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The Holocene
1 –12
© The Author(s) 2016
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DOI: 10.1177/0959683616652702
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Introduction
Investigations of sediment records recovered from lakes in Costa
Rica have provided important insights into the nature of tropical
climate and landscape change and refined our understanding of
tropical climate variability in southern Central America during the
latest Pleistocene and Holocene. Multi-proxy paleoenvironmental
studies document a notable decrease in mean annual air tempera-
ture (MAAT) during the Last Glacial Maximum (LGM) and con-
siderable climate variability during the Holocene (Horn, 2007).
For example, pollen analysis of sediment extracted from La
Chonta bog (2310 m a.s.l.), located in the northern Cordillera
de Talamanca of Costa Rica, suggested that MAAT was 7−8°C
lower than the present between ~50,000 and 15,600 cal. yr BP
(Hooghiemstra et al., 1992; Horn, 2007), in close agreement with
estimates from reconstructions of paleoglaciers at higher eleva-
tions in the Cordillera de Talamanca (Lachniet and Seltzer, 2002;
Orvis and Horn, 2000). At La Chonta bog, the late glacial (15,600–
12,900 cal. yr BP) was characterized by an increase of approxi-
mately 4.6°C in MAAT and increases in annual precipitation and
the upper forest line (Hooghiemstra et al., 1992), although warm-
ing was interrupted by an event thought to be correlative with the
Younger Dryas (12,900–11,600 cal. yr BP). Pollen evidence from
La Chonta and La Trinidad bogs indicates that the early- and mid-
dle-Holocene (10,700–5200 cal. yr BP) at mid-elevations in the
Cordillera de Talamanca was characterized by higher effective
moisture and relatively stable thermal conditions (Horn, 2007;
Islebe et al., 1996). Pollen and charcoal stratigraphies developed
for lakes on the glaciated Chirripó massif of Costa Rica indicate
that these uplands were dominated throughout the Holocene by
treeless páramo similar to the present and characterized by peri-
odic fires (Horn, 1993; League and Horn, 2000), especially during
the late-Holocene. Increased burning during the late-Holocene is
interpreted to reflect drier conditions as compared with the mid-
dle-Holocene, when charcoal is sparse in lake sediments. This
interpretation is in keeping with evidence of vegetation changes at
the bog sites (Islebe et al., 1996) and with carbon and hydrogen
isotope records from Morrenas Lake 1 (Lane and Horn, 2013;
Lane et al., 2011a). In the sediments of Lago Chirripó, two distinct
layers of macroscopic charcoal suggest intervals of lower lake
level at about 1100 and 2500 cal. yr BP (Horn, 1993) that may be
associated with regional droughts (Hodell et al., 2000, 2001; Horn,
2007; Lane et al., 2014).
Although much valuable paleoecological and paleoclimato-
logical research has been conducted in Costa Rica, an outstand-
ing opportunity exists to extend our current understanding of
late-Holocene climate change in the southern Pacific region
using sub-fossil chironomid analysis to develop quantitative
reconstructions of late-Holocene thermal conditions. Sub-fossil
chironomid analysis has proven to be valuable in developing
A chironomid-based reconstruction
of late-Holocene climate and environmental
change for southern Pacific Costa Rica
Jiaying Wu,1 David F Porinchu1 and Sally P Horn2
Abstract
A lake sediment profile spanning the last ~3200 years from Laguna Zoncho in the southern Pacific region of Costa Rica was analyzed for sub-fossil
chironomids. Notable shifts in chironomid assemblages occurred during the late-Holocene. A distinct chironomid community, dominated by Tanypodinae
such as Procladius and Labrundinia, appeared after ~550 cal. yr BP (~1400 CE). Prior to this time, the chironomid assemblage was more diverse, with taxa
such as Paratanytarsus, Tanytarsus type N, and Cladotanytarsus important constituents of the chironomid community. A chironomid-based inference model
for mean annual air temperature (MAAT), developed using partial least squares (PLS 2-component), was applied to sub-fossil chironomid assemblages
from Laguna Zoncho to reconstruct late-Holocene thermal variability for the region. The key findings from this study are as follows: (1) chironomid-
inferred MAAT at ~2740–1220 cal. yr BP (790 BCE–730 CE) was 1.2°C higher than the late-Holocene (~3200 cal. yr BP to present in this study) average
of 21.3°C; (2) MAAT at ~470–90 cal. yr BP (1480–1860 CE) was 1.3°C lower than the late-Holocene average, potentially reflecting ‘Little Ice Age’ (LIA)
cooling; and (3) evidence for an extended period of low lake levels between 1220 and 840 cal. yr BP (730–1110 CE) possibly indicated the influence of the
Terminal Classic Drought (TCD) in southern Costa Rica. This study pioneers the use of sub-fossil chironomid remains to develop quantitative estimates
of Holocene thermal variability and environmental change in Central America.
Keywords
chironomid, Costa Rica, Late-Holocene, ‘Little Ice Age’, paleoclimate, temperature, Terminal Classic Drought
Received 12 June 2015; revised manuscript accepted 18 April 2016
1Department of Geography, The University of Georgia, USA
2Department of Geography, The University of Tennessee, USA
Corresponding author:
Jiaying Wu, Department of Geography, The University of Georgia,
Athens, GA 30605, USA.
Email: wu1092@uga.edu
652702HOL0010.1177/0959683616652702The HoloceneWu et al.
research-article2016
Research paper
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2 The Holocene
quantitative temperature reconstructions for the late Quaternary
(Porinchu and MacDonald, 2003; Walker, 2001; Walker and
Cwynar, 2006). Chironomids are sensitive indicators of past tem-
perature and offer great potential to provide independent estimates
of regional climate conditions during intervals of transition
(Axford et al., 2009; Engels et al., 2008; Levesque et al., 1997;
Reinemann et al., 2014). The existence of strong, statistically sig-
nificant correlations between chironomid assemblages and tem-
perature (surface water and air) has facilitated the development of
chironomid-based inference models from the high-, mid-, and low-
latitudes (e.g. Barley et al., 2006; Heiri et al., 2011; Porinchu et al.,
2010; Self et al., 2011; Wu et al., 2015). Application of inference
models to sub-fossil chironomid stratigraphies has enabled
researchers to develop quantitative paleotemperature reconstruc-
tions with high temporal resolution in much of North America,
Eurasia, and increasingly in Asia, Africa, and South America
(Eggermont et al., 2010; Ilyashuk and Ilyashuk, 2007; Ilyashuk
et al., 2011; Massaferro et al., 2009; Reinemann et al., 2009);
however, qualitative and quantitative chironomid-based paleoen-
vironmental reconstructions in Central America remain limited.
Wu et al. (2015) documented the modern relationship between
chironomid distribution and limnological and climatic parame-
ters in Costa Rica. Direct gradient analyses indicated that MAAT
is strongly correlated to the distribution of chironomid taxa in
the 39 Costa Rican lakes included in the study. The robust per-
formance statistics of the two-component partial least squares
(PLS) chironomid-based MAAT inference model ( rjack
2094=.,
RMSEP = 1.73°C) supports using this model to reconstruct Holo-
cene thermal regimes in Costa Rica and potentially elsewhere in
Central America. In this paper, we apply the quantitative chirono-
mid-based inference model developed by Wu et al. (2015) to sub-
fossil chironomid assemblages extracted from a sediment core
recovered from Laguna Zoncho in southern Pacific Costa Rica to
develop a qualitative reconstruction of environmental change and a
quantitative estimate of thermal variability for this region spanning
~3200 years. Comparison of the sub-fossil chironomid-based
reconstructions to existing paleoclimate records from the region
enables us to determine the degree of correspondence between ther-
mal variability and landscape change and to explore how chirono-
mid evidence of late-Holocene climate at Laguna Zoncho links to
other evidence of climate and environmental changes in Costa Rica
and the broader tropical region.
Study site
Laguna Zoncho (8.8121°N, 82.9607°W) is a small (0.75 ha), mid-
elevation (1190 m a.s.l.) lake located on the eastern end of the Fila
Costeña mountain range, near the Panamanian border in southern
Pacific Costa Rica (Figure 1). The site is within the Diquís subre-
gion of the Greater Chiriquí archaeological region, which includes
southern Pacific Costa Rica and western Panama. Laguna Zoncho
is underlain by Tertiary volcaniclastic sedimentary rocks (Ortiz,
2005) and formed at 3200 cal. yr BP by large-scale slumping,
faulting, or both (Clement and Horn, 2001). The lake is on the
grounds of a private nature reserve, Finca Cantaros, which is open
to the public and includes an ornamental garden and slopes refor-
ested over the last two decades following earlier cattle grazing
and coffee cultivation at the site (Horn and Haberyan, 2016).
The lake and surroundings fall within the premontane wet forest
and premontane rain forest transition life zones according to the
Holdridge bioclimatic classification (Holdridge et al., 1971;
Ortiz, 2005). While much of the area has been cleared since the
1950s for pasture and crop cultivation, the Organization for Trop-
ical Studies’ Las Cruces Biological Station located 3 km south of
the lake includes several hundred hectares of remnant and regen-
erating forests (http://ots.ac.cr/index.php?option=com_content&
task=view&id=220&Itemid=422). Following the Köppen classifi-
cation, the region can be characterized as having a humid, equatorial
climate with a dry winter. Data from the Loma Linda meteoro-
logical station located 9 km SSE of Laguna Zoncho (8.8121°N,
82.9607°W; elevation 1180 m a.s.l.) show mean annual tempera-
ture and mean annual precipitation for the period 1988–2007 are
20.5°C and 3359 mm, respectively (http://www.ots.ac.cr/meteoro/
default.php?pestacion=3). During that period, 88% of the annual
precipitation fell during the wet season from May to November.
Methods
We analyzed chironomids in samples from the lacustrine section
(0–2.9 m) of a 5.9-m sediment core that was recovered from
near the center of Laguna Zoncho in March 1997. At the time of
coring, the lake level was low, with a depth of only 2.3 m at the
core site, but repeated visits for limnological sampling (Horn and
Haberyan, 2016) and additional coring (Taylor et al., 2015) have
revealed lake levels up to 2 m higher. In 1997, the watery upper
Figure 1. Locations of paleoenvironmental records referenced in the text, the 51 lakes in the Costa Rican chironomid calibration set and
Laguna Zoncho (Wu et al., 2015).
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Wu et al. 3
sediments (0–1.13 m) were sampled using a plastic tube fitted
with a rubber piston and extruded, sub-sectioned, and bagged in
the field at 2 cm intervals, and deeper sediments were recovered
using a Colinvaux-Vohnout locking piston corer (Colinvaux et al.,
1999). Core sections were returned to the Laboratory of Paleoen-
vironmental Research at the University of Tennessee still encased
in the original 1-m-long aluminum coring tubes. In the lab, the
tubes were opened on a modified router and the core sections
were sliced longitudinally for sampling, initially for pollen and
microscopic charcoal and loss-on-ignition (Clement and Horn,
2001) and later for stable carbon isotopes (Lane et al., 2004), dia-
toms (Haberyan and Horn, 2005), phosphorous (Filippelli et al.,
2010), and finally chironomids (this study).
The 1997 Laguna Zoncho sediment core contains a lower sec-
tion composed mainly of mineral regolith with poor pollen preser-
vation and a pollen-rich lacustrine section (0–290 cm) (Clement
and Horn, 2001). Chronological control for the lacustrine profile is
based on three AMS dates on small wood fragments or samples
containing a mixture of charcoal, seeds, and leaf fragments and a
well-dated tephra layer from the volcano Barú in western Panama
(Table 1 and see Clements and Horn, 2001 for additional details).
For this paper, the radiocarbon dates were converted to calendar
ages using CALIB 7.1 and the data set of Reimer et al. (2013). Ages
were estimated using a linear age model and the weighted means of
the probability distributions of the calibrated ages. Based on the age
model, the lacustrine sediment in the core accumulated slowly
(0.034 cm yr−1) between ~3100 and 2100 cal. yr BP (284–249 cm)
and more quickly (0.084 cm yr−1) between 2100 and 560 cal. yr BP
(247–120 cm) and (0.194 cm yr−1) from 540 cal. yr BP to the pres-
ent. The highest sedimentation rate (1.013 cm yr−1), which occurred
between 560 and 540 cal. yr BP (120–99.75 cm), was likely due to
the eruption of Volcano Barú (located 35 km east of the study site).
Chironomid samples were analyzed at ~10 cm resolution in the
upper 290 cm of the core. A minimum of 50 head capsules were
analyzed for each interval, with the exception of samples at 272 and
287 cm for which 46 and 44 head capsules were enumerated, respec-
tively. The chironomid analysis was conducted following standard
procedures outlined in Walker (2001). The sediment was treated
with 5% KOH solution to facilitate the break-up of colloidal matter.
A known volume of sediment (3–8 mL) was placed in a beaker with
50 mL of 5% KOH and heated at 50°C for approximately 30 min.
The deflocculated sediment was washed through a 95-µm mesh and
rinsed using distilled water. The material retained on the mesh was
backwashed into a beaker. A dissection microscope at 50× magnifi-
cation and a Bogorov plankton counting tray were used to separate
the chironomid head capsules from the sediment matrix. The chi-
ronomid head capsules were permanently mounted on slides in
Entellan© for identification. Taxonomic identification was con-
ducted at 400× magnification, typically to genus, relying primarily
on larval keys for Florida and North and South Carolina (Epler,
1995, 2001), with Brooks et al. (2007), Eggermont et al. (2008), and
Cranston (2010) providing additional diagnostic information.
The chironomid percentage diagram, plotted using C2 (Juggins,
2003), was based on the relative abundance of all identifiable
chironomid remains. Taxon richness was determined by counting
the number of distinct taxonomic groupings in each sample. We
numerically zoned the chironomid percentage diagram based on
optimal sum-of-squares partitioning, using the R-based rioja
package (Juggins, 2014). The degree of turnover in the sub-fossil
chironomid assemblages can be indicated by the shift of the
assemblage data-based detrended correspondence analysis (DCA)
curve. A chironomid-based inference model (2-component PLS)
for MAAT (Wu et al. 2015) was applied to this data set. This infer-
ence model is based on 45 chironomid taxa in surface samples
from 39 lakes that span seven ecosystem regions and elevations
from 10 to 3520 m a.s.l. and provides robust performance statis-
tics ( rjack
2094=., RMSEP = 1.73°C). Application of the inference
model to the Laguna Zoncho chironomid stratigraphy provides a
means to develop a quantitative temperature reconstruction span-
ning the late-Holocene for the region. Reconstructions are based
on sub-fossil chironomid assemblages that have >95% of the sub-
fossil taxa present in the calibration set, which are considered
reliable (Birks, 1998). The reliability of the quantitative chirono-
mid-based reconstruction was also evaluated by determining (1)
the dissimilarity between each Zoncho chironomid sample and its
closest modern analog using a modern analog technique (MAT-
ech) approach based on a minimum dissimilarity chord distance
and (2) square residual goodness-of-fit (GOF) of each Zoncho
chironomid assemblage to the first ordination axis in a canonical
correspondence analysis (CCA) constrained solely by MAAT.
The second and fifth percentiles of the distribution of dissimilari-
ties, based on the calibration set samples incorporated in Wu et al.
(2015), were used to define the cut-off for ‘no close’ and ‘no
good’ analogs, respectively (Birks et al., 1990; Engels et al., 2008;
Heiri et al., 2003). Samples with a squared residual distance
greater than the 90th, 95th and 99th percentiles of the residual
distances of the calibration set samples were identified as having
a ‘poor fit’ or ‘extremely poor fit’ with temperature, respectively
(Birks et al., 1990). The trajectory of change in the sub-fossil
chironomid assemblages in Laguna Zoncho was determined
using correspondence analysis (CA), with the sub-fossil chirono-
mid assemblages plotted passively against the modern calibration
set samples (Wu et al. 2015).
Results
Sub-fossil chironomid head capsules were well preserved in the
lacustrine sediments of the Zoncho core except for the interval
between 176 and 144 cm. A total of 24 chironomid taxa were
identified, with 23 taxa present (>95%) in the modern calibration
set (Wu et al., 2015). Tanytarsus type W (Figure 3) is the only
taxon, present in the sub-fossil chironomid assemblages from
Laguna Zoncho, not found in the modern calibration set. Chirono-
mid richness varied between 4 and 18 with the highest taxon rich-
ness occurring at approximately 1400 cal. yr BP (550 CE).
Chironomus, Labrundinia, and Procladius were the most abun-
dant chironomid taxa. The chironomid stratigraphy is divided into
four zones (Figure 2).
Table 1. AMS
14C dates available for the Laguna Zoncho lake sediment core. Analyses were performed by Beta Analytic Laboratory on small
wood fragments (W) or mixtures of charcoal, seeds and leaf fragments (M). Radiocarbon ages were calibrated using CALIB 7.02 and the data
set of Reimer et al. (2013).
Lab no. Depth (cm) Material dated Uncalibrated
14C age (14C yr BP)
± Median (cal. yr BP) Calibrated age ± 1σ
(cal. yr BP)
Calibrated age ± 2σ
(cal. yr BP)
β-122556 118–122 M 540 50 558 518–558 505–569
602–629 582–650
β-122555 248–250 M 2110 50 2086 2004–2028 1949–2180
2035–2145 2241–2303
β-115186 283–284.5 W 2940 50 3095 3005–3015 2948–3238
3021–3065 3313–3316
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4 The Holocene
Figure 2. Sub-fossil chironomid stratigraphy for Laguna Zoncho. Taxa on the left side of diagram are arranged according to MAAT optima as determined by Wu et al. (2015), with Procladius having the lowest MAAT
optima and Cladotanytarsus having the highest MAAT optima. The taxa observed in the ‘Low Chironomid Head Capsule Recovery’ interval are symbolized by asterisks. The locations of the radiocarbon dates are
indicated by red bars on the right margin of the figure, labeled with uncalibrated 14C age and calibrated age range (cal. yr BP ±1σ) in brackets (see Clements and Horn, 2001 for additional details). The location of
tephra layer is marked by brown bar.
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Wu et al. 5
Zone LZ-1 (~3220–2740 cal. year BP or 1270–790
BCE; 288–271.5 cm)
This zone is dominated by two taxa: Chironomus and Labrun-
dinia. Lesser amount of taxa belonging to the sub-tribe Tanytar-
sini – Cladotanytarsus, Paratanytarsus, and Tanytarsus type N
– are also present. Taxon richness is relatively low with only nine
taxa present in this zone (Figure 2). Chironomus is most abun-
dant in mid-elevation lakes (1000–2000 m a.s.l.) today in Costa
Rica (Wu et al., 2015). The other taxa present in LZ-1, such as
Dicrotendipes, Labrundinia, Beardius type A, and Paratanytar-
sus, are most abundant and common in warm, low-elevation
lakes (Wu et al., 2015). The DCA suggests that limited faunal
turnover occurred during this interval (Figure 4a).
Zone LZ-2 (~2740–1220 cal. yr BP or 790
BCE–730 CE; 271.5–176 cm)
This zone is characterized by a more diverse chironomid com-
munity with taxon richness reaching a profile maximum of 18 at
approximately 1400 cal. yr BP (550 CE). Thermophilous taxa are
particularly abundant. The assemblage in LZ-2 is dominated by
Chironomus, which co-occurs with less amounts of Labrundinia
(Figure 2). Taxa that are associated with warm, low-elevation
lakes, such as Paratanytarsus, Polypedilum type N, Micropsec-
tra, Labrundinia, Cladotanytarsus and Tanytarsus type N, are
frequent in this zone (Wu et al., 2015). Taxa such as Parachi-
ronomus, Procladius, Cricotopus, and Corynoneura are also
present in this zone, but at low relative abundance. The increase
in the relative abundance of Procladius along with the decrease
in the relative abundance of Labrundinia and Tanytarsus N type
at the very top of LZ-2 is inferred to indicate the onset of a
decrease in air temperature at the termination of this zone. Today,
Procladius is most abundant in cold, high-elevation lakes (>3000
m a.s.l.) in Costa Rica, whereas Labrundinia and Tanytarsus type
N are most abundant in mid- and low-elevation lakes (Wu et al.
2015). A previously undescribed chironomid taxon is also
observed in LZ-2: Tanytarsus type W is characterized by sharply
sloping first lateral teeth, which serves as the key diagnostic fea-
ture (Figure 3). Additional diagnostic features include the pres-
ence of a single narrow median tooth with one pair of tiny
supplementary teeth, a mandible with two inner teeth, and an
antennal pedestal with a round and pointed spur. Cladopelma, a
chironomid absent today from lakes located higher in elevation
than Laguna Zoncho, appears for the first time in the record at
the top of LZ-2, immediately prior to a distinctly different inter-
val in the Zoncho chironomid record. The DCA indicates that the
chironomid community experienced limited faunal turnover until
~1220 cal. yr BP (Figure 4a). Low chironomid interval (~1220–
840 cal. yr BP or 730–1110 CE; 176–144 cm) between zones
LZ-2 and LZ-3 in the Zoncho profile is a section of sediment
with only sparse chironomid head capsules indicating an interval
of low chironomid abundance. The number of sub-fossil chiron-
omid head capsules recovered in samples from this interval
failed to meet the screening criteria required for further statistical
analysis, precluding the use of these samples in deriving quanti-
tative estimates of MAAT. However, the taxa that are present
between 1220 cal. yr BP and 840 cal. yr BP, such as Cladotany-
tarsus, Tanytarsus type P, and Rheotanytarsus, typically occur in
the littoral of warm, productive, low-elevation lakes in Costa
Rica today (Wu et al., 2015).
Rapid compositional change is often a signal of notable fluc-
tuation in limnological and/or environmental conditions. The
DCA of the chironomid percentage data indicates that the chi-
ronomid community at Laguna Zoncho experienced one major
interval of rapid turnover during the late-Holocene (Figure 4a).
This interval occurred during the transition between LZ-2 and
LZ-3. The chironomid community in LZ-2 was dominated by
taxa associated with warm, low-elevation lakes such as Paratany-
tarsus, Polypedilum type N, Micropsectra, Labrundinia, Cladota-
nytarsus, and Tanytarsus type N. Following LZ-2, the relative
abundance of Micropsectra and Cladotanytarsus declined and the
relative abundance of taxa commonly associated with aquatic
macrophytes and wetland habitats such as Stenochironomus and
Beardius reissi type increased in the LZ-3 (Figure 2). It is impor-
tant to note that the chironomid community during the transition
between LZ-2 and LZ-3 was characterized by a ~400-year-long
interval of low lake productivity with sub-fossil chironomid
remains extremely scarce (average <1.6 heads/mL) in the sedi-
ments between ~1220 and 840 cal. yr BP (~730–1110 CE).
Zone LZ-3 (~840–550 cal. yr BP or 1110–1400 CE;
144–108 cm)
Taxon richness decreases to an average value of 12 in Zone LZ-3
(Figure 2). The decrease in taxon richness is largely driven by a
reduction in the relative abundance of taxa typically associated
with warm, low-elevation lakes (e.g. Micropsectra and Cladota-
nytarsus), and the local extirpation of multiple Tanytarsus taxa,
for example, Tanytasus type P, Tanytarsus type L, and Tanytarsus
type N; Tanytarsus type P, and Tanytarsus type N, is most com-
monly associated with low-elevation lakes (Wu et al., 2015).
Zone LZ-3 is also characterized by a notable increase in Beardius
reissi type and the appearance of Stenochironomus taxa that are
often associated with submerged macrophytes and littoral vegeta-
tion, respectively (Cranston, 2010). The modern calibration set
documents that Stenochironomus and Beardius reissi type are
most abundant in and commonly associated with warm, low-ele-
vation lakes in Costa Rica (Wu et al., 2015). Chironomus (50%)
continues to dominate this zone. The DCA suggests that a notable
amount of faunal turnover characterized the chironomid commu-
nity during the transition through the interval with extremely low
chironomid head capsule recovery (Figure 4a).
Zone LZ-4 (~550 to −47 cal. yr BP or ~1400–1997
CE; 108–0 cm)
The relative abundance of Chironomus, which continues to be
high in this zone, fluctuates between 20% and 60% of identifi-
able chironomid remains (Figure 2). This zone is characterized
by a noticeable increase in Cricotopus, a taxon commonly asso-
ciated with high-elevation lakes (Wu et al., 2015). Warm water
taxa, such as Labrundinia, Chironomus, Dicrotendipes, and
Figure 3. Photomicrograph of Tanytarsus type W, a previously
undescribed chironomid taxon in Costa Rica. The key diagnostic
features of this taxon are the sloping angle of the outer margin of
the first pair lateral teeth and the presence of an antennal pedestal
with a round and pointed spur (arrow).
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6 The Holocene
Figure 4. (a) Detrended correspondence analysis (DCA) first axis score based on the sub-fossil chironomid assemblages in the Laguna
Zoncho core. (b) Chironomid-inferred MAAT reconstruction for Laguna Zoncho site (solid line with gray error bars). (c) Square chord
dissimilarity distance (sq. chord. d) to the nearest modern analogs (MATech) in Wu et al. (2015). (d) The goodness-of-fit (GOF) of the fossil
samples to a canonical correspondence analysis (CCA) constrained solely to temperature. Horizontal lines in the GOF analysis indicate the
90th (dash line), 95th (dash-dot-dot line), and 99th (dash-dot line) percentiles in residual distances of the modern samples to the first axis
in a constrained CCA and are defined as ‘poor fit’, ‘very poor fit’, and ‘extremely poor fit’, respectively (Birks et al., 1990). Vertical brown bar
represents the Barú tephra layer observed between 99 and 100.5 cm and dated to ~538 cal. yr BP (Taylor et al., 2013a).
Ablabesmyia, are also observed in this zone, indicating the exis-
tence of a diverse chironomid community during this interval
(Wu et al., 2015). The local extirpation of Stenochironomus and
Rheotanytarsus and a notable reduction in the relative abundance
of Beardius reissi type occurs in LZ-4. Stenochironomus and
Beardius reissi type are often associated with submerged aquatic
macrophytes and wetland vegetation (Jacobsen and Perry, 2000;
Pinder and Reiss, 1983), whereas Rheotanytarsus is commonly
found in inflowing water (Pinder and Reiss, 1983). Taxon rich-
ness increases to 18 in LZ-4; however, the recently deposited
sediment (~1950–1997 CE; 10–0 cm) is characterized by initially
decreased and then increased taxon richness (Figure 2). The DCA
indicates relatively limited faunal turnover characterizes LZ-4,
although sample-to-sample fluctuations are observed (Figure 4a).
The chironomid-based quantitative MAAT reconstruction for
Laguna Zoncho is presented in Figure 4b. The temporal resolu-
tion of the sub-fossil chironomid assemblage analysis and the
associated quantitative temperature reconstruction vary from sub-
centennial scale (550 cal. yr BP–present) to centennial scale
(2300−550 cal. yr BP) and to multi-centennial scale (3200−2300
cal. yr BP). Reconstructed MAAT varied from a minimum of
18.8°C at 180 cal. yr BP (~1770 CE) to a maximum of 24.1°C at
around 1770 cal. yr BP (~180 CE), with 21.3°C as the average
MAAT for late-Holocene (~3200 cal. yr BP to the present in this
study). The chironomid-inferred MAAT reconstruction suggests
that Laguna Zoncho experienced warmer interval between ~2740
and 1220 cal. yr BP (Figure 4b, LZ-2, highlighted in light red) and
colder interval from 550 to −47 cal. yr BP (~1400–1997 CE,
Figure 4b, LZ-4, highlighted in light blue). Specifically, the
reconstructed MAAT from ~2740 to ~1220 cal. yr BP (790 BCE–
730 CE) was averagely 22.5°C, that is, 1.2°C higher than the late-
Holocene average, and from 550 to −47 cal. yr BP was 20.4°C,
that is, 0.9°C lower than the late-Holocene average (Figure 4b).
Sample-specific error estimates ranged between 2.0°C and 2.3°C.
Analyses of the reliability of the quantitative chironomid-based
temperature reconstruction using modern analog technique
(MATech) and GOF approaches gave mixed results (Figure 4c
and d). The GOF analysis indicates that the sample scores fluctu-
ate below the 90th percentile cut-levels and therefore have a good
fit to temperature and can be considered reliable. However, the
MATech analysis indicates that close analogs in the modern cali-
bration set (Wu et al., 2015) are limited to the chironomid assem-
blages recovered from the uppermost (~1950 CE) and lowermost
(~3200 cal. yr BP) portions of the profile.
Discussion
Paleoecological and paleolimnological analyses of the sediment
of Laguna Zoncho have provided important insight into late-
Holocene environmental change for the southern Pacific region
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Wu et al. 7
of Costa Rica (Clement and Horn, 2001; Filippelli et al., 2010;
Haberyan and Horn, 2005; Lane et al., 2004; Taylor et al.,
2013b). These studies documented the onset and magnitude of
prehistoric agriculture, the timing of subsequent forest recovery,
and the changes in catchment conditions and lake productivity
that result from agricultural and associated anthropogenic activi-
ties. Aspects of these records also suggested possible climate-
driven shifts in environment and human activities (Taylor et al.,
2013a). Deciphering the paleoecological information contained
within the sub-fossil chironomid assemblages extracted from
Laguna Zoncho provides an additional means to characterize
late-Holocene environmental and climate change in the neo-
tropics and application of a quantitative inference model for
MAAT (Wu et al., 2015) to the Laguna Zoncho chironomid stra-
tigraphy may help elucidate the role of temperature in driving the
inferred changes. The resolution of the original chronology
(Clement and Horn, 2001), based on three radiocarbon dates, has
been increased with the inclusion of an additional date, which is
associated with the presence of a tephra layer (dated at ~538 cal.
yr BP) from the eruption of volcano Barú located at western
Panama. It is important to recognize that chronological control
for the Laguna Zoncho sediment sequence is most robust at cen-
tennial-scale resolution.
The trajectory of change in the sub-fossil chironomid assem-
blages in Laguna Zoncho was determined by passively plotting the
sub-fossil chironomid assemblages against the modern calibration
set samples using correspondence analysis (CA) (Figure 5). The
black crosses represent the movement of the chironomid assem-
blages relative to the modern chironomid community (classified
by elevation) at a multi-centennial time scale during the late-
Holocene (Wu et al., 2015). The modern chironomid assemblage
recovered from Laguna Zoncho is most similar to assemblages
found today in mid-elevation lakes with moderate MAAT. The
ordination diagram, which separates cold, high-elevation lakes
from warmer, low-elevation lakes along CA axis 1, suggests that
the sub-fossil assemblages found throughout the Laguna Zoncho
record are most similar to assemblages found in warm, produc-
tive, low, and mid-elevation lakes today and that the changes in
MAAT that occurred at Laguna Zoncho during the past 3200 years
were relatively muted. However, the CA indicates that notable
changes in the trajectory of the sub-fossil chironomid assem-
blages with respect to temperature occurred between 1300 and
800 cal. yr BP and between 600 and 400 cal. yr BP. The move-
ment of the sub-fossil chironomid assemblage at 1300 cal. yr BP
toward the assemblages that characterize the lowest elevation
lakes today is inferred to reflect an increase in MAAT during this
interval. The sub-fossil chironomid assemblages, which move
toward the assemblages that characterize high-elevation sites
today between 400 and 200 cal. yr BP, are inferred to reflect a
decrease in MAAT during this interval.
It is important to recognize that changes in the lake catchment,
related to agricultural activity and vegetation composition, may
have influenced the chironomid community at Laguna Zoncho. A
DCA of the pollen data, which captures the magnitude and timing
vegetation change, was used to assess the correspondence
between the variations in the chironomid assemblages and agri-
cultural activity (Figure 6). The results of the DCA suggest that
there is limited correspondence between faunal turnover and the
changes in catchment vegetation during the late-Holocene.
Although not definitive, the DCA does provide additional support
that fluctuations in temperature are the main driver of chironomid
community composition rather than agriculture-related nutrient
enrichment.
The chironomid assemblages present at the base of the Laguna
Zoncho profile, which are relatively depauperate, are dominated
by taxa typically associated with warm, productive lakes found at
middle and low elevations today (Wu et al., 2015). The presence
of Labrundinia together with lower amounts of Beardius type A
and Dicrotendipes in LZ-1 is suggestive of the presence of aquatic
macrophytes. It is important to note that only three chironomid
assemblages are available in LZ-1; therefore, the observed varia-
tions in the sub-fossil chironomid assemblages may not capture
full range of environmental change during this interval. Zone
LZ-1 corresponds to pollen zone 5 in the pollen, microscopic
charcoal and stable carbon isotope records developed by Clement
and Horn (2001) and Lane et al. (2004). The presence of maize
pollen together with high percentages of Poaceae pollen, low tree
pollen percentages, enriched sedimentary 13C values, and high
charcoal concentrations and charcoal:pollen ratios indicates an
early interval of forest clearance and maize agriculture at Laguna
Zoncho that began no later than the time of lake formation. From
the chironomids, we can infer relatively warm conditions from
~3220–2740 cal. yr BP (Figure 6) during this initial agricultural
period.
Figure 5. Time-trend CA biplot comparing sub-fossil chironomid assemblages from Laguna Zoncho with the modern chironomid assemblages in
the 51-lake calibration set from Costa Rica (Wu et al. 2015). The calibration data set lakes are classified according to elevation (m a.s.l.). Numeric
values near the black crosses in the diagram represent Zoncho sub-fossil assemblages at ~0.2 kyr interval from 2.0 k cal. yr BP to modern time and
at 0.3–0.5 kyr interval from 3.2 to 2.0 k cal. yr BP (coarser time interval is due to lower temporal resolution at the base of the core).
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8 The Holocene
LZ-2 zone (~2740–1220 cal. yr BP or 790 BCE–730 CE;
271.5–176 cm) is characterized by an abrupt increase in chirono-
mid richness. The increase in chironomid community diversity is
driven by thermophilous taxa such as Dicrotendipes, Polypedilum
type N, Micropsectra, and Tanytarsus type N. The presence of
Polypedilum and Microtendipes, taxa inhabiting littoral and sub-
littoral sediments, together with the presence of thermophilous
taxa and taxa associated with submerged vegetation indicate that
this interval was characterized by higher temperatures, increased
lake productivity, and the expansion of littoral habitat. This inter-
pretation is supported by the quantitative chironomid-based
reconstruction, which indicates the existence of elevated MAAT
from ~2740 to 1220 cal. yr BP. The low sampling resolution in the
basal portion of the LZ-2 precludes our ability to make definitive
conclusions about the nature of the environmental change from
2740 to 1220 cal. yr BP. Changes in the diatom flora reveal that
lower lake depth and more acidic lake water characterized Laguna
Zoncho during this interval (Haberyan and Horn, 2005). The pol-
len record suggests that the catchment surrounding Laguna Zon-
cho experienced forest regrowth during the lower portion of LZ-2
(corresponding to pollen zone 4), followed by increased maize
cultivation, forest clearance, and fire beginning ~1700 cal. yr BP
(pollen zone 3) (Clement and Horn, 2001; Lane et al., 2004; Fig-
ure 6). The lower δ13C values in LZ-2 likely reflect increased C3
vegetation in the catchment resulting from reforestation (Lane
et al., 2004).
The extremely low head capsule recovery that characterizes
the chironomid stratigraphy from 176 to 144 cm (~1220–840 cal.
yr BP or 730–1110 CE) likely reflects marked changes in limno-
logical conditions at Laguna Zoncho. The near absence of chi-
ronomids during this interval could be explained by anthropogenic
activities, climate change, or some combination of the influences
of both climate change and anthropogenic disturbance. The pol-
len, charcoal, and stable carbon isotope records from the 1997
Zoncho core show evidence of increased intensity of maize culti-
vation during part of the interval of low chironomid productivity;
this evidence includes the most enriched 13C values, highest
charcoal:pollen ratios, and the highest percentages of Asteraceae
and Amaranthaceae pollen, interpreted as agricultural weeds, in
the profile. Increased maize cultivation and burning may have
led to increased erosion and sediment input to the lake, which
in turn may have altered habitat and food availability for the
Figure 6. Summary diagram depicting pollen DCA scores, pollen abundance for select taxa, charcoal:pollen ratio, δ13C values, the abundance
of diatom taxon Aulacoseira sp. 3, and the chironomid-inferred MAAT reconstruction along with DCA first axis score (Clement and Horn, 2001;
Haberyan and Horn, 2005; Lane etal., 2004; modified by Wu). Gray dotted lines indicate the chironomid zones. The gray dashed lines indicate the
zonation based on diatom, pollen, and charcoal analyses, used in Clement and Horn (2001), Haberyan and Horn (2005), and Lane etal. (2004).
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Wu et al. 9
chironomids. This inference is supported by the presence of
Rheotanytarsus, albeit at low numbers, during this time. Increased
erosion in the catchment would have facilitated the construction
of larval cases consisting of coarser sediment such as sand by
Rheotanytarsus (Pinder and Reiss, 1983).
Alternatively, it is possible that a decrease in effective mois-
ture between ~1220 and 840 cal. yr BP lowered lake level to the
point that only a limited pool of standing water remained at the
core site, which would have decreased the chironomid diversity
and abundance. This interval of low head capsule recovery may
be linked to the impact of the Terminal Classic Drought (TCD),
which consisted of a series of multi-decadal droughts that
affected Central and South America and the Caribbean between
1200 and 850 cal. yr BP (750 and 1100 CE) (Lane et al., 2014)
and may have contributed to the disintegration of the classic
Maya civilization (Haug et al., 2003; Hodell et al., 2005a) as well
as to cultural and environmental change throughout the circum-
Caribbean region (Lane et al., 2009, 2011b, 2014). That the inter-
val of low chironomid productivity at Laguna Zoncho may have
been associated with a climate-mediated lowering of lake level is
supported by high-resolution δ13C records (Taylor et al., 2013a,
2015) of five sediment cores recovered from Laguna Zoncho in
2007. These analyses, carried out at much higher resolution than
our chironomid, pollen, or prior isotope analyses, revealed two
periods of basin-wide depletion of 13C inputs interpreted to indi-
cate agricultural decline associated with droughts from 1150 to
970 cal. yr BP (800–980 CE) and 860 to 640 cal. yr BP (1090–
1310 CE). Finally, additional evidence of dry climate and low-
ered lake level in Costa Rica at this time is provided by lenses of
macroscopic charcoals deposited at ~1100 cal. yr BP (850 CE) at
Laguna Gamboa located 4 km southwest of Laguna Zoncho
(Horn, 2007 and unpublished) and at Lago Chirripó in the Cor-
dillera de Talamanca (Horn, 1993). Given the additional evi-
dence of droughts at Laguna Zoncho and other sites in Costa
Rica and the wider circum-Caribbean at this time, we find it rea-
sonable to interpret the interval of low chironomid productivity
in our record to reflect at least in part the influence of the TCD
on hydroclimatology in Costa Rica; anthropogenic activities in
the watershed may have also contributed.
In LZ-3 (~840–550 cal. yr BP or 1110–1400 CE), the diver-
sity of the chironomid assemblage is reduced relative to LZ-2.
The decrease in taxon richness is largely driven by the local
extirpation of a number of taxa typically associated with warm,
low-elevation lakes, for example, Tanytarsus type P and Tanytar-
sus type N (Wu et al., 2015). The notable increase in Beardius
and the appearance of Stenochironomus likely reflect an expan-
sion of aquatic macrophytes cover, the continued existence of
relatively low lake levels, and elevated lake productivity (Crans-
ton, 2010). The diatom flora in this zone is also very diverse,
with an increase in Eunotia minor and the absence of Aulaco-
seira sp. 3, a planktonic diatom taxon; these aspects of the
diatom assemblage together with relatively high values for
the shallow water ratios of phytoliths:diatoms and sponge
spicules:diatoms support the chironomid-based inference of
lower lake level during this interval (Haberyan and Horn, 2005).
Clement and Horn (2001) interpreted this time period in the
upper part of pollen zone 3 as one of continued maize cultivation
and agricultural burning, as shown by pollen and microscopic
charcoal indices, but Taylor et al. (2013a, 2015) found evidence
of declining 13C ratios between 840 and 560 cal. yr BP in five
cores collected from Laguna Zoncho in 2007 and interpreted
these isotope shifts to indicate a period of population and agricul-
tural decline in the watershed prior to the Spanish Conquest.
Earlier analyses of stable carbon isotopes in the 1997 profile
had also shown a drop in 13C ratios within pollen zone 3, but
the possible significance of this drop as an indicator of agricul-
tural decline was not recognized. The interpretation of this
agricultural decline as resulting from drought is consistent with
the interpretation from the chironomid stratigraphy of low lake
level.
Zone LZ-4 (~550– (−47) cal. yr BP or ~1400–1997 CE) is char-
acterized by an increase in the relative abundance of taxa associated
with relatively cold, high-elevation lakes such as Procladius and a
decrease in taxa typically found in warm, productive low-elevation
lakes such as Paratanytarsus, Polypedilum type N, and Cladotany-
tarsus (Figure 2). Today in Costa Rica, Procladius dominates high-
elevation, glacial lakes (e.g. Lake Morrenas 0, 1, 2, 3, 4; Lago
Chirripó; and Lago Ditkebi) and Cricotopus is mainly found in
deep glacial lakes (Wu et al., 2015). The reduction of many littoral
taxa, together with the increase in Procladius and Cricotopus,
appears to indicate the onset of colder, less productive conditions at
Laguna Zoncho. The shift in the chironomid assemblages in LZ-4
could reflect an increase in lake depth; however, based on auteco-
logical information and the lack of a statistically significant rela-
tionship between chironomid distributions and depth in the modern
calibration set (Wu et al., 2015), we believe that the LZ-4 assem-
blages are reflective of a lowering of air temperature. The average
chironomid-inferred MAAT declines to 20.3°C during LZ-4 period,
which is 1.0°C lower than the late-Holocene chironomid-inferred
MAAT average of 21.3°C. Additionally, notable excursions in
MAAT occur at 1410 CE (18.9°C) and 1770 CE (18.8°C) (Figures
4 and 6). The existence of depressed MAAT (1.3°C lower than the
3200-year average) between 1480 CE and 1860 CE (470–90 cal. yr
BP) may reflect the manifestation of the ‘Little Ice Age’ (LIA) in
southern Costa Rica. Evidence of low-latitude cooling and drought
during the ‘LIA’ has been documented at several sites in the cir-
cum-Caribbean (Böhm et al., 2002; Glynn et al., 1983; Hodell
et al., 2005b; Lane et al., 2009, 2011b; Taylor et al., 2013a, 2015;
Watanabe et al., 2001; Winter et al., 2000) and from the tropical
Andes, where ice cores suggest marked cooling between 1400 CE
and 1900 CE (Thompson et al., 2006). Lake and marine records
recovered from study sites in the southern hemisphere also indi-
cate the occurrence of ‘LIA’ cooling (Polissar et al., 2006; Rabatel
et al., 2008). High atmospheric aerosol concentrations, resulting
from several large volcanic eruptions and sea-ice/ocean feedbacks
(Crowley and Lowery, 2000; Mann et al., 2009; Miller et al.,
2012), have been implicated as the drivers responsible for the
‘LIA’. On the basis of chironomid assemblages, we suggest that
the ‘LIA’ in southern Costa Rica was characterized by ~1.3°C
depression in MAAT relative to the late-Holocene MAAT average
(Figure 6).
The diatom, pollen, δ13C, and charcoal records document
that Laguna Zoncho and the surrounding catchment were char-
acterized by increasing lake levels, regeneration of forest,
reduced burning, and a marked decrease in human activities
beginning at ~1450 CE (Clement and Horn, 2001; Haberyan and
Horn, 2005; Lane et al., 2004) or earlier (Taylor et al., 2013b,
2015). The Spanish Conquest greatly reduced indigenous popu-
lations in Costa Rica, and changes in terrestrial proxies in the
Zoncho record may have been driven largely by these popula-
tion reductions. However, agricultural declines prior to the Con-
quest were likely associated with drought, and colder and drier
conditions during the ‘LIA’ would have affected people and
environments in southern Pacific Costa Rica as in other areas of
circum-Caribbean (e.g., Hodell et al., 2005b; Lane et al., 2011b).
Based on a compilation of charcoal records from across the
Americas, Power et al. (2013) concluded that cooling during the
‘LIA’ played a greater role in reducing fire occurrence after
1500 CE on a global to continental scale than did the post-con-
tact collapse of indigenous populations. We recognize and in no
way mean to discount the large and devastating cultural and
environmental changes associated with the Spanish Conquest in
Costa Rica. However, based on the indications of ‘LIA’ cooling
in our chironomid record, and following Lane et al. (2011b), we
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10 The Holocene
suggest that climate change coincident with the arrival of Euro-
peans deserves more attention as a factor that affected local and
regional ecosystems, environments, and potentially human
activities in Costa Rica.
Conclusion
This study pioneers the use of sub-fossil chironomid analysis in
developing qualitative inferences of environmental change and
quantitative estimates of thermal variability in Central America.
Distinct shifts in chironomid community composition were
observed during the late-Holocene. The notable shift in chirono-
mid community composition that occurred at ~550 cal. yr BP
(~1400 CE), characterized by an increase in the relative abun-
dance of Procladius and a decrease in Polypedilum, Cladotany-
tarsus, and Paratanytarsus, suggesting that Laguna Zoncho was
likely characterized by cooler, less productive conditions
between ~1480 CE and 1860 CE. The chironomid-inferred
MAAT during this interval shows 1.3°C below the ~3200-year
MAAT average, providing evidence for the regional manifesta-
tion of the ‘LIA’ in Costa Rica. The near absence of sub-fossil
chironomid remains between ~1220 and 840 cal. yr BP (~730–
1110 CE), which likely reflects changes in habitat availability
due to lowered lake levels, may be associated with decreased
effective moisture during the TCD. Anthropogenic activities in
the catchment such as forest clearance and cultivation of maize
during the TCD may have also contributed to low chironomid
abundance. The development of temperature reconstructions for
Costa Rica will benefit from additional analyses of chironomid
stratigraphies, especially at sites of more limited human impact
during the Holocene.
Acknowledgements
We thank the late Luis Diego Gómez for encouraging Sally P
Horn to begin work at Laguna Zoncho, former land owner Mi-
chelle Cloud and current landowners Gail Hewson Hull and
Harry Hull for permission for repeated sampling at the lake, Lisa
Kennedy and Brandon League for assistance with sediment cor-
ing, and Professor Maureen Sánchez of the University of Costa
Rica and the Organization for Tropical Studies for logistical sup-
port. We also thank Scott Reinemann, Xuebin Wei, and Yu Luo
for providing comments on the manuscript.
Funding
A University of Georgia Provost’s Summer Research Award and
a University of Georgia SEC Visiting Faculty Travel Grant to
David F Porinchu provided support for this research. Sediment
coring at Laguna Zoncho and radiocarbon dates were funded by a
grant to Sally P Horn and RL Sanford, Jr, from The A.W. Mellon
Foundation, and core sampling for chironomids was supported by
the Initiative for Quaternary Paleoclimate Research at the Univer-
sity of Tennessee.
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