Water Air Soil Pollut (2012) 223:1125–1136
DOI 10.1007/s11270-011-0930-6
Atmospheric Deposition of Nitrogen to a Caribbean Coastal
Zone (Cayo Coco, Cuba): Temporal Trends and Relative
Importance as a Nitrogen Source
Roberto González-De Zayas &
Martín Merino-Ibarra & Felipe Matos-Pupo &
Martín F. Soto-Jiménez
Received: 22 March 2011 / Accepted: 9 August 2011 / Published online: 3 September 2011
# Springer Science+Business Media B.V. 2011
Abstract Nitrogen (N) deposition to the ocean is
thought to be increasing worldwide, but the amount of
coastal and open ocean measurements is very limited. In
this paper, we assess N deposition in the coastal zone of
Cayo Coco, in central Cuba, during a multi-annual period
(2005–2007). Wet and dry N depositions were estimated
R. González-De Zayas
Posgrado en Ciencias del Mar y Limnología.
Instituto de Ciencias del Mar y Limnología, UNAM,
Ciudad Universitaria, Coyoacán,
Mexico DF, Mexico 04510
e-mail: cuba_robe@yahoo.com
R. González-De Zayas
e-mail: roberto@ciec.fica.inf.cu
M. Merino-Ibarra (*)
Unidad Académica de Ecología Marina, Instituto de
Ciencias del Mar y Limnología, Universidad Nacional
Autónoma de México, Circuito Exterior s/n,
Ciudad Universitaria, Coyoacán,
Mexico DF, Mexico 04510
e-mail: mmerino@cmarl.unam.mx
M. Merino-Ibarra
e-mail: mmerino56@gmail.com
R. González-De Zayas : F. Matos-Pupo
Centro de Investigaciones de Ecosistemas
Costeros, Cayo Coco,
Morón, Ciego de Ávila, Cuba 69400
M. F. Soto-Jiménez
Unidad Académica Mazatlán, Instituto de Ciencias del Mar
y Limnología, Universidad Nacional Autónoma de México,
Av. Joel Montes Camarena S/N, Apartado Postal 811,
82040 Mazatlán, Sinaloa, Mexico
based on the NH4+ and NOx– concentrations in the rain.
Cold fronts and troughs, coming from the west,
contributed most to rain (41%) and to N deposition,
followed by tropical waves and storms coming from the
east, which caused 31% of the rain. Average concen-
trations of NH4+ and NOx– in the rain were 8.8 and
8.3 μM. NOx– presented a clearly decreasing trend
(0.26 μM per month), decreasing by half during 2005–
2007. Total N deposition averaged 3.23 kg N
ha−1 year−1, similar to that found in Virgin Islands and
Puerto Rico, but lower than previously measured in Cuba
and in nearby areas of the USA and than model
predictions for the oceanic region around Cuba. These
low values and the decreasing trend found are attributed
to drastic reduction of fossil fuel and fertilizer use in Cuba
since 1990. Because land input has decreased even more
drastically, deposition seems to be nowadays the most
important N source to the coastal zone of Cayo Coco.
The δ15N range of seagrass (Thalassia testudinum) and
macroalgae (Penicillus dumetosus) in the area (−1.83‰
to 3.02‰ and +1.02‰ to +4.17‰, respectively) sustain
that atmospheric sources (deposition and N2 fixation)
comprise 70–90% of the N budget.
Keywords Rainfall . Nutrient . Budget . Isotopic .
Nitrogen deposition . Seagrass
1 Introduction
Humans have impacted the global nitrogen (N) cycle
through their input of reactive N to the Earth’s surface
1126
(Schlesinger 2009). Anthropogenic N emissions to
the atmosphere have significantly increased since
the mid-1800s, and future increases are expected
(Duce et al. 2008). Atmospheric deposition of N to
the ocean has also increased over the past 200 years
following the intensification of anthropogenic N
production from agricultural and fossil fuel sources
(Duce et al. 2008; Galloway et al. 2008). Because of
this, atmospheric transport and deposition is an
increasingly important pathway for reactive N
entering the open ocean, often poorly represented
in analyses of open ocean anthropogenic impacts
(Duce et al. 2008).
Models predict that anthropogenic emissions will
continue to increase, and by 2030, a ∼4-fold increase
in total atmospheric N deposition to the ocean could
be reached (Dentener et al. 2006), although there is a
high uncertainty in the predictions. The spatial
distribution of atmospheric deposition to the ocean
has also changed greatly, and it is expected to keep
changing (Duce et al. 2008). It is therefore important
to monitor N depositions, particularly in the oceans
and coastal areas. However, the amount of coastal and
open ocean measurements is very limited, and it is
difficult to estimate their representativeness for larger
regions (Dentener and Prospero 2009).
On the other hand, N is an essential nutrient in
terrestrial and marine ecosystems. The availability of
reactive N limits primary production, the conver-
sion of inorganic carbon to organic carbon, in
much of the ocean. N sources to the surface ocean
include upwelling and subsurface water entrain-
ment or mixing, continental runoff, N2 fixation,
and atmospheric deposition (Duce et al. 2008). The
North Atlantic is particularly affected by increased
deposition because it is very nutrient impoverished
and it is located downwind of highly industrial-
ized continents (Zamora et al. 2010). It is estimat-
ed that N deposition has become an increasingly
important new source of N to marine organisms
there (Krishnamurthy et al. 2007).
In the most oligotrophic regions of the Atlantic,
such as the Caribbean Sea, the relative importance
of N deposition may be greater. In Cuba, due to
the absence of important coastal upwelling, and to
the low tidal ranges, the main sources of nutrients
to the coastal zone are limited to atmospheric
deposition, N2 fixation and river, groundwater, and
wastewater discharges (Baisre 2006). Nevertheless,
Water Air Soil Pollut (2012) 223:1125–1136
the economic crisis that started in Cuba since the
collapse of the socialist countries in 1989 has caused
drastic reductions of N inputs in the country,
including the use of fertilizers, industrial activity,
and fossil fuel consumption. These changes, coupled
to the extensive river damming throughout the
country (Baisre and Arboleya 2006), have caused a
significant decrease of continental nutrient inputs to
coastal Cuban waters (Baisre 2006). Therefore, it is
expected that the other N sources, including deposi-
tion, have increased their relative contribution to the
coastal N budget. Assessing the relative contribution
of N deposition is particularly important for coastal
ecosystems dominated by coral reefs and seagrasses,
such as the Sabana-Camagüey Archipelago in the
North coast of Cuba (Alcolado et al. 1999).
Stable N isotope (δ15N) analysis has proven to be
a powerful tool in discriminating between natural
and anthropogenic sources of N in marine and
aquatic environments (Sherwood et al. 2010). The
utility of this tool arises from the different δ15N
signatures of different N sources. For example,
newly fixed N has a δ15N from −3‰ to 0‰, while
dissolved inorganic N derived from sewage has a
much higher δ15N in the range from +6‰ to +22‰
(Heaton 1986; Sherwood et al. 2010). Primary
producers that assimilate N over periods of weeks
or longer, such as seagrasses and macroalgae, are
ideal for identifying N sources because their δ15N
signatures closely reflect the δ15N of source nutrients
(Carrouthers et al. 2005; Sherwood et al. 2010).
This article follows atmospheric N deposition in
the coastal zone of Cayo Coco, Cuba, for a multi-
annual period (2005–2007) to evaluate its recent
temporal trends, as well as to assess the relative
importance of deposition in the N budget to coastal
waters in this zone and similar ones in the Caribbean
Sea. Stable N isotope (δ15N) analysis of seagrass and
macroalgae is used as an independent means of
assessing N sources.
2 Methods
2.1 Study Area
The coastal zone of Cayo Coco is part of the Sabana-
Camagüey Archipelago, which extends through the
central section of the northern coast of Cuba
Water Air Soil Pollut (2012) 223:1125–1136
(Fig. 1). Cayo Coco and adjacent keys separate the
shallow Los Perros and Jigüey Bays from the
deeper water. On the outside of the keys, a
submerged coral reef runs parallel to the coast.
Seagrass (Thalassia testudinum) meadows extend
between the reef and the shoreline, which is mostly
formed by sand beaches. The shallow (0–2.5 m) Los
Perros and Jigüey behave like extensive hypersaline
lagoons because freshwater input is very low and the
bays are also quite isolated from the open sea. In
land, there are two freshwater lagoons, La Leche and
La Redonda, where most of the annual 130 million
m³ land drainage from rainfall is collected and
retained. Freshwater reaches the bays only under
extreme rainfall events (Alcolado et al. 1999).
The average atmospheric temperature is 25.0°C in
the region. Dominant wind direction is from the East.
Diverse synoptic processes (SP) bring air masses from
other directions, mainly from the west and the
northeast. The average annual rainfall is 1,129 mm.
Fig. 1 Map of the coastal
zone of Cayo Coco, Cuba
showing the location of
Meteorological Station 339.
The stars indicate the two
areas where seagrass and
macroalgae were collected
for δ15N at ten different
sites. Depths are in meters
1127
Most of these rains fall from May to October,
although there is also a high variability associated to
the SP taking place.
2.2 Rainfall, N Concentrations, and Deposition
Daily reports from station 339 of the Cuban Net of
Meteorological Stations, located in Cayo Coco
(Fig. 1), were used to study the temporal evolution
of rainfall. The period 1990–2003 was used as a
historical reference. During 2005–2007, rainfall sam-
ples for N determinations were randomly taken,
corresponding to 20% of the reported rainfall during
2005, 40% during 2006, and 50% during 2007. The
weather associated to each sample was analyzed to
identify the SP that caused the rain and the related air
mass trajectory.
The samples were analyzed for dissolved inorganic
N (NH4+ and NO3− +NO2− (NOx−)) with a Skalar
SAN Plus segmented flow analyzer using standard
1128 Water Air Soil Pollut (2012) 223:1125–1136

methods adapted by Grasshoff et al. (1983) and
analytical circuits suggested by Kirkwood (1994). The
precision obtained with this system was 0.1 μM. Total
inorganic N (TIN) was calculated as (NH4+ +NOx–).
Wet N deposition was calculated on a monthly basis by
multiplying the accumulated rainfall by the mean
monthly concentrations (NH4+, NOx−, and TIN).
Because of the difficulties involved in its determination
(Clark and Kremer 2005), dry N deposition was
estimated from wet deposition. It is generally accepted
(Valiela et al. 1997) that dry N deposition can be
assumed to amount about the same than wet deposition.
In this case, we calculated it using the dry/total
deposition proportion (40%) found specifically for the
Cuban region by Cuesta et al. (1998). Cross-
correlations were used to evaluate the effect of rainfall
and concentration variations over deposition variations.
2.3 N Budget for the Coastal Zone of Cayo Coco
2.4 N Isotopic Composition of T. testudinum
and Penicillus dumetosus
The stable N isotopic composition of the seagrass T.
testudinum and the seaweed P. dumetosus, found
throughout the area, was used to assess the dominant
sources of N in the coastal zone of Cayo Coco.
Samples were collected at ten sites located east
and west of Cayo Coco station (Fig. 1) during the
dry (February) and rainy (October) seasons of 2007.
The collected vegetal tissue was rinsed with distilled
water and dried to constant weight at <80°C. After
finely grounded in a porcelain mortar, the dry
samples were exposed to a HCl–saturated atmosphere
for 4 h, and dried again in an oven. Homogenized
sample aliquots (∼5 mg) were packed in aluminum
vials and sent to the Isotope Analysis Laboratory
of the University of California at Davis for N
isotopes determination.

In order to assess the relative importance of N

deposition at Cayo Coco, the magnitude of the main
N fluxes to its coastal zone was estimated. The land N
fluxes (runoff, groundwater, and wastewater) were
calculated using the data of the N and phosphorus
budget of the Land–Ocean Interactions in the
Coastal Zone program at Laguna Larga, Cayo Coco
(Gonzalez-De Zayas and Merino-Ibarra 2010). N2
fixation was estimated using rates from the literature
(Howarth et al. 1988). Oceanic N fluxes to the coastal
zone were considered negligible.
3 Results
3.1 Rainfall and Synoptic Processes
Yearly rainfall during 2005, 2006, and 2007 was
slightly below the historical mean of the 1990–2003
period (Fig. 2). Annual rainfall was the lowest (86%
of the historical mean) during 2006. Monthly rainfall
distribution (Fig. 3) showed higher rainfall peaks
during the rainy season in 2005 (September–October)

Fig. 2 Annual accumulated Annual acumulated rain (1990 - 2003) Historical average rain (1990 - 2003)
rainfall for 1990–2007 at the
coastal zone of Cayo Coco,
1400
Cuba. The continuous line
Historical Average: 1129.2 mm
depicts the annual historic
1200 1125.1
average (1990–2003) 1004.5
1000 976.7
Rainfall (mm)

800

600

400

200

0
90

91

92

93

94

95

96

97

98

99

00

01

02

03

04

05

06

07
19

19

19

19

19

19

19

19

19

19

20

20

20

20

20

20

20

20

Year
Water Air Soil Pollut (2012) 223:1125–1136 1129

Fig. 3 Monthly rainfall Monthly Rain (2005 - 2007) Monthly historic average (1990 - 2003)
(bars) at the coastal zone 350
of Cayo Coco, Cuba during
2005–2007 compared to 300
the monthly historical
(1990–2003) average 250

Rainfall (mm)
(dark line)
200

150

100

50

0
J F MA M J J A S O ND J F MA M J J A S O N D J F MA MJ J A S ON D
2005 2006 2007
Months

and 2007 (May–June). Six different kinds of SP that
caused rain were identified, aside from local processes
(Table 1). Cold fronts and troughs coming from the
west were the most frequent, and they accounted for
41% of the rain, followed by tropical waves and
storms coming from the east, which caused 31% of
the rain.
3.2 Nitrogen Concentrations in Rainfall
NH4+ concentration in the rainfall sampled in Cayo
Coco averaged 8.8 μM (Table 2) and showed a clearly
seasonal pattern during 2005–2007 (Fig. 4). Annual
maxima occurred before the peak of the rainy season
(21.1 μM in June of 2005, 19.0 μM in May of 2006,
and 18.1 μM in April of 2007) and declined
sharply to an annual minimum during the follow-
ing months. The lowest values (0.3 μM) occurred
in September of 2005 and August of 2006. A
slight decreasing trend in NH4+ concentration was
found over the 2005–2007 period.
Rainfall NOx− concentration was on average
slightly lower (8.3 μM) than NH4+ concentration
(Table 2). NOx− content in the rainfall also showed
important variations (range, 1.6 to 18.7 μM), but it
did not show a defined seasonal pattern (Fig. 4). A
clear decreasing trend of the NOx− content took place
over the three sampled years with an average decrease
of 0.26 μM per month. Annual NOx− concentration
diminished by half from 2005 (12.1 μM) to 2007
(6.1 μM, Table 2). Average TIN was 17.15 μM in
Cayo Coco during 2005–2007. The temporal trend of
TIN showed both the seasonal pattern found in NH4+
and the long-term decrease observed in NOx− (Fig. 4).
3.3 N Deposition
On a monthly basis, the relative contribution of
NH4+ and NOx− to N wet deposition varied in time
(Fig. 5), but their rates were on average quite similar
(NH4+ 0.098 kg N ha−1 month−1 and NOx− 0.095 kg
N ha−1 month−1). The calculated N dry deposition

Table 1 Synoptic processes identified, their air mass trajectories, rain contribution, and N concentrations at the coastal zone of Cayo
Coco, Cuba

Synoptic processes Air mass trajectory Number of cases % of accumulated rain NH4+ (μM) NOx− (μM)

Cold fronts and troughs From the west 19 41 9.8 7.1

Tropical waves and storms From the east 13 31 7.1 8.1
Anticyclones From the northeast 2 12 17.8 8.0
Wave–depressions interactions From the west and the east 2 8 6.9 0.6
Low pressures From the south 5 5 9.0 7.7
(from the Gulf of Mexico)
Local processes Local 9 3 6.7 7.3
1130 Water Air Soil Pollut (2012) 223:1125–1136

Table 2 N rainfall concentrations (micromolar) measured at the coastal zone of Cayo Coco, Cuba and at other sites around the world

Site Period Concentration Reference

NH4+ NOx− TIN

Cayo Coco, Cuba Average (2005–2007) 8.8 8.3 17.1 This study
2005 10.1 12.1 22.2
2006 7.0 7.3 14.3
2007 9.7 6.1 15.8
Havana, Cuba 1986–1991 29.8 29.4 59.2 Cuesta et al. 1998
Falla, Cuba (rural) 1986–1991 21.8 8.2 30.0
Colón, Cuba (rural) 1986–1991 24.4 9.9 34.3
La Palma, Cuba (rural) 1986–1991 4.0 9.9 13.9
Santiago, Cuba 1986–1991 1.2 22.3 23.5
Sarasota, Florida, USA 1999–2000 2.1–29.0 3.7–56.0 5.7–85.0 Dillon and Chanton 2005
Tampa Bay, FL, USA 1996–2004 5.0–1,150 0.0–157.8 5.0–1,307.8 Strayer et al. 2007
Mullica River–Great Bay 2004–2005 44.6 26.3 70.9 Ayars and Gao 2007
Estuary, NJ, USA
Delaware Coast, USA 2001–2003 17.0 17.2 34.2 Scudlark et al. 2005
Maracaibo Lake, Venezuela 1991–1994 45.0–102.8 10.6 51.6–111.4 Morales et al. 1999
Fiji Islands 1990 3.6–33.6 3.6–19.3 7.2–52.9 Waterloo et al. 1997

averaged 0.128 kg N ha−1 month−1, and total N
deposition averaged 0.320 kg N ha−1 month−1. Total
N deposition also exhibited important time variations
on a monthly basis, mainly following the variations
in rainfall intensity. Cross-correlation between these
two parameters yielded a high (r=0.81) and signif-
icant (p < 0.05) correlation at 0 lag. Correlation
coefficients at other lags were not significant, but
indicated a coupled annual pattern for both param-
eters. Total N deposition also reflected the seasonal
pattern of NH4+ and the decreasing trend of NOx−,
but significant cross-correlation coefficients between
N concentration and deposition were much smaller
(0.37 at lag=0 and 0.38 at lag=2). On a yearly basis,
total N deposition in Cayo Coco averaged 3.85 kg
N ha−1 year−1 during 2005–2007. N deposition
decreased from 4.84 kg N ha−1 year−1 in 2005 to
3.29 and 3.41 kg N ha−1 year−1 in 2006 and 2007,
respectively (Table 3).
3.4 N Budget for the Coastal Zone of Cayo Coco
Table 4 allows comparison of the N deposition flux to
the coastal zone of Cayo Coco with the other
fluxes estimated for the area. The N flux through
runoff was estimated at 411 g N ha−1 year−1, while
the groundwater and wastewater fluxes were much
smaller (0.25 and 3.59 g N ha−1 year−1, respectively).
For N2 fixation estimation, a range was considered
since fixation can vary significantly depending on the
benthic community (Howarth et al. 1988). Lower
values are for bare sediments (370 g N ha−1 year−1),
while the higher values correspond to seagrass meadows
(1,000 g N ha−1 year−1).
3.5 Nitrogen Isotopic Composition of T. testudinum
and P. dumetosus
The stable N isotope ratio (δ15N) measured in T.
testudinum leaves ranged from −1.83‰ to +3.02‰
and averaged 0.55‰ (Table 5). The seagrass pre-
sented slightly lower δ15N values in the dry season
sampling (mean, +0.37‰) than during the wet season
sampling (mean, +0.72‰), although this difference
was not statistically significant (p<0.05). The macro-
algae P. dumetosus samples presented higher δ15N
values than the seagrass (T. testudinum), ranging from
+1.02‰ to +4.17‰ and averaging +2.67‰. On a
seasonal basis, δ15N in the macroalgae exhibited an
inverse pattern: it was slightly higher (mean, +3.14‰)
in the dry sampling (February 2007), than in the rainy
one (mean, +2.25‰).
Water Air Soil Pollut (2012) 223:1125–1136 1131

Fig. 4 Monthly wet concen- NH4 (2005 - 2007) Trend line (NH4 (2005 - 2007))
trations (micromolar) of
NH4+, NO3− + NO2− (NOx−) y = -0.0911x + 10.615
and total N (TIN) at the 20 2
R = 0.0245
coastal zone of Cayo Coco,
Cuba during 2005–2007.
Dashed line and equations in
15
box show the results of a

NH4 - (µM)
linear regression for each
data set
10

0
J FMAMJ J ASONDJ FMAMJ J ASONDJ FMAMJ J ASOND
2005 2006 2007 Months
NO3 + NO2 (2005 - 2007) Trend line (NO3+NO2 (2005 - 2007))

y = -0.2604x + 13.392
20 2
R = 0.4239
(µM)

15
-
2
+ NO

10
-
NO 3

0
J FMAMJ J ASONDJ FMAMJ J ASONDJ FMAMJ J ASOND
2005 2006 2007
Months
N (Wet) Trend line (N (2005 - 2007))

35 y = -0.3515x + 24.007
2
R = 0.1972
30

25
TIN (µM)

20

15

10

0
J FMAMJ J ASONDJ FMAMJ J ASONDJ FMAMJ J ASOND
2005 2006 2007 Months
1132 Water Air Soil Pollut (2012) 223:1125–1136
Fig. 5 N deposition 1.4
(kilograms N per hectare N (Dry)
per month) at the coastal 1.2 NOx (Wet)
NH4 (Wet)

N Deposition (kg ha month )

zone of Cayo Coco, Cuba

-1
during 2005–2007. Asterisk
wet deposition for these 1.0
months (when concentra-

-1
tions were not measured) 0.8 *
was calculated using the
measured rainfall and the
0.6
interpolated concentration
(except for January and
February of 2005, when 0.4
*
the March concentrations
were used) 0.2 *
* * * *
* * *
* * *
0.0
J F M AM J J A SO N D J F M A M J J A S O N D J F MA M J J A S O ND
2005 2006 2007
Months

4 Discussion were the most important SP, accounting for 43% of

the NH4+ and 57% of the NOx−. The most likely
In terms of their contribution to N deposition in Cayo sources of the N deposited by tropical waves are the
Coco, tropical waves and storms coming from the east cities of Moa, Nicaro and Nuevitas, located east of

Table 3 N deposition rates (kilograms N per hectare per year) measured at the coastal zone of Cayo Coco, Cuba and at other sites
around the world

Site Period N deposition Reference

NH4+ NOx− Wet Dry Total

Cayo Coco, Cuba Average (2005–2007) 1.17 1.14 2.31 1.54 3.85 This study
2005 1.33 1.58 2.90 1.94 4.84
2006 1.00 0.98 1.97 1.32 3.29
2007 1.19 0.86 2.04 1.36 3.41
Falla, Cuba 1986–1991 3.8 2.0 4.0 1.9 5.9 Cuesta et al. 1998
Colón, Cuba 1986–1991 9.0 1.8 2.7 6.0 8.7
Santiago, Cuba 1986–1991 5.5 1.6 7.1 4.2 11.3
Havana, Cuba 1986–1991 7.7 2.5 10.2 6.5 16.7
La Palma, Cuba 1986–1991 4.0 1.8 5.8 2.8 9.6
Tampa Bay, Florida, USA 1996–2004 1.75 2.30 4.05 – – Strayer et al. 2007
Virgin Islands – – – 1.06 0.94 2.0 Navato 2007
Puerto Rico Island – – – 2.70 1.22 3.92 Ortiz-Zayas et al. 2006
Mullica River-Great Bay 2004–2005 6.21 3.77 9.98 1.54 11.52 Ayars and Gao 2007
Estuary.NJ, USA
Massachusetts, USA – – – 7.5 7.5 15.0 Valiela et al. 1997
New York State, USA 2002–2004 2.7–4.3 3.2–5.6 5.9–9.8 – – Golden and Boyer 2009
Delaware Coast, USA 2001–2003 1.6–2.9 2.3–2.8 3.9–5.7 – – Scudlark et al. 2005
Maracaibo Lake Venezuela 1991–1994 4.2 1.2 5.4 – – Morales et al. 1999
English East Coast – – – – 17.08 Spokes and Jickells 2005
Fiji Island 1990 1.5–5.9 0.9–2.9 2.4–8.8 – – Waterloo et al. 1997
Coasts of Cuba (from models) 2000 – – – – 5.6–7.0 Dentener et al. 2006;
Duce et al. 2008
Water Air Soil Pollut (2012) 223:1125–1136 1133

Table 4 Estimated N budget (grams N per hectare per year) reach Cayo Coco in the center of the island. The
and relative contribution of N sources at the coastal zone of
relatively high (for a rural area) N deposition rates
Cayo Coco, Cuba
found at La Palma (Table 2), which is located near the
N source N flux (g N ha−1 year−1) Relative western tip of Cuba, are consistent with this possibility.
contribution (%) The average N concentrations in the rainfall of
Runoff 411 8–10
Cayo Coco during 2005–2007 are lower than most of
the previous measurements reported for diverse places
Groundwater 0.25 <0.05
throughout the world (Table 2). In highly industrial-
Wastewater 3.59 <0.1
ized regions, N concentrations can be one to two
N2 fixationa 370–1,000 9–22
orders of magnitude higher than in Cayo Coco, like in
Deposition 3,230 70–80
the Yellow and East Seas of China (up to 1,360 μM of
Total 4,020–4,065 100
NH4+ and 396 μM of NOx–, Zhang et al. 2007). N
a
Calculated after Howarth et al. (1988) for bare sediments and concentrations are also much higher in nearby cities of
seagrass meadows, respectively Florida (Table 2), such as Tampa (Strayer et al. 2007)
and Sarasota (Dillon and Chanton 2005). Our measure-
Cayo Coco, where there are numerous emitting ments are also lower than those previously reported
industries (Cuban Government 1999). The transport within Cuba, particularly in the case of NOx−. Between
and deposition of N emissions towards the west may 1986 and 1991, Cuesta et al. (1998) found NOx− levels
be a previously unconsidered impact of tropical of up to 29.4 and 22.3 μM at Havana and Santiago, the
hurricanes as they sweep Cuba. Cold fronts and two major cities in Cuba, and much lower in rural
troughs coming from the west contributed next, with stations, like La Palma (9.9 μM), Colón (9.9 μM), and
28% of the NH4+ and 19% of the NOx− deposited. N Falla (8.2 μM). The NOx− concentrations found in
deposition associated to cold fronts could be impor- Cayo Coco during 2005 are similar to those previously
tant because the N concentrations in the Southeastern found in rural areas of Cuba, but the averages for
USA are high (Table 2). However, because the fronts 2006 and 2007 were 30% lower. In the case of
sweep Cuba from the west, it is likely that they NH4+, the levels in Cayo Coco are also much smaller
discharge an important fraction of their N before they to those previously found in other rural areas like

Table 5 Nitrogen isotopic

composition (δ15N, ‰) of Source δ15N (‰) References
Thalassia testudinum and
Penicillus dumetosus Mean Range
at Cayo Coco, Cuba com-
pared to other sites in the Thalassia testudinum
Caribbean and to common Cayo Coco, Cuba +0.55 −1.83 to +3.02 This study
N sources to the
Bahamas −0.2 Kieckbusch et al. 2004
coastal zone
Biscaine Bay, Florida +1.4 Kieckbusch et al. 2004
Puerto Morelos Reef, Mexican Caribbean +1.69 Carrouthers et al. 2005
Akumal and Xaak, Mexican Caribbean +6.5 +6 to +7 Mutchler et al. 2007
Nichupté Lagoon, Mexican Caribbean +9.06 Carrouthers et al. 2005
Penicillus dumetosus
Cayo Coco, Cuba +2.67 +1.02 to +4.17 This study
Bahamas +0.9 Kieckbusch et al. 2004
Biscaine Bay, Florida +1.7 Kieckbusch et al. 2004
Akumal Bay, Mexican Caribbean +4 Mutchler et al. 2007
N sources
Deposition (rain) −12.0 to +4.0 Sherwood et al. 2010
N fixation −3.0 to 0
Sewage +6.0 to +22.0
1134
Colón (24.4 μM) and Falla (21.8 μM). As a result,
the total N concentrations found at Cayo Coco
(overall mean, 17.1 μM; 2006 mean, 14.3 μM) are
smaller than all previously reported in Cuba during
1968–1991, with the only exception of one measure-
ment (at rural La Palma, 13.9 μM). The low concen-
trations found in Cayo Coco during 2005–2007, and the
decreasing trend they show, are likely due to an
important decrease of N emissions to the atmosphere
in Cuba after the drastic reductions in fossil fuel
consumption and use of fertilizers, due to loss of imports
from the Soviet Union after 1990 (Baisre 2006).
N deposition in Cayo Coco during 2005–2007 was
also lower than previously measured in Cuba, and
than in most other studied areas of the Americas
(Table 3). Only the Virgin Islands exhibit lower N
deposition rates (2.0 kg N ha−1 year−1, Navato 2007)
than Cayo Coco, while similar values (3.92 kg N
ha−1 year−1, Ortiz-Zayas et al. 2006) have been found
recently in Puerto Rico Island. It is to be noted that
these two cases are islands of a much smaller size
than Cuba, and therefore, lower depositions would be
expected because of higher oceanic influence (see for
example Dentener et al. 2006). Furthermore, our data
also show an important decrease (by 30%, Table 3) in
N deposition during 2005–2007. Since annual rainfall
was very similar for these 3 years (Fig. 2), the decrease
of deposition was not due to rainfall variations. A 50%
decrease of the NOx− concentration during this period
(Fig. 4, Table 2) was the main cause for the observed
decrease of the N deposition in the area.
In most parts of the planet, there is an increasing
trend in fuel consumption and fertilizer application, and
N deposition rates are increasing (Duce et al. 2008;
Galloway et al. 2008). In contrast, the decrease in fuel
consumption and fertilizer use in Cuba caused by the
economic crisis (Baisre 2006) is likely responsible for
the decreasing trend in N deposition we found at the
coastal zone of Cayo Coco. The N deposition measured
in Cayo Coco is about half of that predicted by models
for the region around Cuba (5.6–7.0 kg N ha−1 year–1,
Table 3). As the decrease in consumption affects the
entire Cuban archipelago (Baisre 2006), the observed
decreasing trend is likely representative of the
region around Cuba. Our data can be useful to
make better assessments of N deposition within the
Caribbean and to feed the models to obtain the more
precise global N deposition rates that are needed
(Dentener and Prospero 2009).
Water Air Soil Pollut (2012) 223:1125–1136
On the local scale, however, the relative impor-
tance of N deposition seems to be increasing because
generalized river damming and fertilizer shortage
have drastically reduced land inputs of N to coastal
ecosystems in Cuba (Baisre and Arboleya 2006). This
conclusion is sustained by the N budget estimates at
Cayo Coco (Table 4), which indicate that runoff,
groundwater, and wastewater make up a minor part of
the budget, and that atmospheric sources (deposition
and N2 fixation) may currently constitute nearly 90%
of the total input to the coastal zone of Cayo Coco.
Our stable N isotope analyses support that wastewater
is not an important source of N for the Cayo Coco coastal
zone. The low δ15N values observed in T. testudinum
and P. dumetosus at Cayo Coco are far away from the
much higher δ15N signature (+6‰ to +22‰, Sherwood
et al. 2010) found in sewage. Our δ15N data for T.
testudinum (mean, 0.55‰ and range, from −1.83‰ to
+3.02‰) clearly contrast with those found in coastal
areas of the Mexican Caribbean where there are sewage
and groundwater inputs (Table 5). For example, T.
testudinum growing in the eutrophic Nichupté Lagoon,
where wastewater discharge is important (Merino et al.
1992), shows much higher δ15N values (+9.06‰,
Carrouthers et al. 2005), which are clearly within the
range reported for sewage.
The low δ15N values found in T. testudinum at
Cayo Coco also support that N deposition is likely the
most important source of N in the area because they
fall clearly within the range of δ15N values reported for
N deposition (−12‰ to +4‰, Sherwood et al. 2010).
Moreover, our results are also consistent with
those of previous studies in the region (Table 5)
which have also suggested that N deposition is the
main source of N at coastal areas of Bahamas
(Kieckbusch et al. 2004), Florida (Sherwood et al. 2010),
and Bermuda (Hastings et al. 2003).
However, because the δ15N signatures of N
deposition and N2 fixation have overlapping ranges
(−12‰ to +4‰) and N2 fixation (−3‰ to 0‰, Table 5),
other data are needed to determine the exact proportion
between these two sources. One option is the direct
measurement of N2 fixation in the coastal zone of
Cayo Coco. Another is to include the isotopic
composition of other elements in the analysis, to solve
the proportion between these two sources. In any case,
our data all together show that atmospheric sources are
nowadays dominant (70% to 90%) in the N budget of
the coastal zone of this region of central Cuba.
Water Air Soil Pollut (2012) 223:1125–1136
5 Summary
During 2005–2007, rainfall in Cayo Coco was
slightly below average. Cold fronts and troughs
coming from the west accounted for 41% of the rain
and were the most frequent of the six kinds of SP that
caused rain in Cayo Coco. Tropical waves and storms
coming from the east caused 31% of the rain, but
were the most important SP in terms of their
contribution to N deposition in Cayo Coco, account-
ing for 43% of the NH4+ and 57% of the NOx−. NH4+
concentration in the rainfall of Cayo Coco showed a
marked seasonal pattern, peaking at the start of the
rainy season and declining sharply in the following
months. Rainfall NOx− concentration presented a
clear decreasing trend, diminishing by half during
the 3 years of observations. As a consequence, TIN
concentration in the rainfall and N deposition also
decreased during this period. N concentrations and
deposition observed in Cayo Coco are lower than
most of those reported throughout the world, includ-
ing the nearby cities of Florida. Our measurements are
also lower than those previously reported within
Cuba, particularly in the case of NOx−. This decreas-
ing trend is attributed to the drastic reduction of fossil
fuel and fertilizer use in Cuba caused by the loss of
imports from the Soviet Union after 1990. Deposition
rates at Cayo Coco show that there are regions where
N deposition is decreasing and can therefore be useful
for a better assessment of global predictions. Because
land runoff input has also been drastically reduced, N
deposition seems to dominate the N budget of the
coastal zone of Cayo Coco.
Acknowledgments The authors are grateful to R. Almira and
Sergio F. Castillo for nutrient analysis and to L. Castellanos, A.
Fernández, M. Reiné, L. Vázquez, and A. Fernández for rain
collection. We acknowledge Marcos Merino and Patricia M.
Valdespino for their thorough revision of the English. We are
grateful to the editor and reviewers who helped improve
significantly this paper through their sound revision of the
manuscript submitted.
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