Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214

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Palaeogeography, Palaeoclimatology, Palaeoecology

j o u r n a l h o m e p a g e : w w w. e l s ev i e r. c o m / l o c a t e / p a l a e o

A 370,000-year record of vegetation and re history around

Lake Titicaca (Bolivia/Peru)
Jennifer A. Hanselman a,, Mark B. Bush a, William D. Gosling a,1, Aaron Collins a, Christopher Knox a,
Paul A. Baker b, Sheri C. Fritz c
a
Department of Biological Sciences, Florida Institute of Technology, 150 W. University Blvd., Melbourne, FL 32901, USA
b
Division of Earth and Ocean Sciences, Duke University, 301 Old Chemistry, Durham, NC 27708, USA
c
Department of Earth and Atmospheric Sciences, University of Nebraska-Lincoln, 214 Bessey Hall, Lincoln, NE 68583, USA

a r t i c l e i n f o a b s t r a c t

Article history: Fossil pollen and charcoal analyses of sediments from Lake Titicaca, Peru/Bolivia, provide a record of
Received 18 May 2009 palaeoclimatic variation spanning four full glacial cycles. Pollen, aquatic microfossils, and charcoal, as well as
Received in revised form 3 March 2011 previously published data including diatom assemblages, carbonate content, and stable carbon isotopic ratios
Accepted 8 March 2011
of organic carbon, indicate that interglacials were warm and dry whereas the peaks of glacials were cold and
Available online 11 March 2011
wet. Each of the interglacials documented in the record are somewhat different, with those of MIS 5e and MIS
Keywords:
9 inducing lower lake levels and a drier vegetation signature than those of MIS 7 and 1. The presence of
Andes charcoal particles in sediments deposited during previous interglacials provides evidence of the long-term
Climate change role of re in shaping Andean ecosystems.
Fire 2011 Elsevier B.V. All rights reserved.
Fossil pollen
Fossil charcoal
Interglacial

1. Introduction
In terms of dislocation of human society, utilization of natural
resources, and loss of biodiversity, the tropics potentially face extreme
consequences of climate change. Because of their steep environmental
gradients, and a projected 21st century warming of ca. 27 C (Urrutia
and Vuille, 2009), the tropical Andes may experience profound
rearrangement of biological communities (Malcolm et al., 2006). That
biota in the Andes responded to past climate change associated with
glacialinterglacial cycles and millennial-scale droughts is well
established (Gonzalez et al., 1966; Van der Hammen, 1974;
Hooghiemstra, 1984; Gosling et al., 2008). However, no continuous
pollen record exists from the southern hemisphere tropical Andes that
spans the Holocene and multiple glacial cycles.
For many years it was assumed that glacial stages were dry and
interglacial stages were wet in the Neotropics (Prance, 1982). The
paleoecological records from the High Plains of Bogot (4.5N) provided
the empirical data necessary to test this hypothesis in the northern
tropics (e.g. Van der Hammen, 1974; Hooghiemstra, 1984; Hooghiemstra
Corresponding author at: Present address: Department of Biology, Westeld State
University, 577 Western Ave., Westeld, MA 01086, USA.
E-mail address: jhanselman@wsc.ma.edu (J.A. Hanselman).
1
Present address: Department of Earth and Environmental Sciences, CEPSAR, The
Open University, Walton Hall, Milton Keynes, MK76AA, UK.
and Van der Hammen, 2004). The Fuquene and Funza records, which
span almost 3 million years (Myr), demonstrated that migration of forest
and grassland elements in response to climate change in northern South
America was paced by Milankovitch orbital forcing, and that glacial-age
climates were as much as 8 C cooler than modern (Hooghiemstra et al.,
1993; Van der Hammen and Hooghiemstra, 2000). The Colombian lakes
were not particularly sensitive to changes in precipitation (Torres et al.,
2005).
Here we provide a second long Andean paleoecological record, one
that has continuously accumulated sediment across four glacial
interglacial cycles (Fritz et al., 2007). Lake Titicaca straddles the
border between Peru and Bolivia and lies in a high, dry, setting in the
Altiplano. The late glacial and Holocene environmental history of Lake
Titicaca reects past changes in environmental conditions from peri-
glacial to Andean woodland in a location that is sensitive to changes in
precipitation (Paduano et al., 2003).
1.1. Climate change on the Altiplano
The existence of large paleolakes on the Altiplano has been known
for a long time (Minchin, 1882; Bowman, 1914). Meltwater discharge
from local montane glaciers was offered as an explanation for these
lacustrine highstands (Moon, 1939; Servant and Fontes, 1978). It has
subsequently been shown that implausible amounts of ice melt would
be needed to create the highstands (Hastenrath and Kutzbach, 1985;
Blodgett et al., 1997). Moreover, the highstands coincide with wet

0031-0182/$ see front matter 2011 Elsevier B.V. All rights reserved.
doi:10.1016/j.palaeo.2011.03.002
202 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214
Table 1
A description of ve vegetational communities surrounding Lake Titicaca (Bolivia/Peru). Descriptions were adapted from Troll (1968) and Graf (1981) and further described in
Paduano et al. (2003).
Vegetation type Vegetation description
Puna brava Scanty herbs and grasses
Dry puna Tufted grasses, shrubs, and herbs
Moist puna Herbaceous, Poaceae dominates
Sub-puna/shrub belt Greater diversity and density of shrubs and ferns
Andean forest Trees
phases rather than with major melt events (Blodgett et al., 1997;
Baker et al., 2001a). Instead, lacustrine highstands on the Altiplano
and glacial advances in the adjacent Andes largely coincide (Baker et
al., 2001a, 2001b; Fritz et al., 2007; Zech et al., 2007), not surprisingly
since both record positive mass balance of precipitation minus
evaporation (lakes) or melting (glaciers) (e.g. Blodgett et al., 1997;
Kull et al., 2002).
A long sedimentary record from the Salar de Uyuni that spans
several glacial cycles contains millennia-long, lacustrine sedimentary
intervals separated by salt beds indicative of dry salt pan conditions
(Baker et al., 2001a; Fritz et al., 2004). Over the past two major lake
phases (Minchin and Tauca), wet conditions have been paced by
summer insolation forcing of the South American summer monsoon
at precessional time-scales (Baker et al., 2001a, 2001b). The wettest
phases, hence highest lake levels (Bills et al., 1994; Sylvestre et al.,
1999; Placzek et al., 2006) were additionally forced by North Atlantic
sea surface temperature (Baker et al., 2001a, 2009). The precessional
pattern of alternation between wet (mud) and dry (halite) is not
readily evident in the drill core from Salar de Uyuni prior to about
60 ka, (Fritz et al., 2004), likely because of geologic changes
(unrelated to climate) in the hydrologic threshold between Lake
Titicaca and Lago Poopo. Placzek et al. (2006) published a very
different chronology for paleolake highstands for the last 120 ka based
upon U/Th dates of carbonate tufas. Their chronology does not match
that based upon dating lacustrine sediment intervals within the drill
core (Baker et al., 2001b; Fritz et al., 2004). In drill cores there is no
evidence for an Ouki highstand interval at 12098 ka (a term coined
by Placzek et al., 2006). And Placzek et al. (2006) cite no evidence for a
highstand equivalent to the Minchin paleolake that deposited about
12 m of organic-rich mud between 50 and 32 Ka (Baker et al., 2001b;
Fritz et al., 2004). It is not the purpose of the present manuscript to
evaluate these different conclusions, sufce it to state here that we
believe the older U/Th dates of Placzek et al. (2006) to be erroneous,
most likely due to diagenetic loss of uranium from the older, highly
altered carbonate deposits.
Previously published data from Lake Titicaca, both seismic
reection proles and data from drill cores LT01-1A and LT01-2B in
Lago Grande and drill core LT01-3B in Lago Huiaimarca revealed a ca.
200 m lowering of lake levels during MIS 5e (c. 125 ka). The fossil
pollen assemblage from MIS 5e in core LT01-2B (Hanselman et al.,
2005), contained high concentrations of Amaranthaceae (formerly
Chenopodiaceae or Amaranthaceae) while a sedimentary hiatus or
slow sedimentation rate is suspected in the core from the shallow
water, Huiaimarca sub-basin (see Gosling et al., 2008, 2009). Fossil
diatom and geochemical data also supported lower lake levels and
increased salinity during MIS 5e (Fritz et al., 2007) and is supported in
seismic reection proles (D'Agostino et al., 2002).
1.2. Vegetation communities of the Altiplano and
palaeoecological signicance
The distribution of the ve main terrestrial vegetation communi-
ties found on the Altiplano today is strongly related to elevation and
precipitation (Table 1; Troll, 1968; Graf, 1981). The continuous tree
Characteristic taxa Elevation (m)
Caryophyllaceae, Asteraceae (Mutisiae, Tubulifoloriae), 53004500
Apiaceae (Azorella), Malvaceae, Valeriana
Poaceae, Ephedra, Cactaceae, Fabaceae, Iridaceae 45003800/3700
Poaceae, Asteraceae, Acaena, Gentianaceae, Ericaceae, Polygonaceae 45003800/3700
Asteraceae, Melastomataceae, Polylepis, Polypodiaceae 3800/37003200
Alnus, Podocarpus, Weinmannia, Myrsine b3200
line lies close to the upper limit of where cloud formation provides
moisture year round (c. 3200 meters above sea level [masl]). The
upper Andean forest is a diverse community in which Alnus and
Podocarpus are important genera. Above the continuous tree line the
landscape is dominated by grasslands, but contains patches of
woodland. These isolated woodlands are often dominated by Polylepis
and Gynoxys and grow in settings sheltered from re. In general these
woodlands become less frequent and smaller with increasing
elevation and decreasing temperature (Fjelds, 1992, 2002). Regional
climate variation means that within the high Andean grasslands
(Puna) there are numerous different oristic communities; however,
palynological differentiation of these communities is problematic
because key taxa (e.g. Asteraceae, Cyperaceae and Poaceae) cannot be
differentiated below family level using a light microscope.
Two key components of the aquatic vegetation in the high Andes
are Isotes (a shoreline quillwort) and Myriophyllum (a milfoil). Isotes
species can reside in moist gullies but the largest population densities
are at lake margins. The upper elevational limit of Isotes has been
suggested to be c. 4200 masl because it cannot survive where ice
forms along the lake margins (Edwards et al., 2000; Macluf et al.,
2003; Liu et al., 2005). When found in lacustrine sediments, it has
been suggested that N90% Isotes (relative to the terrestrial pollen
sum) may be interpreted as near-shore conditions (Edwards et al.,
2000). The persistence of Isotes during past glacial times at Lake
Titicaca (Paduano et al., 2003) may reect local microclimatic
conditions that prevented the lake from freezing (Bush et al., 2010).
Myriophyllum sp. are typically submerged aquatic plants that can
grow in c. 3 m water depth; although a terrestrial species, M. quitense,
has been found in Bolivia (Ritter and Crow, 1998). Under frost-free
conditions Myriophyllum spp. are capable of spreading quickly by
vegetative or sexual reproduction (Ritter and Crow, 1998).
Within Andean lakes and freshwater streams non-motile green
algae of the genus Pediastrum sp. are common (Hutchinson, 1967).
The presence/absence of Pediastrum has been used in paleoecological
reconstructions as an indicator of changing water levels (Jankovsk
and Komrek, 2000) and nutrient availability (Foster et al., 2006).
Pediastrum spp. require fresh water, therefore the presence Pedia-
strum spp. can be interpreted as indicative of higher lake levels in Lake
Titicaca.
1.3. Fire on the Altiplano
Fire shapes modern Andean terrestrial vegetation communities,
and is of considerable importance in planning conservation and
carbon-sequestration strategies in the High Andes (Granda, 2005).
Understanding whether Puna systems burned prior to the arrival of
humans is critical to establishing accurate models of tree-line, carbon
storage and wildlife potential of these mountain areas. Prior to this
study, it was unknown whether the Puna burned naturally before
human occupation.
Charcoal in sediment cores provides evidence of local and regional
re events (Clark and Hussey, 1996; Clark and Royall, 1996; Clark and
Patterson, 1997) and can be used to interpret re histories (Whitlock
and Larsen, 2001). Paduano et al. (2003) documented charcoal at c.
 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214 203

14.3 ka in the previous Titicaca record, NE98-1PC, but based on that
age could not denitively state whether those res were natural or
evidence of extremely early human occupation.
1.4. Research questions
Core LT01-2B from Lake Titicaca provides the longest continuous
terrestrial paleoecological record derived from a lake in the Southern
Hemisphere, providing the rst opportunity to study vegetation
change over the past c. 370 ka. This study asks the following
questions: (1) How did the paleoclimates of Lake Titicaca inuence
vegetation change over the last c. 370 ka? (2) Is re a natural
phenomenon on the Altiplano during the past c. 370 ka? The present
results will be interpreted within the broader climate context
previously presented by Fritz et al. (2007).
2. Methods
2.1. Site description
The Altiplano covers c. 200,000 km2 of the Central Andes. The
climate of the Altiplano is semi-humid in the north with increasing
aridity towards the south (Lenters and Cook, 1997). Precipitation
across the Altiplano is strongly seasonal with c. 80% precipitation
occurring between December and March (Roche et al., 1992). In the
northern portion of the Altiplano, at Lake Titicaca, the average
precipitation is c. 889 mm/year and mean annual temperature is c.
8 C. South of Lake Poop the salars (saltpans) of Uyuni and Coipasa lie
within a zone of extreme aridity and the modern landscape is sparsely
vegetated. The major vegetation type of the Altiplano is Puna
grassland (Fig. 1).
Lake Titicaca (Bolivia/Peru; 3810 m above sea level; 1617S,
68.570W) currently covers c. 8500 km2 of the Altiplano and lies
between the Cordilleras Oriental and Occidental. The lake is composed
of a main basin (Lago Grande) and a sub-basin (Lago Huiaimarca).
Today, Lake Titicaca is nearly a closed basin lake with evaporation
accounting for 90 to 100% of the water loss from the lake in the period
of instrumental measurement (Roche et al., 1992). The remainder of
the water loss occurs via discharge into the sole outlet of Lago
Huinaimarca, the Rio Desaguadero. Lago Grande and Lago Huinai-
marca are joined by the Straits of Tiquina with a sill depth of
3788 masl, approximately 22 m below the mean water level of
modern Lake Titicaca. The sill that regulates outow from Lago
Huinaimarca is at 3804 masl. When the water level of Lago
Huinaimarca drops below this level, outow ceases and can reverse
(Wirrmann et al., 1992). Ouow from Lago Huinaimarca has been

Fig. 1. Core LT01-2B, east of Isla del Sol, was raised from the deepest point of Lake Titicaca, c. 235-m water depth. Whereas NE98-1PC was in 140 m water depth and LT01-3B was in
40 m water depth. The lowest lake-level in the late-Pleistocene at ca. 33 ka is marked, at which time Huiaimarca would have been dry.
Figure adapted from D'Agostino et al. (2002).
204 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214
regulated articially since the completion of a dam at Desaguadero in
2001.
Each year, the level of Lake Titicaca rises and falls by approximately
one meter due to wet season inow and dry season evaporation.
There has been considerable inter-annual variability of lake level in
the 20th century with a total range more than 6 m. In the late
Holocene (since 3500 ka), Lago Huinaimarca lake level varied
approximately 20 m between wet and dry periods (Abbott et al.,
1997). In the middle Holocene dry period, the level of Lago Grande
was as low as 90 m below present (Cross et al., 2000). And, as
mentioned previously, during MIS 5, lake level fell to approximately
200 m below present (D'Agostino et al., 2002; Fritz et al., 2007).
2.2. Sediment samples recovered from piston coring and drilling of Lake
Titicaca
Several coring expeditions recovered piston cores from deep
waters in Lake Titicaca using the R/V Neecho. These results are
summarized in Baker et al. (2001b) and Rowe et al. (2003). In 2001,
Paul Baker, Sheri Fritz and Geoffrey Seltzer led a coring expedition to
Lake Titicaca that recovered sediments using the GLAD800 system
(Fig. 1). Duplicate cores were raised from three different coring sites.
Recovery in each core was nearly complete. Sedimentologic, geo-
chemical, and diatom assemblage date were used to reconstruct lake
levels and to discuss the causes of paleoclimate variations at orbital
(Fritz et al., 2007) and millennial (Fritz et al., 2010) timescales. Core
LT01-2B was recovered from the deepest coring site in Lake Titicaca
(235 m water depth; the maximum depth of Lake Titicaca is 285 m).
Two other cores recovered from Lake Titicaca are also discussed in this
paper: i) drill core LT01-3B was recovered from the Chua basin, the
deepest portion of Lago Huinaimarca, and ii) piston core NE-98-1PC
was raised from Lago Grande from 140 m water depth.
Fig. 2. Total pollen concentration data for fossil pollen from Lake Titicaca are plotted with fossil charcoal, CaCO3 and MS data are from Fritz et al. (2007). Interglacials are characterized
by pollen concentrations N20,000 grains/cm3 and higher charcoal presence.
2.3. Chronology
The LT01-2B age model was developed using a radiocarbon
chronology for the upper 26 m of the core. A cluster of four U-series
ages from calcium carbonate (CaCO3) rich sediments at 45.44 m
below lake oor (mblf) provided a mean age of 122.8 7.7 ka (Fritz et
al., 2007; Fritz et al., 2008). The chronology beyond the U-series dates
was based on tuning the Lake Titicaca calcium carbonate record to the
Vostok CO2 chronology (Fritz et al., 2007). Details regarding
construction of the chronology can be found in Fritz et al. (2007).
Sampling for pollen and charcoal was conducted at 20 cm intervals
throughout the glacial portions of the core and at 10 cm resolution
throughout the interglacial portions. The sampling interval averages c.
500 years during glacials and c. 150 year during interglacials.
2.4. Subsample preparation and analysis
Samples were processed using standard palynological protocols
(Faegri and Iversen, 1989), including a density separation using
sodium metatungstate (specic gravity 2.10). Concentration data for
terrestrial and aquatic pollen and spores were calculated using a spike
of exotic Lycopodium spores (18,588 spores/tablet; Stockmarr, 1972).
A sum of 300 pollen grains (excluding aquatic pollen and spores) or
2000 exotic Lycopodium was counted using a Zeiss Axioskop
photomicroscope (40 and 100). Percentage abundance for all
taxa (including aquatic pollen and spores) was calculated relative to
the main sum.
Pollen was identied using the Florida Institute of Technology
modern pollen reference collection of N3000 Neotropical pollen types,
an electronic pollen database (Bush and Weng, 2007), and a selection
of photographs from previous studies (Paduano, 2001) and published
descriptions (Hooghiemstra, 1984). Pollen types of Asteraceae were
separated into c. 40 morphotypes during counting, but those types
 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214
were re-combined for statistical analysis. Pollen taxa were classied
as Andean forest, Puna, shrub belt, and aquatic categories based on
Troll (1968) and Graf (1981).
Approximately 530 samples were counted to provide the abun-
dance of all pollen types, and an additional 100 samples were counted
at a lower taxonomic resolution (12 key types and others) to
provide total pollen concentration and Isotes abundance data.
Charcoal was processed for each sample using standard processing
procedures (Clark and Patterson, 1997), with sample ltration to
retain particles N180 m. Although charcoal was processed across all
MIS stages, a majority of the samples focused on the interglacial
periods. Each processed charcoal sample was quantied under an
Olympus photomicroscope at a magnication of 20. Charcoal area
was calculated using NIH-image software. Charcoal was analyzed in
314 samples.
These results were graphed in C2 (Juggins, 2003). TILIAGRAPH and
CONISS software were used to calculate the pollen zones (Grimm,
1987, 1992). Due to size constraints of the TILIAGRAPH program,
approximately 100 glacial-age samples were excluded from the
cluster analysis. These samples were not transition samples; therefore
their elimination did not bias any zoning results. Detrended
Correspondence Analysis (DCA) (Hill, 1979; Ter Braak, 1988), using
the PC-ORD 4 software version (McCune and Mefford, 1999), was run
on percentage data for taxa represented at N3% in the record.
3. Results
3.1. Pollen concentration and charcoal
Fossil pollen concentrations in the 630 samples examined from
LT01-2B were generally low (mean pollen concentration = 4097 -
grains/cm3) with the highest recorded value c. 86,000 grains/cm3.
Charcoal was common during each interglacial, but rare during glacial
phases. Pollen concentrations fell to values below 200 grains/cm3
during glacial periods, but increased to values typically greater than
Fig. 3. Percentage diagram for taxa with N3% representation. Grouped taxa were based on vegetation descriptions of Troll (1968) and Graf (1981). Shaded areas represent
interglacials MIS 1 (Holocene), MIS 5e, MIS 7, and MIS 9.
205
24,000 grains/cm3 during interglacial periods (Fig. 2). A rare instance
of glacial-age charcoal was found c. 60 ka. The rst appearance of
charcoal in the record was approximately 320 ka, during MIS 9. Pollen
and charcoal concentrations were closely correlated, likely due to
both dilution and productivity changes.
3.2. Description of pollen zones
Sixteen local pollen zones (each containing at least four samples)
were determined for the LT01-2B sediments using CONISS (sum of
squares values N50) on the complete pollen data set. The strongest
division within the record separates samples older/younger than
138 ka. In the description of zones, Puna taxa were ubiquitous
throughout the pollen record, unless otherwise noted. In the interest
of brevity, comments will be limited to taxa occurring at N3% of each
pollen sum (Fig. 3).
3.2.1. LT-1 (135125 mblf; 370330 ka)
Pollen concentrations of this section ranged from c. 100 grains/cm3
at c. 366 ka to 5682 grains/cm3 at c. 359 ka (mean = 1081.26 grains/
cm3). The pollen of Puna taxa were consistently present, ranging from
80 to 100%. Asteraceae pollen averaged 24% of the pollen sum,
however, they peaked (c. 70%) at c. 346 ka. Amaranthaceae pollen
consistently accounted for 5% of the sum, with a maximum of 28% at c.
330 ka. Poaceae pollen averaged approximately 45% of the taxa. Of
note are Caryophyllaceae pollen, which averaged around 8% repre-
sentation, but peaked at 30% at c. 362 ka and Polylepis pollen that
reached 7.6% at c. 357 ka.
Representation of aquatics varied during this zone. Myriophyllum
pollen were only present at 330 ka, while Pediastrum algal cell
concentrations were low throughout this time. Isotes spore concen-
trations peaked around c. 362 ka with 2263 spores/cm3, percentages
of Isotes spores N100% occurred at 336 (108%) and 362 ka (121%;
Fig. 4). Charcoal particulates were absent within this zone (Fig. 2).
206 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214

Fig. 3 (continued).
J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214 207

Fig. 3 (continued).
208 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214
Fig. 4. Percent occurrence of fossil Isotes spores relative to pollen sum in Lake Titicaca,
Bolivia/Peru, plotted against time.
3.2.2. LT-2 (125115 mblf; 330301 ka (MIS 9))
Pollen concentrations varied from 21 grains/cm3 to 57,080 grains/cm3
at the peak of the interglacial (c. 324 ka), with a mean concentration of
8653 grains/cm3. Puna taxa pollen were present from 50 to 100%. Around
310 ka, Andean forest taxa pollen percentages were c. 23%. Alnus pollen
representation reached 21% at c. 314 ka, and Juglans pollen had their
highest value in the core at c. 311 ka (10.2%), while Podocarpus
representation did not exceed 7%. There was a peak in Amaranthaceae
pollen concentrations and percentages around c. 322 ka (60%). Polylepis
was consistently present from c. 327301 ka, at between 2 and 33% (peak
at c. 327 ka) during MIS 9.
Myriophyllum pollen concentrations were consistently high during
this time, with a spike at 33,054 grains/cm3 at c. 324 ka. Isotes
representation was greater than the total pollen sum from c. 327 ka
(Fig. 4). Pediastrum algal cell concentrations exceeded 100,000 colo-
nies/cm3 at c. 327 and 320 ka. Charcoal was present at 0.6 mm2/cm3 at
c. 320 ka and declined to trace amounts by 315 ka, mirroring the
decline of pollen concentration within this zone (Fig. 2).
3.2.3. LT-3 (115110 mblf; 301288 ka)
Pollen concentrations varied from 619 to 4990 grains/cm3
(mean = 1585 grains/cm3). Puna taxa pollen representation domi-
nated this time period, with percentages ranging from 55 to 90%.
Juglans pollen was present (up to 12.5%) in a few samples at c. 307 ka,
while Alnus pollen reached 18.5% at c. 297 ka. Podocarpus pollen was
consistently present in the samples, but in small numbers. Polylepis
pollen was also consistently present between c. 301 and 297 ka, with a
maximum percentage of 22% at c. 300 ka.
The aquatic pollen and spore concentrations were low with the
exception of concentrations of Pediastrum algae. Pediastrum concen-
trations increased to c. 10,000 colonies/cm3 from c. 299 to 296 ka and
then declined. Charcoal values were very low within this zone, never
exceeding 0.15 mm2/cm3 (Fig. 2).
3.2.4. LT-4 (110100 mblf; 288260 ka)
Pollen concentrations ranged from 501 to 3615 grains/cm3
(mean = 1615 grains/cm3). Pollen representation was dominated by
Puna taxa.
Myriophyllum pollen representation peaked at 61,373 grains/cm3
at c. 274 ka Pediastrum algal concentrations were low except at
c. 285 ka when concentrations reached 4335 colonies/cm3. Isotes
spore concentrations peaked at c. 260 ka during a period of high
pollen concentrations (mean = 261 spores/cm3). No charcoal parti-
cles were recorded within this zone (Fig. 2).
3.2.5. LT-5 (10093 mblf; 260238 ka)
Pollen concentrations were variable, ranging from 589 to
10,042 grains/cm3 during this time period, with a majority of samples
N2000 grains/cm3. Pollen of aquatic taxa were present, but in low
concentrations. No charcoal particles were recorded within this zone
(Fig. 2).
3.2.6. LT-6 (9376 mblf); 238211 ka (entering into MIS 7)
Pollen concentrations varied during this time period, ranging from
128 to 20,023 grains/cm3, with the highest concentrations between
c. 213 and 211 ka. Polylepis pollen occurred frequently, with peaks as
high as 30% between c. 237 and 236 ka, with a mean occurrence of 9%.
Charcoal area also sporadically increased within this zone, reaching
values of 2.0 mm2/cm3 around 211 ka. (Fig. 2).
Myriophyllum pollen representation was low except at c. 238 ka,
when it reached 19,292 grains/cm3. Pediastrum algal concentrations
reached a peak of 12,334 colonies/cm3 and Isotes spore concentra-
tions also peaked (1509 spores/cm3) at c. 215 ka.
3.2.7. LT-7 (7670 mblf); 211196 ka (MIS 7)
Pollen of Puna taxa increased from about 80 to 90% in this zone.
Polylepis pollen was frequently present, peaking at 34% around
c. 205 ka.
Pediastrum algal concentrations had a few spikes in which
concentrations were c. 1000 colonies/cm3: c. 209 ka, 199 ka, and
196 ka. Isotes spore representation was N100% at c. 204 ka (Fig. 4), but
the highest concentrations occurred at c. 210 ka (1064 spores/cm3).
Charcoal particles were abundant in this zone (Fig. 2).
3.2.8. LT-8 (7057 mblf; 196150 ka)
Pollen concentrations ranged from 21 to 3054 grains/cm3. Podo-
carpus pollen representation was frequently b10%, but increased to
1425% from 181 to 180 ka. Polylepis pollen representation reached
22% at c. 155 ka (mean = 5%).
Pediastrum algal concentrations were variable, ranging from 1751
to 3780 colonies/cm3 from c. 194 to 192 ka, which coincided with the
highest total pollen concentrations. Charcoal was not found within
this zone.
3.2.9. LT-9 (5754 mblf); 150139 ka (transition into MIS 5e)
Pollen concentrations were lowranging from 299 to 8373 grains/cm3
(mean = 1060 grains/cm3). Pollen of Puna taxa were consistently
represented at 8090% except for a decline c.150 ka. Asteraceae pollen
representation varied every c. 1000 years, ranging from 5 to 21%.
Podocarpus pollen representation ranged from 8 to 13% from c. 150 to
146 ka, but peaked at 22% at c.144 ka. Polylepis pollen representation
increased to approximately 15% around c. 139 ka.
 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214
Isotes spore concentrations averaged c. 284 spores/cm3, but
increased to 1881 spores/cm3 at c. 139 ka Pediastrum algal concentra-
tions were N2000 colonies/cm3 from c. 142 to 141 ka. Unidentied
trilete spores were high during this zone, averaging 1459 spores/cm3.
There was no charcoal found during the transition into MIS 5e (Fig. 2).
3.2.10. LT-10 (5445 mblf); 139115 ka (MIS 5e)
Total pollen concentration was extremely high at c. 127 ka
(75,632 grains/cm3). The average concentration was 15,165 grains/
cm3, much higher than any other portion of this record. A sharp
decrease in the pollen of Puna taxa marked the beginning of this time
period, but then increased by 133 ka. Poaceae pollen representation
did not dominate the spectra (mean = 13%), rather Amaranthaceae
pollen rose to 84% between c. 133 and 123 ka. Polylepis pollen
representation ranged from 20 to 28% from c. 138136 ka, and
reached c. 30% at c. 125 ka and 123 ka. At the same time, charcoal area
increased to over 0.8 mm2/cm3. However, charcoal area maintained
low values after 123 ka and for the rest of the zone (Fig. 2).
Aquatic taxa pollen and spore concentrations were low during MIS
5e, but Pediastrum algal concentrations were extremely high at the
beginning of this zone, ranging from 64,420 to 296,890 colonies/cm3
from c. 137 to 136 ka.
3.2.11. LT-11 (4540 mblf; 11563 ka)
Total pollen concentrations ranged from 288 to 15,836 grains/cm3.
Concentrations remained high until c. 103 ka and subsequently
dropped to less than 6000 grains/cm3 until c. 73 ka, and then
b2000 grains/cm3 until c. 63 ka. Although Puna taxa were dominant
in the pollen spectra, Polylepis pollen representation reached two
main peaks during this time19% at c. 114 ka, and 18% at c. 93 ka.
Isotes spore percentages were N100% at c. 65 ka (Fig. 4).
Concentrations were high as well, with a maximum 2418 spores/
cm3 at c. 73 ka. The unidentied monolete and other trilete spores
were relatively high, with concentrations at 221 and 1057 spores/cm3,
respectively. Only two charcoal samples were taken during this zone,
and both resulted in low charcoal area, less than 0.2 mm2/cm3 (Fig. 2).
3.2.12. LT-12 (4035 mblf; 6355 ka)
Total pollen concentrations ranged from 128 to 3463 grains/cm3
(c. 62 ka). The pollen of Puna taxa, mainly Asteraceae and Poaceae,
were common.
Isotes spore percentages were high (115120%) at c. 62 and 59 ka
(Fig. 4). Concentrations were high at 62 ka (1293 spores/cm3) and from
c. 60 to 59 ka (10251110 spores/cm3). Trilete spores were high during
this time period, averaging 2182 spores/cm3, peaking at 11,417 spores/
cm3 at c. 62 ka. During peak pollen and Isotes concentrations, charcoal
was also present. Although in small values overall, charcoal did increase
to approximately 0.4 mm2/cm3 c. 60 ka (Fig. 2).
3.2.13. LT-13 (3525 mblf; 5542 ka)
Total pollen concentrations were b2000 grains/cm3, ranging from
2221996 grains/cm3. There were three occasions when Isotes spore
percentages were N100%: 53 ka (123%); 52 ka (105%); 50 ka (189%)
(Fig. 6). At 50 ka, that was also the time of the highest spore
concentration (1473 spores/cm3). Charcoal was not found within this
zone (Fig. 2).
3.2.14. LT-14 (2515 mblf; 4228 ka)
Pollen concentrations were low, ranging from 779 to 2274 grains/
cm3 (mean = 1302 grains/cm3). Spore concentrations were high,
especially the triletes, which averaged about 2045 spores/cm3.
Charcoal was not found within this zone (Fig. 2).
3.2.15. LT-15 (154 mblf; 2820 ka)
Pollen concentrations were lowranging from 177 to 5085 grains/
cm3. Caryophyllaceae pollen representation was high from c. 25
209
24 ka, ranging from 8 to 13%, but was rare after that time. Podocarpus
pollen representation was c. 17% at c. 23 ka, and then 14% at c. 24 ka.
Isotes spore concentrations were N2100 spores/cm3 and its percen-
tages were N100% several times, with an average of 105% for this zone
(Fig. 4). Charcoal was not found within this zone (Fig. 2).
3.2.16. LT-16 (40.1 mblf; 203 ka)
Pollen concentrations were low ranging from 224 to 1626 grains/cm3
until c. 17 ka and then increased to 20,279 grains/cm3 at c. 9 ka. Trilete
spores concentrations averaged 1076 spores/cm3 during this zone.
Polylepis pollen representation was low, but increased to 24% at c. 17 ka.
Aquatic pollen and spore concentrations uctuated throughout
this time period. Pediastrum algal concentrations peaked at 1810 col-
onies/cm3 at c. 3 ka. Isotes spore concentrations were high during this
zone, especially from c. 149 ka. Charcoal was only found at c. 3 ka
with area calculated as less than 0.2 mm2/cm3.
3.3. Detrended correspondence analysis
The positive extreme of Axis 1 of the DCA was characterized by
Amaranthaceae, while the negative extreme was characterized by
Polylepis and Andean forest elements. Axis 2 is characterized by
Poaceae at the positive extreme and Polylepis and Andean forest taxa
at the negative extreme. A summary diagram of the DCA of percentage
pollen data from Lake Titicaca (Axis 1 versus Axis 2) revealed a
clustering of several zones (Fig. 5). The data points from 115 to 139 ka
isolated MIS 5e from all other samples. Glacial-age samples have high
scores on Axis 2 and low scores on Axis 1, whereas interglacials
generally have low scores on Axis 2. The Holocene samples group
close to those of MIS 7 and transitional samples between glacial and
interglacial conditions for MIS 5e and 9.
Axis 1 and 2 values were plotted against age to identify trends in
the dataset over time (Fig. 6). DCA Axis 1 provides a range of values
with glacial-aged samples generally exhibiting values from 70 to 180.
Interglacial values are generally higher with many values N200. It can
be seen that MIS 7 (max value 210) and MIS 1 (180) values are not as
high as those of the other interglacials (e.g. MIS 9 = 295, 5e = 305).
High pollen concentration values (N24,000 grains cm3) are only found
in interglacial intervals.
The plot of Axis 2 versus time reveals periods in which general
trends toward positive values are apparent corresponding to glacial
episodes and the interglacials marked by more negative values.
While MIS 9 and 5e are represented by strongly clustered responses
on both axes (but particularly on Axis 1), MIS 7 is represented by
two or three strong changes (according to the precise denition of
the boundary), which is consistent with a protracted interglacial
comprised of three warm events as recorded from higher latitudes
and marine settings (Shackleton, 2000; Siddall et al., 2003; Tzedakis
et al., 2004).
4. Discussion
4.1. Do palynological data complement LT01-2B geochemical data?
The chronology developed by Fritz et al. (2007, 2008) was built
around both 14C and U/Th ages, and cross-correlation with the Vostok
ice-core record. Because the LT01-2B chronology was constructed
independently from any biological data some basic predictions that
could be tested with palynological data were: i) glacial periods will
have very low pollen concentrations, ii) interglacial periods will have
high pollen concentrations, and iii) interglacial periods will have a
higher abundance of thermophilous taxa than glacials, e.g. increased
representation of arboreal elements, and decreased occurrence of
Isotes spores (Paduano et al., 2003).
Previous studies have suggested that high magnetic susceptibility
within the LT01-2B core corresponds with glacial and elevated CaCO3
210 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214
Fig. 5. Summary results of sample scores for the DCA ordination of taxa with N 3% representation. The mean Axis 1 and Axis 2 scores were plotted for each of the zones, and the size of
the circle represents the number of samples (487) in each zone. Interglacial zones are shown in red circles, glacial zones in blue, and transition zones are plotted in purple.
with interglacial periods (Fritz et al., 2007, 2010). Our fossil pollen
data show low concentrations related to high magnetic susceptibility
and high concentrations related to high CaCO3; which conforms to our
prediction of biotic response to glacial and interglacial changes. In
each transition from glacial to inferred interglacial, the Andean treelet,
Polylepis, appears in the pollen record amid clear indications of
ecosystem change, e.g. rise in pollen concentration N25,000 grains/
cm3, increased charcoal occurrence, Podocarpus sp. representation
increasing from 8% to 22%, and sudden positive shifts in the Axis 1 DCA
scores. The timing of the proposed interglacials based on the CaCO3
concentrations used by Fritz et al. (2007) is consistent with the peak
of warming inferred from the fossil pollen record associated with each
event. Because the pollen is more sensitive than the chemical record
to glacial termination, i.e. before the lake level is lowered sufciently
to precipitate carbonate, and continues to provide a sensitive proxy as
lake level rises, the duration of interglacial warming indicated by the
pollen record is longer in each instance than that of the CaCO3 record.
If the chronology suggested by Fritz et al. (2007) is accepted, the rst
signs of Polylepis expansion that continue into MIS 5e occur begin as
early as 144 ka. Polylepis tarapacana can grow to elevations of c.
5000 m under modern conditions, and its principal limitation is re.
Although still cold at 144 ka, the expansion of Polylepis at 144 ka
would be consistent with a wet event at the peak of MIS 6 that
allowed development of woodlands near the lake.
Onset and termination of most of the interglacials is fairly distinct,
however, MIS 7 was unique in a number of ways. First, it developed
from what was, at least locally, a glacial (MIS 8) that was not quite as
cold as MIS 10, 6 or 2. Second, as in other locations, MIS 7 did not build
to a simple thermal peak (Shackleton, 2000; Siddall et al., 2003), but
had two strong and one weaker warming events, leading to a longer
overall event (Tzedakis et al., 2004).
Representation of the Andean treelet, Polylepis, which is an early
marker of all the other interglacials, is evident as early as c. 238 ka.
However, the peak pollen concentrations of MIS 7 (N24,000 grains/
cm3) do not occur until c. 218 ka. This record parallels the event as it
was recorded in Europe, where MIS 7 consists of pulses of warming as
opposed to a long continuous trend (Roucoux et al., 2008).
4.2. What were the paleoclimates of Lake Titicaca in the last 370 ka?
Axis 1 of the DCA is interpreted to represent decreasing moisture
availability, whereas Axis 2 is decreasing temperature. The placement
of MIS 5e samples at the positive extreme of Axis 1 and the negative
extreme of Axis 2 identies this as the warmest and driest period of
the last 370 ka, a nding consistent with data from core LT01-3B
(Gosling et al., 2008) and with diatom data from LT01-2B (Fritz et al.,
2007). When plotted against time, Axis 1 scores for the period
included in MIS 9 were also N300, well above the maximum MIS 7 and
Holocene values. According to this analysis, MIS 9 and MIS 5e were
drier than MIS 7 and the Holocene. The transition zones between full
glacial and the interglacial conditions of MIS 9 and 5e were clustered
with the milder interglacials MIS 1 (the Holocene) and MIS 7. This
similarity suggests broadly similar communities form at the transition
from glacial to interglacial, but some glacial periods go on to more
extreme conditions.
A striking difference between glacial and interglacial stages was in
the productivity of the landscape. Glacial times were represented by
very low pollen concentration as the local environment became a
peri-glacial habitat. In many cases, pollen concentration values were
b100 grains/cm3. As no taxa reliably distinguish Puna Brava from
Puna palynologically (Troll, 1968; Graf, 1981), concentrations rather
than percentages are probably the best means to distinguish these
systems.
Glacial-age deposits of MIS 10 formed the base of core LT01-2B.
Low pollen concentrations and the Puna Brava signature in these
sediments suggested strong cooling. The modern Puna Brava lies
between c. 4500 and 5300 m, implying a glacial-age descent of
vegetation of c. 7001500 m, equivalent to a cooling of c. 48 C,
which is within the range of the c. 8 C cooling inferred for the High
Plains of Bogot at this time (Hooghiemstra, 1984).
The transition into the MIS 9 interglacial was characterized by
increasing temperature and by an increasing moisture balance.
Andean woodlands were represented by Polylepis and, at the height
of the interglacial, a community dominated by Amaranthaceae.
Between the increases of Polylepis and Amaranthaceae there was a
 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214
Fig. 6. Sample scores for DCA Axes 1 and 2 of fossil pollen data from Lake Titicaca
plotted against time.
peak of Myriophyllum. This genus is generally a submerged aquatic of
shallow, fresh to slightly brackish aquatic systems. Myriophyllum
quitense, the species that occurs at Lake Titicaca has been observed to
ower as a terrestrial plant under frost-free conditions (Ritter and
Crow, 1998). The huge abundances of Myriophyllum and relative
scarcity of Isetes in MIS 9 suggests lowered lake level with substantial
shallows or exposed mudats. The peak of Amaranthaceae probably
represents dry, possibly salt at, conditions along the lake margin, and
their presence coincides with evidence of increased lake solute
concentration and sedimentary deposition of carbonate (Fritz et al.,
2007). The transition from Myriophyllum to Amaranthaceae is
consistent with a declining P:E ratio. Warm, dry conditions appear
to have dominated the peak of MIS 9, consistent with a warming of 1
2 C.
Later in the interglacial, a surprising presence is that of Juglans at
percentages as high as 12.5%. Although peaking at 307 ka, Juglans
occurred intermittently at low values throughout interglacial phases.
This genus, which has very distinctive pollen, generally favors warm
moist conditions and is seldom documented above 2500 m elevation.
Although the presence of this taxon could be used to infer a c. 1300 m
upslope migration, which could translate into a c. 6 C warming, no
other Andean forest species is documented responding in the same way.
We do not consider this amount of warming likely. Two other
possibilities could explain this observation. First, Juglans may have
changed its ecological range through adaptation. Second, the species
represented by the pollen may have been a high elevation species that is
now extinct. While there is no clear evidence of a species changing its
elevational adaptation radically across a glacial cycle, at least one tree
with a fossil range outside that of most extant species of that genus is
known to have gone extinct during the LGM (Jackson and Weng, 1999).
Whatever the explanation for Juglans growing close to Lake Titicaca it is
apparent that this was a community without modern analog. Abundant
charcoal in the sediment during MIS 9 indicates that there was fuel for
re in the nearby landscape (e.g. Juglans and Polylepis).
MIS 8 was cold, however, probably not as cold as other glacial
sections of the core. Normal Puna conditions remained in the
Altiplano until the transition into MIS 7. Based on insolation to the
upper atmosphere, seasonality in MIS 7 was stronger than in MIS 9
(Berger, 1978). This period appears to have been climatically unstable
211
with conditions that were, intermittently, as warm as the mid-
Holocene (possibly 12 C warmer than modern). However, the
climate never stabilized either as a warm, wet system (productivity
would have been higher) or as a warm, dry system (lake level would
have been lower). While MIS 7 was not necessarily a cooler
interglacial than the others, it may have been a more mixed climate
with periods of warmth interrupted by cooler conditions, and equally
unstable precipitation patterns.
MIS 6 was apparently a very cold period, with pollen concentra-
tions declining to very low levels (b1000 grains/cm3). Puna taxa were
present, but scarce local vegetation allowed for stronger representa-
tion of pollen transported from Andean forests (Bush, 2000).
Increased magnetic susceptibility values indicate increased rates of
detrital sediment input during this time, likely due to local glacial
advance (Fritz et al., 2007).
MIS 5e was similar to MIS 9 in intensity, but the duration of peak
interglacial conditions was longer. As in MIS 9, Polylepis and
Pediastrum marked the transitions into and out of MIS 5e, indicating
warm and wet conditions. Polylepis attains its highest representation
of c. 32% during transitions between glacial and interglacial condi-
tions. These percentages are similar to those from modern lakes that
are completely surrounded by Polylepis (Hanselman, unpublished
data) and supports the view that Polylepis was once more abundant
than it is today (Ellenberg, 1979; Fjelds, 1992). Myriophyllum does
not exhibit peaks during MIS 5e. The interglacial was arid, with
unparalleled concentrations of Amaranthaceae (Hanselman et al.,
2005). High concentrations of CaCO3, low ratios of planktic-to-benthic
diatoms indicate very low lake levels and very dry climate (Fritz et al.,
2007). The pollen data are consistent with this conclusion. When the
level of Lago Grande dropped below the Tiquina sill, Lago Huinaimarca
desiccated. Bush et al. (2010) emphasized the importance of positive
feedback mechanisms on microclimates at Lake Titicaca and how
contracted lake level could further enhance aridity.
By ca. 6255 ka, full glacial conditions were once again established
on the Altiplano. Although extreme cold results in very little pollen
deposition during the last glacial, the interpretation of cold and wet
conditions is supported by the presence of aquatics and Polylepis.
Although Polylepis was commonly associated with interglacial transi-
tions, it can be distributed up to c. 4800 m elevation (Fjelds, 1992;
Fjelds and Kessler, 1996; Fjelds, 2002), which is close to the snow-
line with an average modern temperature of approximately 0 C. The
inference of a c. 48 C cooling, similar to that of MIS 10 would apply
to MIS 2 and is consistent with the LGM record from Lake Pacucha
(Valencia et al., 2010). Despite such cooling, Polylepis survived.
Throughout this record Polylepis has its strongest occurrences in the
termination and inception of ice ages rather than at the peak of
interglacial climates; an observation that was further explored by
Gosling et al. (2009).
4.2.1. Comparison with regional records
The Fuquene and Funza records from the High plain of Bogot,
provide replicate cores spanning the period represented in Titicaca LT01-
2B (Van der Hammen and Gonzlez, 1959; Van der Hammen, 1974;
Hooghiemstra, 1984; Hooghiemstra and Cleef, 1995; Hooghiemstra and
Van der Hammen, 2004). As in our Titicaca record, the chronology of the
Colombian sites was established with some radiometric dating, but the
detailed chronology of the Colombian sites was established through
tuning to the ODP site 677 record (Van 't Veer and Hooghiemstra, 2000).
Direct comparisons of the sites are complicated by their different
elevations (the Colombian records are from c. 2550 m elevation
whereas Titicaca lies at 3810 masl) which results in interglacial
glacial migrations between forest and Puna in the Colombian records,
but Polylepis-Puna and Puna brava at Titicaca. Also, one of the most
important taxa found in Colombia, Quercus, which increased in
abundance to N35% of the pollen sum during the last glacial period, is
not found south of the equator. In the Colombian records, interglacial
212 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214
peaks of Alnus alternated with glacial peaks of Poaceae (e.g.
Hooghiemstra, 1984). The basic pattern of warm interglacials leading
to forested settings with species migrating steadily upslope found in
the Colombian records is not replicated in Titicaca. Indeed, the upper
montane cloud forests of one interglacial look much like another in
the Colombian record, and the climatic differences are less than the
biogeographic ones associated with the Quercus invasion in the late
Pleistocene (Van 't Veer and Hooghiemstra, 2000). In contrast to this
pattern, in the Titicaca basin, the initial warming of each interglacial
produces an upslope migration of woodland taxa, but in MIS9 and MIS
5e that migration stalled as the system ipped to a re-prone, arid,
state as local climatic effects apparently prevented further expansion
of forest (Bush et al., 2010).
The difference in vegetation history points to another major
distinction between the Colombian and Peru/Bolivian sites. Histori-
cally, the most extreme interglacials (MIS 5e and 9) at Lake Titicaca
were the driest times in the record based on seismic data (D'Agostino
et al., 2002), CaCO3 deposition (Baker et al., 2005; Fritz et al., 2007);
halophytic diatom assemblages (Tapia et al., 2003), and the fossil
pollen history. In contrast to this pattern, it was the cold, full-glacial
periods in Colombia that had the lowest lake level (Velez et al., 2003),
while the interglacial periods appear to have been warm and wet.
To have the two hemispheres out of phase in terms of precipitation
is a basic expectation based on precessional forcing of their respective
wet and dry seasons (Baker, 2002; Clement et al., 2004; Ballantyne et
al., 2005). That precession strongly inuences tropical rainfall in the
lowlands is evident in speleothem (Wang et al., 2004; Cruz et al.,
2009) and lake records (Bush et al., 2002). A statistically signicant
peak of precessional inuence is also evident in the Funza 1 record
(Hooghiemstra et al., 1993). However, moisture availability, precip-
itation minus evaporation, is a strong function of surface air
temperature (as well as insolation, wind, and relative humidity).
Surface temperature has changed on eccentricity timescales over the
past ca 1 Ma with the waxing and waning of large ice sheets on the
continents and the rise and fall of atmospheric CO2 concentrations.
Fig. 7. Comparison of Isotes concentrations and proportion of biogenic silica in sediments from Lake Titicaca, Peru/Bolivia (Fritz et al., 2010) for the period between 20 and 60 ka.
Also shown is the O18 record for Greenland (NGRIP) (NGRIP, 2004). Peaks of biogenic silica are highlighted and reveal a general correspondence to troughs in Isotes concentration
and peaks in O18 abundance in Greenland. Heinrich events H2-6 are marked.
Thus, the time variation of moisture availability should contain
strong power at both precessional (precipitation, out of phase north
versus south) and eccentricity (temperature, globally symmetric)
frequencies.
South American tropical moisture availability also varies strongly
on millennial timescales with the northern and southern tropics
antiphased with respect to precipitationnorthern hemisphere dry,
southern hemisphere wet (e.g. Baker et al., 2001b; Peterson and Haug,
2006; Wang et al., 2004)in response to north Atlantic cold events
(Baker et al., 2009). This is illustrated in the Lake Titicaca record by the
correlation of biogenic silica minima, grain size maxima, and maxima
of carbon isotopic ratios of organic carbon with cold events recorded
in the Greenland ice cores (Fritz et al., 2010). Here we nd that Isetes
spore concentrations are high when biogenic silica contents are low
(Fig. 7). Because Isetes lives primarily as a nearshore submerged
aquatic and generally releases its spores underwater, offshore
transport of the spores is best achieved by the hyperpycnal ows
attributed by Fritz et al. (2010) to the wet phases.
4.3. Fire is a natural occurrence on the Altiplano
Charcoal is differentially transported, according to its size, to a
lake. Hence the size fractions of charcoal recovered from sediment can
be used as a proxy for the distance to re. An abundance of smaller
particles (65179 m) could indicate long-distance transport and
remote res, while larger particles (N180 m) could indicate res
close to the lake basin (Clark and Patterson, 1997; Paduano et al.,
2003). The measurement of charcoal particles N180 m used in this
study is an efcient technique to describe the occurrence of re close
the Lake Titicaca.
Core NE98-1PC correlated re with a possible early entry date for
human arrival of approximately 14.5 ka (Paduano et al., 2003). Here
we present the rst documentation of the presence of re in prior
interglacials.
 J.A. Hanselman et al. / Palaeogeography, Palaeoclimatology, Palaeoecology 305 (2011) 201214
Conditions have to be favorable for re (warm and dry) but there
also needs to be sufcient fuel for re (wet). During glacial periods the
Puna Brava vegetation that we infer based on pollen concentrations
apparently did not provide sufcient fuel load to carry a re despite
relatively high moisture availability. Probably at this stage, the
principal limitation on fuel development was cold rather than lack
of moisture. The exception to this observation is a period at c. 60 ka
when, apparently, climatic conditions did lead to re. Indeed for the
last 370 ka, the Altiplano was primarily composed of Puna Brava
vegetation, except for periods of transition into and out of inter-
glacials. These transitions were times of locally increased productivity,
resulting in sufcient fuel (e.g., Polylepis, Juglans) that lightning strikes
could initiate re. Fire is thus seen to be a natural component of
interglacial conditions in the tropical Andes within the modern Puna
zone.
5. Conclusions
A detailed fossil pollen analysis of a 370 ka record from Lake
Titicaca characterized four full glacial cycles. Each interglacial was
characterized by increased pollen (N20,000 grains/cm3) and charcoal
concentrations. Interglacials were warm and dry, but varied in
intensity. MIS 9 and MIS 5e were the most intense (driest)
interglacials, with high representation of dry taxa such as Amar-
anthaceae. MIS 7 was a protracted interglacial warming, though the
ora that developed around Lake Titicaca was more similar to that of
MIS 1 than MIS 5e or 9.
Though ice-free, the littoral regions of Lake Titicaca, were largely
unproductive during glacial periods. Lack of vegetation cover may
have contributed to the increased detrital sedimentation rates
observed during glacial times (Fritz et al., 2007). Pollen assemblage
changes are consistent with previous estimates of approximately 5 C
glacial cooling (Paduano et al., 2003).
Fire was a natural component of Andean ecosystems long before
the arrival of the rst human inhabitants. Fire was more frequent
during interglacial stages than during glacial. Apparently the cool, wet
conditions of glacials reduced fuel loads and decreased the likelihood
of ignition.
Acknowledgments
We are grateful to the staff at LacCore for their help with core
sampling and core logging. We wish to thank two anonymous
reviewers who improved earlier versions of this manuscript and also
our editor Peter Kershaw for his thoughtful comments. This study was
funded by NSF grant ATM 0317539 (to Bush), NSF grant AGS 0602329
(to Baker), and NSF Grant EAR 0602154 (to Fritz). This is publication
#3 of the Institute for Research on Global Climate Change at the
Florida Institute of Technology.
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