy
between the two hemispheres. To restore energy balance the
Hadley circulation (represented by blue and red circulation boxes)
moves northward in order to increase the cross equatorial transport
of energy from north to south (Eflux). The ITCZ (represented by
vertical blue dashed lines) follows the location of convergence of
air from both hemispheres and is therefore drawn northward with
the expansion of tree cover in midlatitudes in (b). This mechanism
is presented in Swann et al. (2012) for the present day and proposed
by this study to act in the mid-Holocene in response to changes in
vegetation patterns between the two time periods.
a consequent shift in the location of the ITCZ (Fig. 9).
This northward shift in the ITCZ gives an added boost to
precipitation over North Africa on top of increases from
local grass in the Sahara. Third, the shift of the Hadley
circulation leads to teleconnections and an impact on
plant productivity in eastern North America and central
and southern Brazil.
Our underlying explanatory framework is consistent
with the emerging hypothesis of ITCZ shifts from al-
tering atmospheric cross-equatorial energy transports
(Chiang and Friedman 2012). The results presented here
show the response of a single climate model and the
4867
FIG. 10. Anomalous northward atmospheric energy transport:
Zonally averaged latitudinal profile of the annually averaged
anomalous northward energy transport by the atmosphere com-
paring each experiment against present day vegetation and orbital
conditions (CPI). The red dashed line represents the effects of
orbital forcing (CMH–CPI), the green dash-dotted line represents
the effects of adding grasslands in the Sahara and orbital forcing
(SVMH–CPI), the brown line with crosses represents the effects of
expanding forests in Eurasia and orbital forcing (EVMH–CPI),
and the thick blue line represents the combined effect of both
vegetation changes and orbital forcing (SEVMH–CPI).
dependence of these results on the model itself cannot
be ruled out; however, this gross behavior is robust
across a number of modeling studies. Quantitatively,
however, the strength of the teleconnection may differ
from model to model. The response of the carbon cycle
and plant productivity through ecoclimate teleconnections
has not yet been tested across models in experiments
similar to those presented here and represents a new
finding of this study.
We propose that an additional forcing, from the re-
mote effects of Eurasian forest cover changes, could
help shift the ITCZ northward over Africa and increase
precipitation in the Sahara during the mid-Holocene.
Given that the change in precipitation from Eurasian
forests occurs globally, this new hypothesis is testable by
additional information about climate from proxy re-
cords around the globe. The work presented here sug-
gests that the interpretation of paleo-proxy records
based on plant cover should consider ecoclimate
TABLE 2. Changes to atmospheric energy transport: Change in
annually averaged northward energy flux by the atmosphere at each
latitude is shown in each row for an experiment compared against
the preindustrial control (CPI). The last row (SEVMH–SVMH)
shows the additive effect of expanding forests in Eurasia. Columns
show the total change in energy flux (EFlux) in petawatts, the change
represented in percent of the preindustrial control flux at the
equator, and the p value of significance. Significance at the 99%
confidence level or above is indicated by boldface values.
Experiment DEFlux (PW) DEFlux (%) p value
CMH–CPI 0.076 4.2 0.0003
SVMH–CPI 0.020 1.1 0.2786
EVMH–CPI 20.003 20.1 0.8680
SEVMH–CPI 20.054 23.0 0.0080
SEVMH–SVMH 20.091 0.0008
4868 JOURNAL OF CLIMATE
FIG. 11. Changes to top-of-atmosphere net energy fluxes: The
change in top of atmosphere net radiative flux of energy is shown as
a function of latitude for the total flux (thick solid line), contribu-
tions from clouds (thin solid line), and contributions from noncloud
factors (dashed line) including albedo, longwave and shortwave
forcing from water vapor, and changes in temperature. These fluxes
are shown for the combined effect of Eurasian forests and Saharan
grass (blue lines, SEVMH–CMH) and the additive effect of
Eurasian forests in the combined experiment (magenta lines,
SEVMH–SVMH). Colored overbars show the latitudes at which
grass was added to the Sahara in SEVMH and SVMH (green) and
forests were added to Eurasia in SEVMH (brown).
teleconnections—the fact that local vegetation reflects
both local climate conditions and may imply global
circulation patterns. By considering these ecoclimate
teleconnections, knowledge of vegetation cover in one
region in the past can lead to additional, testable hy-
pothesis about climate in other parts of the globe.
Acknowledgments. We would like to acknowledge
the help of Gordon Bonan and Samuel Levis for assist-
ing us with model setup. Dargan Frierson provided
comments and Brian Rose provided helpful discussions.
We acknowledge National Science Foundation Awards
ATM-0628678 and AGS-1143329 to the University of
California, Berkeley, and National Science Foundation
Award AGS-1321745 to the University of Washington.
This work was partially conducted while A.L.S.S. was
a Giorgio Ruffolo Fellow in the Sustainability Science
Program at Harvard University, for which support from
Italy’s Ministry for Environment, Land and Sea is
gratefully acknowledged. Some of the computing re-
sources used in this work were provided by the Climate
Simulation Laboratory at NCAR’s Computational and
Information Systems Laboratory (CISL), sponsored by
the National Science Foundation and other agencies.
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