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Wetland Conservation in the Gulf of Mexico:

The Example of the Salt Marsh Morning Glory,
Ipomoea sagittata

Article in Wetlands May 2015

DOI: 10.1007/s13157-015-0662-2

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Wetlands
DOI 10.1007/s13157-015-0662-2
ORIGINAL RESEARCH
Wetland Conservation in the Gulf of Mexico:
The Example of the Salt Marsh Morning Glory,
Ipomoea sagittata
Guillermo Huerta-Ramos 1 & Patricia Moreno-Casasola 2 & Victoria Sosa 1
Received: 15 December 2014 / Accepted: 6 May 2015
# Society of Wetland Scientists 2015
Abstract Global climate change will have major effects on
wetlands, ecosystems with elevated biodiversity and of
enormous economic importance. Using ecological niche
modeling and genetic data from three plastid DNA markers
sequenced from 96 plants, we studied the salt marsh morn-
ing glory, Ipomoea sagittata, to understand the impact that
future global warming and increasing sea level may have on
aquatic plant conservation, distribution and genetic connec-
tivity on the Gulf of Mexico. Data suggest that genetic
variation is low and lacks structure; probable causes in-
clude high gene flow, clonal reproduction or use of ineffec-
tive molecular markers. Global warming models for its po-
tential distribution in the year 2080 predict a loss of suitable
habitat in its northern inland distribution (Cuatro Cinegas
Basin), while its coastal and southern habitats increase.
Genetic connectivity decreases along the coast owing to a
rise in sea level (Yucatn, Laguna Madre Basin,
Usumacinta Basin). Three zones are identified, each requir-
ing a different conservation strategy: 1) A saltwater intru-
sion zone where most protected areas are; 2) A stability
zone which may offer optimal conditions for the creation
of protected areas; 3) A zone of range expansion that may
cause ecological instability, reducing species richness and
promoting colonization by opportunistic species.
* Victoria Sosa
victoria.sosa@inecol.mx
1
Biologa Evolutiva, Instituto de Ecologa A.C., Carretera antigua a
Coatepec 351, El Haya, Xalapa 91070, Veracruz, Mexico
2
Ecologa Funcional, Instituto de Ecologa A.C., Carretera antigua a
Coatepec 351, El Haya, Xalapa 91070, Veracruz, Mexico
Keywords Climate change . Cuatro Cinegas Basin .
Ecological connectivity . Ecological niche modeling . I.carnea
subsp. fistulosa . Sea level rise . Yucatan coast
Introduction
There is extensive evidence confirming that global climate
change results from human activity, primarily from the in-
crease in greenhouse gas emissions (e.g., Oreskes 2004;
Anderegg et al. 2010). The consequences of global climate
change and additional anthropogenic disturbances have
caused an extinction crisis and a concomitant biodiversity loss
(Bellard et al. 2012). Forecasting suggests that global climate
change and the increase in sea level will critically affect wet-
lands through mechanisms such as erosion, flooding, seawater
intrusion and changes in climate cycles (Nicholls and Tol
2006). Climate simulation models for the next 100 years pre-
dict that wetlands will decrease by 40 to 70 % worldwide; the
most affected regions comprise the Mediterranean, Southeast
Asia, the Gulf of Mexico and the Caribbean (Nicholls et al.
1999; Craft et al. 2009; Landgrave and Moreno-Casasola
2012). Recent simulations following the climate scenarios
for 2050 and 2080 by the Intergovernmental Panel for
Climate Change (IPCC) anticipate a reduced habitat for sev-
eral species restricted to the Gulf of Mexicos coastal dunes.
Furthermore, the effect of an increase in sea level together
with the unsuitability of inland environmental conditions will
create a coastal squeeze pattern, greatly reducing the area of
sustainable wetlands (Mendoza-Gonzlez et al. 2013).
Together, ecological niche modeling for future global climate
change scenarios and genetic diversity estimates provide a
broader perspective of the consequences of climate change
on species distribution and their viability (reviewed in Pauls
et al. 2013). The inclusion of genetic data allows us to propose

future conservation strategies for cryptic evolutionary lineages
and preserve evolutionary potential (Blint et al. 2012).
Severe changes in distribution areas are expected to cause
local population extinction, genetic diversity reduction or the
loss of the species as a whole (Fitzpatrick et al. 2008; Sork
et al. 2010; Habel et al. 2011). Recent studies in
Mediterranean and Tropical areas suggest that shifts in species
distribution will have heterogeneous consequences: a mixture
of future distribution expansion and contraction highly de-
pends on how conditions vary with elevation and latitude,
and therefore this lack of a pattern may hinder extrapolation
to other species in need of conservation (Temunovi et al.
2013; Valle et al. 2014).
Changes in the genetic variation and connectivity of popu-
lations are likely to occur depending on the ecological hetero-
geneity and variations in habitat suitability. Some species will
be negatively affected, where populations become isolated,
thus reducing their ecological and genetic capability of
responding to climate change (Neel 2008; Velo-Antn et al.
2013). Although a reduction in connectivity can take a long
time to affect populations, drastic phenomena like a rise in sea
level will alter the distribution of organisms extensively as
rapid changes in ecological factors exceed the normal scope
of the capacity of organisms to adapt (Gilman et al. 2008;
Hoegh-Guldberg and Bruno 2010). In addition, populations
may be undermined by increased competition, further reduc-
ing their range, while highly competitive species are positively
selected reaching a state of ecological dominance (Klanderud
and Totland 2005).
One of the main approaches for the conservation of
Mexican wetlands has been in situ conservation in Protected
Natural Areas (PNA) (CONANP, http://www.conanp.com.
mx/; Bezaury-Creel and Gutirrez-Carbonell 2009). Ideally,
PNAs are designed taking into consideration high species di-
versity, endemism and the presence of endangered species,
other important criteria like threat level, biotic networks and
local community participation are often considered as well
(Eken et al. 2004; Rodrigues et al. 2004). However, global
climate change effects include the displacement of some spe-
cies from their original range of distribution, thus modifying
reserve connectivity and reducing conservation effectiveness
(Arajo et al. 2004). Conservation strategies should therefore
include forecasts and the evaluation of both habitat suitability
and spatial connectivity (Hannah et al. 2007; Arajo 2009;
Alagador et al. 2014).
To assess the impact of global climate change and rising
sea levels on the distribution, genetic connectivity and genetic
diversity of the Gulf of Mexico wetlands we selected the
climbing vine Ipomoea sagittata Poir. that occurs in undis-
turbed oligohaline marshes. This morning glory is an amphi-
Atlantic subtropical species. It may be native to the Americas;
however, it was introduced to Europe primarily as an orna-
mental (Austin and Human 1996; Elorza et al. 2004; Austin
Wetlands
2014). I. sagittata has been naturalized throughout the
Mediterranean where its populations are apparently declining
due to wetland degradation (Medagli et al. 1992; Rhazi and
Grillas 2010; Austin 2014). I. sagittata is a locally abundant
plant throughout the wetlands of the Gulf of Mexico (south-
eastern USA, eastern Mexico and the Yucatan Peninsula) and
it has also been reported for the Greater Antilles. There are
some isolated inland populations in the Pnuco River Basin in
the Sierra Madre Oriental mountain range and also alongside
ponds in the Chihuahuan Desert in northern Mexico (Pinkava
1984; Lot et al. 1999). This species is a perennial climbing
vine, restricted to banks along marshes, so it is considered a
subaquatic species (Novelo-Retana 2006). It flowers from
May to August, with lilac colored flowers that are diurnal,
funnel-shaped and probably bee-pollinated; it mainly repro-
duces by seed though it can reproduce clonally (McDonald
1991, 1993; Elorza et al. 2004). Seeds have 2- to 5-mm-long
hairs that favor water dispersal (Austin et al. 2001). This
morning glory is mostly distributed on coastal wetlands and
has been found to be highly sensitive to disturbance and
changes in salinity (Flynn et al. 1995; Baldwin and
Mendelssohn 1998; Slocum and Mendelssohn 2008).
In this paper we addressed the following questions: i) What
is the genetic diversity pattern of Ipomoea sagittatas popula-
tions on the Gulf of Mexico?, ii) Which regions will be most
affected by a rise in sea level and global climate change in this
area?, iii) How will global climate change affect ecological
and genetic connectivity among I. sagittatas populations?
and iv) Which protected areas will allow for the conservation
and viability of populations of this morning glory?
Materials and Methods
Sampling
A total of 96 individuals from 10 localities were collected
based on records from the MEXU, ENCB, XAL and CICY
herbaria, as well as geographic reports from literature
(Pinkava 1984; Lot et al. 1999; Novelo-Retana 2006; herbar-
ium abbreviations follow Index Herbariorum, http://sciweb.
nybg.org/science2/IndexHerbariorum.asp) (Table 1). Flower
or leaf material was obtained from five to a maximum of 15
scattered individuals from each locality and dried in silica gel.
Collecting sites were georeferenced. Vouchers are listed in the
Appendix 1.
DNA Extraction, Amplification and Sequencing
Total genomic DNA was extracted following the CTAB 2X
protocol from Cota-Snchez et al. (2006). We utilized a set of
widely used non-coding chloroplast regions that have been
identified as highly variable among several angiosperm
Wetlands

Table 1 Populations of Ipomoea sagittata used in this study and their localities

Locality Code N Haplotypes Latitude Longitude Elevation (m a.s.l.)

Mexico, Campeche, Campeche CAM 15 A (13) D(2) 19 55 90 26 25

Mexico, Coahuila, Poza Azul PAZ 7 A(7) 26 55 102 7 748
Mexico, Coahuila, Rio Mezquites MEZ 6 A(6) 26 36 101 55 726
U.S.A, Texas, Port Arthur PAR 5 E (5) 29 17 94 48 4
Mexico, San Luis Potos, Rio Verde RVE 10 A(10) 21 53 100 3 1003
Mexico, Tabasco, Frontera FRO 12 A(12) 18 25 93 08 4
Mexico, Campeche, Candelaria CAL 9 A (6) B(3) 18 06 91 03 30
Mexico, Quintana Roo, Muyil MUY 10 A(8) C (2) 20 20 87 30 0
Mexico, Quintana Roo, Coba COB 13 A (13) 18 58 87 57 5
Mexico, Campeche, Calkin CAL 9 A (9) 20 30 90 22 10

groups: psbD-trnT (KP734321-KP734323), psbJ-petA
(KP734319, KP734320), psbK-psbI (KP734324, KP734325)
(Shaw et al. 2005, 2007, 2014) and the ITS-1 nuclear region
(primers by Miller et al. 1999). PCR products were cleaned
using QIAquick PCR purification kit (Qiagen, California,
U.S.A.). Clean samples were sequenced using BigDye
Terminator Cycle sequencing kits (Perkin Elmer Applied
Biosystems, Foster City, U.S.A.) in an ABI 310 Automated
DNA sequencer (Perkin Elmer Applied Biosystems, Foster
City, U.S.A.). The Sequencher 4.1 program (Gene Codes
2000) was used to correct ambiguities in contiguous se-
quences. Sequences were manually aligned using Se-Al
v.2.0a11 (Rambaut 2002).
Molecular Data Analyses
Total genetic diversity, population genetic diversity, average
intra-population diversity, and inter-population genetic differ-
entiation were calculated using a concatenated chloroplast se-
quences matrix in the program DNASP v.5. 10 (Librado and
Rozas 2009). The differentiation was quantified using the FST
statistic. To detect population expansion a mismatch distribu-
tion was implemented in ARLEQUIN v. 3.5.3 (Excoffier and
Lischer 2010). The degree of population differentiation was
assessed using the hierarchical analysis of molecular variance
(AMOVA) without group definition (Excoffier et al. 1992).
The statistical significance was tested using a permutation
procedure with 10,000 replications.
Ecological Niche Modeling
We built a database from 42 georeferenced localities of Ipomoea
sagittata. Records were obtained from the National Biodiversity
Information Networks open access databases (REMIB; http://
www.conabio.gob.mx/remib_ingles/doctos/remib_ing.html;
accessed January, 2012) and that of the Global Biodiversity
Information Facility (GBIF, www.gbif.org/) (Edwards 2000),
and from our own collections. We selected only American
collections in order to obtain reliable results for the native
range (Gallien et al. 2010; Petitpierre et al. 2012). To
characterize climatic niche, 19 variables were obtained from
the WorldClim project (www.worldclim.com/; (Hijmans et al.
2008) at a resolution of 0.0083/px (ca. 1 km2).
To reduce over-fitting caused by redundant variables, we
selected the most explanatory and uncorrelated variables,
eliminating the others based on the highest and most signifi-
cant correlation coefficients (Spearman correlation index,
r>0.5 and p<0.05) (Schrag et al. 2008). The importance of
the remaining variables was then assessed using a Principal
Components Analysis (PCA). The current potential distribu-
tion for Ipomoea sagittata was modeled and projected for the
2080 IPCC-SRES B2 and A2 scenarios using the maximum
entropy algorithm implemented in Maxent software (ver. 3.3,
Phillips and Dudk 2008). These simulations of future climate
change were developed by the Canadian Centre for Climate
Modeling and Analysis (CCCma). The A2 scenario projects a
human population of 15 billion by the year 2100, slow eco-
nomic and technological development, lower greenhouse gas
emissions and lower aerosol loadings than at present. The B2
scenario foresees slower population growth (10.4 billion by
2100), with an economy that evolves more rapidly and an
emphasis on environmental protection, with lower greenhouse
gas emissions and less intense future warming (IPCC, Special
Report on Emissions Scenarios 2001, in Gaffin et al. 2001). In
this study 80 % of the locality records were considered train-
ing data and 20 % were considered points of validation; the
maximum iterations value was 1,000 with a convergence limit
of 0.00001 and a regularization value of 1. The continuous
prediction obtained (as a probability between 0 and 1) was
transformed to a binary map (0 for absence and 1 for presence)
using as a criterion the Bminimum training presence^ value for
each species, which included over 90 % of the total number of
records in all cases. With the aim of further minimizing over-
prediction (commission errors) we clipped the current and
future distribution maps generated (Martnez-Meyer 2005)
using information about the potential distribution of wetlands

in Mexico provided by the National Institute of Statistics,
Geography and Informatics (INEGI 2005).
Impact of the Rise in Sea Level
Estimates of the increase in sea level for the year 2100 range
from 0.5 to 2 m, with an expected 4 C rise in temperature,
therefore we selected a conservative model with a rise in sea
level of 1 m (Vermeer and Rahmstorf 2009; Nicholls et al.
2011). We used a simulation with a spatial resolution of
1 km and a vertical resolution of 1 m and overlapped it with
potential distribution scenarios A2 and B2 for the year 2080
(Li et al. 2009). We also added polygons delimiting national
and regional protected areas obtained from Bezaury-Creel
(2005). The areas impacted were identified using Quantum
GIS Software 2.2 (Quantum GIS Development Team 2012).
Changes in Genetic Connectivity Among Populations
Ecological dispersal networks for Ipomoea sagittata were cre-
ated by inverting the continuous models generated by Maxent
to obtain a Bdispersal cost^ layer for current and future sce-
narios; the future dispersal cost layer is clipped by a 1 m sea
level rise model (Li et al. 2009). Then, we generated a chlo-
roplast haplotype network by adding up the least-cost paths
(LCPs) among all shared haplotypes from sampled localities.
Likewise, a total pairwise population connectivity model was
generated using all geographic records collected in Mexico.
Both models were generated using SDM toolbox v1.1 in
ArcMap 10.2 (Brown 2014).
Results
Haplotype Diversity and Population Differentiation
We obtained a 2446 bp sequence matrix for three molecular
markers (psbD-trnT, psbJ-petA, psbK-psbI) excluding ITS se-
quences which were found to be monomorphic; five haplo-
types were identified. Haplotype A is the most common and
frequent haplotype among populations representing 87.5 % of
individuals sampled. Unique haplotypes B, C, and D are re-
stricted to the southeast of the Isthmus of Tehuantepec.
Northern inland regions (the Cuatro Cinegas and Pnuco
basins) include only haplotype A (Fig. 1). Genetic diversity
indexes estimated very low nucleotide and haplotypic diver-
sity with only three populations showing more than a single
haplotype (Table 1).
Population differentiation based on chloroplast DNA vari-
ation (Gst: 0.053, p=0.05) indicates a lack of population struc-
ture supported by a fixation index (Fst) of 0.0834, p<0.05)
and a low degree of differentiation for ordered alleles (Nst:
0.003, p<0.05). AMOVA results for populations indicate no
Wetlands
genetic differentiation (FST: 0.087, p<0.05). Demographic
expansion analysis using Tajimas D returned a value of
1.70, (p<0.05.)
Genetic Connectivity Among Populations
Least-cost path models made for current and future climate
A2 scenarios including a sea level rise projection indicated
changes in ecological connectivity. For the haplotype dis-
persal network (Fig. 2a, b), the central part of the distribu-
tion maintains a high degree of connectivity in spite of the
rise in sea level; however, a major change decreases con-
nectivity along the coast, toward the northern part of the
Yucatan Peninsula. The inland southern part of the penin-
sula increases potential connectivity due to the increase in
habitat suitability that is forecasted. Another major de-
crease in connectivity is predicted for the northern part of
the Mexican wetlands due to sea intrusion into the lower
Laguna Madre Basin. Pairwise population connectivity re-
veals a similar pattern (Fig. 2c, d).
Ecological Niche Modeling
Spearman correlation analysis and the PCA analysis specified
seven climate variables as optimal for identifying the potential
distribution of Ipomoea sagittata: BIO1, Annual mean tem-
perature; BIO5, Max temperature of the warmest month;
B I O 7 , Te m pe r at ur e A nn ua l R an ge ; B I O 8 , M ea n
Temperature of Wettest Quarter; BIO13, Precipitation of
Wettest Month; BIO14, Precipitation of Driest Month;
BIO15, Precipitation Seasonality. These climate variables
were included in Maxent to generate the model based on 42
retrieved localities for I. sagittata with a minimum distance of
1 km between them. Suitable areas for potential distribution
include most of the coast of the Gulf of Mexico, with impor-
tant inland regions, including the Usumacinta and Grijalva
basins, northern Guatemala, Belize, Florida and a small part
of the Chihuahuan Desert. The islands of the Antilles such as
Cuba, Jamaica and the Bahamas were also predicted. In the
Mexican part of the distribution, the addition of the potential
wetland information restricted the distribution to closer to the
coast with a patchy distribution. Larger regions included the
Usumacinta Basin, Papaloapan River Basin and the Laguna
Madre.
Projections for the year 2080 modify the distribution of
Ipomoea sagittata in two ways. The Gulf of Mexico Coast
and Yucatan Peninsula ranges tend to increase inland, while
the northern Chihuahuan Desert distribution decreases its
range. The same pattern is found for both A2 (Fig. 3a, c)
and B2 (Fig. 3b, d) scenarios; however, the changes are more
severe in the A2 scenario.
Wetlands

Fig. 1 Haplotype geographic

distribution for studied
populations of Ipomoea sagittata
(PAZ Poza Azul, MEZ Mezquites,
PAR Port Arthur, RVE Rio Verde,
FRO Frontera, CAM Campeche,
CAL Calkin, MUY Muyil, COB
Cob, CAN Candelaria). Shaded
area shows the potential wetland
distribution model (INEGI, 2012)
and Ipomoea sagittata potential
distribution model in Maxent.
Darker areas correspond to a
clipped model defined by
potential wetland information
(INEGI 2005). Black dots are
localities used to generate the
model

Sea Level Rise Impact
The addition of the rise in sea level to the model reduced the
Gulf of Mexico coast by 23 %. The most affected Protected
Natural Areas were Yum Balam, Ra Lagartos and Contoy
Island that were entirely covered by the sea. The Laguna de
Trminos reserve decreased in size by 83.3 %, Sian Kaan by
67.4 %, Alvarado Lagoon System 46.7 %, Pantanos de Centla
38.2 %, Los Petenes 33.1 %, Celestn 29.5 % and Laguna
Madre 15.9 % (Table 2, Fig. 4).
Discussion
Genetic Variation
Our data exhibited low genetic variation in both chloroplast
and nuclear markers, differing from the elevated number of
nuclear haplotypes found for other Ipomoea, such as
I. imperati and I. pes-caprae, sand dune coastal species with
wider distributions (Cennamo et al. 2013; Miryeganeh et al.
2014). Moreover it has been indicated that in general, aquatic
plants sustain moderate to high levels of genetic variation
within populations, through a combination of founder effects
and rapid local adaptations (De Meester et al. 2002). Plastid
molecular markers in several aquatic plants have detected high
population structure but low variation within populations,
(Dorken and Barrett 2004; Koga et al. 2008; Volkova et al.
2010; Arrigo et al. 2011). Shaw et al. (2005, 2007, 2014) have
identified several non-coding plastid markers with elevated
polymorphic sites for most groups of plants, at the species
and population levels. However, for Ipomoea sagittata these
DNA regions had less than 1 % variation. We propose that
molecular markers like SNPs or microsatellites be used to
corroborate genetic variation in populations of this sand dune
morning glory, and that the number of individuals sampled per
population be increased to at least 20.
We think it is likely that other causes such as ample dis-
persal and clonal reproduction might have acted simulta-
neously, generating the lack of genetic variation found in
I. sagittata. With regard to ample dispersal, it has often been
found that low inter-population and intra-population genetic
diversity in plastid markers may be caused by elevated seed
dispersal via water, wind or animals, which maintains gene
flow among discontinuous populations (Arafeh and
Kadereit 2006). Ipomoea sagittata has a likely adaptation
for water dispersal in that its seeds have hairs that allow
them to float (Austin 2014). In general, seeds in the genus

Fig. 2 Ecological connectivity model for dispersal networks: a current distribution of the cpDNA haplotype, b future A2 scenario for the cpDNA
haplotype, c predicted distribution based on the population connectivity model and d predicted A2 scenario, based on the population connectivity model
Ipomoea are characterized by their hardy, waterproof coat and
an ability to remain viable, in some cases for more than
30 years (Gomes et al. 2012). For Calystegia soldanella, also
in the Convolvulaceae, it has been reported that seeds are able
to stay afloat and germinate following as long as 18 months in
seawater (Ridley 1930; Arafeh and Kadereit 2006). Thus, the
lack of genetic variation might be due to ample dispersal and
gene flow of populations of I. sagittata in the Gulf of Mexico
through dispersion of seeds. However, in this study we found
genetic similarity between coastal and inland populations sep-
arated by more than 800Km, mountain ranges (>3000 m a.s.l.)
and general lack of available wetlands to connect populations.
The most common haplotype was found throughout most of
the distribution range and the population expansion test may
indicate a rapid population expansion (Tajimas D=1.70)
(Avise 2000). However, this pattern might be also attributable
to ancestral polymorphisms, including a low mutation rate
(Schaal et al. 1998; Avise 2000).
Another possible explanation for the low genetic diversity
is related to clonal reproduction in Ipomoea sagittata. It is
common for aquatic plants to reproduce by stolons or rhizome
fragmentation and this type of reproduction promotes the sur-
vival of stress-tolerant clones and results in low levels of ge-
netic variation among populations (Santamara 2002).
Wetlands
Moreover, these rhizome fragments can be dispersed long
distances by water. We suggest that, rather than resulting from
contemporary gene flow, the shared haplotypes found in the
Chihuahuan Desert and the Gulf of Mexico coastal wetland
populations may be indicative of historical colonization when
more humid conditions prevailed in the past.
Sedimentological, geochemical, and stratigraphic studies of
lacustrine sequences in lakes from the north of Mexico indi-
cate that a gradual drying occurred, reaching a maximum
a r o u n d 6 0 0 0 B P, d u r i n g t h e m i d d l e H o l o c e n e
(Ortega.Ramirez et al. 1998)
Low genetic variation, high dispersal ability and clonal
reproduction match a pattern suggesting that Ipomoea
sagittata behaves like a metapopulation; a model that requires
a high turnover rate and produces continuous bottlenecks that
deplete genetic variability (Harrison and Hastings 1996). A
series of local extinctions and recolonizations could be ex-
plained by wetland dynamics related to climate cycles of dis-
turbance (e.g., hurricanes, seasonal flooding, wrack deposi-
tion, and fire) (Flynn et al. 1995; Guntenspergen et al.
1995). The same lack of genetic variation has been reported
for Ipomoea hederacea, a species with similar turnover char-
acteristics (Campitelli and Stinchcombe 2014). Nonetheless,
for I. imperati, a coastal sand dune species, several nuclear
Wetlands

Fig. 3 Prediction model for the

potential distribution of Ipomoea
sagittata for the year 2080: A2
future scenario (a, b) and B2
scenario (c, d)

haplotypes were found, corresponding to ITS sequences
which we found to be monomorphic (Cennamo et al. 2013).
Distributional Changes and Ecological Connectivity
Most of the variables predicting the potential distribution of
Ipomoea sagittata are related to temperature: BIO1, Annual
Mean Temperature; BIO2, Mean Diurnal Range; BIO3,
Isothermality; BIO5, Max Temperature of the Warmest
Month; BIO10, Mean Temperature of the Warmest Quarter
and BIO12, Annual Precipitation. The latter has been recog-
nized as the main climate factor affecting plants in aquatic
ecosystems (Lacoul and Freedman 2006).
Also, it has been reported for other plant species that their
response to climate change is not equal throughout their entire
distribution (Sork et al. 2010; Temunovi et al. 2013). In
 Wetlands

Table 2 Protected Natural Areas of wetlands on the Gulf of Mexico. increase in sea level our results match those of Greaver and
Current area, area that will be covered according to a 1 m sea level rise
Sternberg (2010) who predicted a landward shift by dune
scenario and for each PNA, the percentage of area that will be covered by
the sea species, and those of Feagin et al. (2010) who reported a
positive impact for an increased sea level when the increase
Protected Natural Area Land Area Future area Percent of was moderate (0.18 m by 2095) for the salt marshes of Texas.
(ha) covered (ha) original area (%)
However, when the increase in sea level was higher, they
Alvarado Lagoon System 275,341 128,601 46.7 found this would have a negative impact on the distribution
Isla Contoy 5126 5126 100 of salt marshes.
Laguna de Trminos 705,516 585,578.28 83.3 Our results of the least-cost path analyses reveal a change
Laguna Madre 307,894 46,184.10 15.9 in the genetic connectivity of Ipomoea populations as a result
Los Petenes 282,587 93,253.71 33.1 of global climate change. Sea invasion near the lower Laguna
Pantanos de Centla 302,706 115,028.28 38.2 Madre Basin and the Usumacinta-Grijalva Basin has major
Ria Celestn 81,482 23,629.78 29.5 effects, as does the overall sea intrusion along the coast of
Ria Lagartos 60,348 60,348.00 100 the Yucatan Peninsula. These last two greatly reduce connec-
Sian Kaan 528,148 353,859.16 67.4 tivity and isolate the I. sagittata populations of the northwest-
Yum Balam 154,052 154,052.00 100.00 ern coast. In contrast, the southern Yucatan Peninsula distri-
bution increased in connectivity due to distribution expansion
creating a new corridor. We obtained a model where the inland
Chihuahuan Desert distribution has an intermediate degree of
Ipomoea sagittata, our distribution model for future scenarios connectivity; a pattern that seems mostly unaffected by future
found an overall expansion, however, the coastal range in- climate change aside from a small decrease in area. The same
creased while the northern inland range contracted and this pattern was found for the haplotype and pairwise population
pattern has been found for other tropical and Mediterranean analyses. Over-prediction of the current and future potential
species (Fitzpatrick et al. 2008; Temunovi et al. 2013). The distribution was identified by the projection of areas where
decrease for the Chihuahuan Desert distribution might be re- Ipomoea sagittata has not been collected. This result may be
lated to the forecasted decrease in rainfall (IPCC, Special explained by the fact that many ecological factors can influ-
Report on Emissions Scenarios, 2001). The increase in distri- ence the establishment of aquatic plants, among them, chem-
bution may be the result of a rise in global temperature, which ical components in the water are the most determinant
increases habitat suitability for this warm climate species. Our (Rabalais 2002; Lacoul and Freedman 2006).
results contrast those of Mendoza-Gonzlez et al. (2013) The expected stable distribution of Ipomoea sagittata
where the distribution of all of the dune plant species from shows that populations will probably decline in the coastal
the Gulf of Mexico decreased, however they based their future estuarine wetlands of the Gulf of Mexico as a result of
ecological modeling on six species. For the scenario with the saltwater intrusion and loss of connectivity, but overall, the

Fig. 4 Potential distribution

model for Ipomoea sagittata in
Maxent, for the year 2080, A2
scenario (green). Model for a 1 m
rise in sea level (blue). Water
intrusion into protected areas
(red). Dotted polygons are
protected areas
Wetlands
distribution will increase by ~30 %, and this may con-
strain the future range to a fragmented distribution charac-
terized by increased competition for new zones (Slocum
and Mendelssohn 2008). This increase does not take into
account a vast transitional zone where habitat availability
for fresh water species will decrease because of increased
erosion and salinity (Craft et al. 2009; Li et al. 2009;
Glick et al. 2013). Additionally, a factor that may inhibit
the expansion of I. sagittatas populations is its low toler-
ance to natural phenomena such as hurricanes, which are
likely to increase in frequency due to global climate
change (Mulholland et al. 1997).
Wetland Conservation in the Gulf of Mexico
During the fieldwork for this study we found that previ-
ously reported localities of Ipomoea sagittata have been
subjected to anthropogenic disturbance and this species
has most probably been locally eradicated from the wet-
lands of the central region of the Gulf of Mexico. This is
an exceptionally well-studied region where current urban
development and agricultural activities have had a tremen-
dous impact on wetlands (Lpez-Rosas et al. 2006;
Siemens et al. 2006; Moreno-Casasola 2008; Vzquez-
Gonzlez et al. 2014). In this region, instead of
I. sagittata, we found a related species with contrasting
ecological characteristics: I. carnea subsp. fistulosa (Mart.
ex Choisy) D.F. Austin, another wetland morning glory
with a broader distribution that grows in natural and an-
thropogenically disturbed habitats, alongside seasonally
flooded areas, and is often considered a noxious weed
Fig. 5 a. Ipomoea carnea subsp. fistulosa potential distribution model in
Maxent. Darker areas correspond to a clipped model defined by potential
wetland information (INEGI 2005). White spots are localities used to
generate the model. b. Prediction model for the potential distribution of
since it is poisonous to cattle (Haase 1999). To further
understand whether the future connectivity of populations
of this species would be affected by climate change in the
same way as I. sagittata (a species that is exclusive to
conserved wetlands), we applied the same methods to gen-
erate future ecological niche models and ecological con-
nectivity models for I. carnea subsp. fistulosa, a plant that
is better adapted to disturbed areas.
Optimal variables for I. carnea subsp. fistulosa were relat-
ed to precipitation, which has already been reported as a major
factor in the establishment of this species on seasonally
flooded wetlands (Haase 1999). Our results indicate that
I. carnea subsp. fistulosa maintains a similar pattern of distri-
bution in both present and future models, with a high degree of
correspondence with potential wetland distribution models
(Fig. 5). The two species had contrasting patterns in the loss
and gain of their distribution areas, therefore areas of ecolog-
ical stability where several species can be considered for con-
servation should be selected with caution, since even for phy-
logenetically and ecologically similar species the conse-
quences of global climate change are different (Hannah et al.
2007). I. carnea subsp. fistulosas broader distribution may be
related to a wider spectrum of adaptation to climate variables,
which have been selected by its affinity to early successional
communities. This high degree of adaptability of I. carnea
subsp. fistulosa to a changing environment, its high clonal
capability, abiotic dispersal and an ecological symbiosis with
ants make it likely to become a dominant ecosystem species
(Haase 1999; Rahel et al. 2008). Therefore, in future niche
models comparing I. sagittata and I. carnea subsp. fistulosa,
the latter is less affected under a climate warming scenario.
Ipomoea carnea subsp. fistulosa in Maxent for the year 2080, A2 future
scenario (IPCC 2007). Green: Stable habitat, red: Habitat loss, purple:
Habitat gain

Warmer conditions in the future may favor dominant species
at the cost of species richness (Houlahan and Findlay 2004;
Vander Zanden and Olden 2008).
Our results of genetic diversity and ecological connectivity
are difficult to reconcile since they address different spatial
and temporal scales. However, they offer a small glimpse into
future problems for wetland viability and conservation. Niche
modeling and connectivity assessment for future climate
change increasingly provide useful information for conserva-
tion. Nevertheless, predictive tools cannot yet fully integrate
the various determinants of species distributions, complicating
the transition from forecasting to decision-making (Hannah
et al. 2002; Thuiller et al. 2008; Alagador et al. 2014).
Based on future climate change models, there have been sev-
eral proposals about how to deal with climate change (e.g.,
multiple species modeling, dynamic protected areas, assisted
relocation, ex-situ conservation, etc.) (Hannah et al. 2007;
Sgr et al. 2011; Sax et al. 2013; Shoo et al. 2013), each with
its own setbacks and advantages (Minteer and Collins 2010).
It is evident that climate change will have major effects on
species distributions and change our classical views of
protected areas. Our predictions for the wetlands of the
Yucatan Peninsula suggest GCC will have major effects on
the Yum Balam, Ria Lagartos and Contoy Island NPAs which
may become entirely covered by sea water. In addition, salt-
water intrusion will probably decrease the range of the popu-
lations of Laguna de Trminos reserve by 83.3 %, Sian Kaan
67.4 %, Alvarado Lagoon System 46.7 %, Pantanos de Centla
38.2 %, Los Petenes 33.1 %, Celestn 29.5 % and Laguna
Madre 15.9 %. All of these reserves are located in the potential
distribution area of Ipomoea sagittata. The unique haplotypes
we found in the Sian Kaan and Los Petenes-Celestn reserves
might be lost. The inland distribution of Ipomoea sagittata
mainly lies in the Cuatro Cinegas Basin, an isolated region
of discontinuous wetlands in the middle of the Chihuahuan
Desert distinguished by high levels of endemicity (Souza et al.
2006). This region has been devastated for more than 50 years
through the abuse of water bodies, extraction from gypsum
deposits and tourism (Contreras-Balderas 1984). Therefore
this will be an increasingly important area for conserving
I. sagittata.
Conclusions
We found a lack of genetic structure in Mexican populations
of I. sagittata. A single haplotype is the most frequent and
occurs in most populations, however, our DNA markers are
not conclusive so an assessment using a different set of
markers could provide better hypothesis for historical popula-
tion evolution and future viability. We did find, however, that
climate change will have a decisive impact on I. sagittatas
future distribution. Inland, in the Chihuahuan Desert its
Wetlands
distribution will be greatly reduced, while it seems that its
coastal distribution will increase and thus conservation areas
should be planned for this area. The protected areas most
affected by the rise in sea level, such as Yum Balam, Ria
Lagartos and Contoy Island, will be completely submerged,
while others (e.g., Celestn, Los Petenes, Laguna de
Trminos, Pantanos de Centla) will be partially covered by
the new expanded distribution of I. sagittata. New problems
will arise as a result of its encounter with anthropogenic ac-
tivities and competition to establish populations in new sites.
Moreover, future changes in distribution and sea level will
diminish genetic flow and landscape connectivity which could
isolate populations of Ipomoea sagittata and reduce its ability
to recolonize. Other effects of global climate change, includ-
ing for example an increase in the frequency of climatological
phenomena like hurricanes, will probably negatively impact
this salt marsh morning glory as it inhabits undisturbed
environments.
Acknowledgments We thank Andrew P. Vovides, Antonio Hernndez,
Carla Gutirrez and Dulce Infante whose reviews significantly improved
this manuscript. We are grateful to Diego Angulo, Ruth Delgado-Dvila,
Eduardo Ruz-Snchez, Andrs Ortz-Rodrguez and Ismael Valdivieso
for assistance in the field. We also thank Arith Prez and Cristina
Brcenas for their help in the laboratory and Anna Armitage for providing
samples. Laboratory and fieldwork were supported by a grant from
CONACyT (106451) to P.M.C. and G.H.R. gratefully acknowledges a
graduate studies scholarship from CONACyT (322444). With this study
G.H.R. has fulfilled his Master of Science degree requirements at the
Instituto de Ecologa, A. C.
Appendix 1
G. Huerta-Ramos 67, Poza Azul, Coahuila (XAL), G. Huerta-
Ramos 68, Rio verde, San Luis Potos (XAL), G. Huerta-
Ramos 69, Muyil, Quintana Roo, (XAL), G. Huerta-Ramos
72, Frontera, Tabasco, (XAL), G. Huerta-Ramos 73, San
Francisco Kobn, Campeche (XAL), G. Huerta-Ramos 74,
Candelaria, Campeche, Paul C. Keen S/N, Port Arthur,
Texas (TAES).
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