Content uploaded by Federico Mosquera Guerra
Author content
All content in this area was uploaded by Federico Mosquera Guerra on Apr 26, 2018
Content may be subject to copyright.
!
1!
Analysis of distribution of river dolphins (Inia and
Sotalia) in protected and transformed areas in the
Amazon and Orinoco basins
!
Federico Mosquera-Guerra 1,2, Fernando Trujillo1, Danni Parks3, Marcelo Oliveira-da-
Costa4, Miriam Marmontel5, Dolors Armenteras-Pascual2, Saulo Usma4, Daphne Willems4,
Juan David Carvajal-Castro6, Hugo Mantilla-Meluk6, Nicole Franco1, Diego Amorocho4,
Roberto Maldonado4, Karina Berg4, Lila Sainz4, Paul A. Van Damme7, Elizabeth Cambell8.!
!
1 Fundación Omacha: Calle 84 No. 21 – 64 - Barrio el Polo - Bogotá, D.C,
Colombia. federico.mosqueraguerra@gmail.com!
!
2 Grupo de Ecología del Paisaje y Modelación de Ecosistemas-ECOLMOD: Departamento
de Biología, Universidad Nacional de Colombia - Cra 30 No. 45-03 - Bogotá D.C,
Colombia.!
!
3 Whitley Fund for Nature : 110 Princedale Road l - London W11 4NH, United Kingdom.!
!
4 WWF – Brazil; Bolivia; Colombia; Ecuador, Peru, Netherlands; Deutschland; United
Kingdom. Rue Mauverney, 28 1196 Gland, Switzerland
!
5 Mamiraguá: Estrada do Bexiga, 2.584 Bairro Fonte Boa Cx. Postal 38 69.553-225 – Tefé
(AM), Brasil.!
!
6 Universidad del Quindiìo: Programa de Biologiìa, Carrera 15 No. 12 Norte. Quindiìo -
Armenia - Colombia.!
!
7 Faunagua: Av. Max Fernández final s/n - Plazuela del Chillijchi (Arocagua) –
Cochabamba, Bolivia.!
!
8 Prodelphinus: Enrique Palacios 630 – 204, Lima 18 – Miraflores - Lima, Peru.!
!
!
!
Abstract
The South American river dolphins have evolved in the continental aquatic
ecosystems of the Amazon, Grande, Iténez - Mamoré, Araguaia - Tocantis and Orinoco
rivers. The spatial and temporal distribution and the habitat use of these cetaceans do in
these systems, are determined by distinct environmental characteristics such as
precipitation regimes, elevation, productivity and biomass specific to each system. In
addition, geomorphological accidents, such as rapids, have emerged as barriers that
separate dolphin populations, potentially promoting processes of speciation. To date, there
is no comprehensive analysis of river dolphins distribution and representativeness in
protected areas or areas transformed by hydroelectric plants. In the present work, through
!
2!
niche and spatial modeling tools, we research the representativeness of both protected areas
and areas transformed by hydroelectric plants in the Amazon and Orinoco basins in the
distribution of river dolphins (Inia and Sotalia). The models presented here were
constructed using the MaxEnt algorithm through the integration of 35,594 georeferenced
records and 19 environmental variables derived from the Bioclim and Hydroshed database,
which were parameterised in the R programme. A good representation of the distribution of
river dolphins within the protected areas was evidenced, although the limited management
of the aquatic ecosystems inside the protected areas does not guarantee the conservation of
these species. A major threat identified for river dolphins in South America is the loss of
habitat and fragmentation as a result of the construction of hydroelectric dams. We
examined the degree of overlap between the distribution of Inia and Sotalia and
hydroelectric projects in construction, operation and planning phases and provided an initial
quantification of this tensor. Finally, we consider that the cumulative impacts
(fragmentation, regulation of the flood pulse, retention of limiting nutrients and alteration in
the levels of productivity) generated by this type of infrastructure at the macrobasin scale
will exacerbate the level of the threats to the conservation of river dolphins and their
habitats in the Amazon and Orinoco basins.
Key words: Conservation,!distribution,!human!impact,!hydroelectric!dams,!
river dolphins.!
!
!
Introduction
The South American river dolphins are represented by the subspecies and species of
the genus Inia and Sotalia. They are widely distributed in the aquatic ecosystems of the
Amazon, Orinoco, Tocantins basins and Orinoco delta (Caballero et al. 2007; Reeves et al.
2011; Secchi et al. 2012; Carvajal-Castro et al. 2015; Caballero et al. 2017). Throughout
their distribution it has been identified that geological interruptions, particularly rapids, act
as effective barriers separating natural populations of river dolphins and promoting
processes of differentiation between populations (Rice 1998).
To date, three subspecies of Inia geoffrensis are recognized with distribution
throughout the Amazon basin (I. g. geoffrensis) and the Orinoco basin (I. g. humboldtiana)
with a suggested vicarious isolation caused by the formation of the Canal del Casiquiare on
the ecological border between the Colombian Orinoquia and the Venezuelan Amazon (Da
Silva & Martin 2000). I. g. boliviensis seems to have been isolated from I. g. geoffrensis
because of the 400 km of rapids from Porto Velho on the Madeira river to Riberalta on the
Beni river in Bolivia (Rice 1998). Currently the Bolivian bufeo (I. g. boliviensis) occurs in
the basin of the Madeira, Iténez and Grande river basins in Bolivia and Brazil (Trujillo et
al.1999; Banguera- Hinestroza et al. 2002; Ruiz-García et al. 2008; Ruiz-García 2010;
Reeves et al. 2011; 2013). However, the taxonomy for the genus Inia has been discussed
and even a new species has been described: the Araguaia river dolphin (Inia
araguaiaensis), present in the hydrographic complex formed by the Araguaia - Tocantins
rivers (Hrbek et al. 2014).
!
3!
The genus Sotalia includes two species based on cranial morphology and genetic
evidence (Madeira et al. 2004; Cunha et al. 2005; Caballero et al. 2007; 2010; 2017).
Sotalia guianensis (P-J. Van Bénedén 1864) presents populations distributed along the
coasts and estuaries of the Atlantic Ocean from Honduras to Brazil, and in the north of
South America, including individuals established in Lake Maracaibo, middle and delta
parts of the Orinoco river in Venezuela. Recent genetic studies determined that there is a
divergence of 600,000 years between continental and coastal populations (Carvajal-Castro
et al. 2015; Caballero et al. 2017). The species Sotalia fluviatilis (Gervais y Deville, In
Gervais 1853) is found in the Amazon river basin (riparian zone) and is sympatric with Inia
geoffrensis in Brazil, Colombia, Ecuador, Peru and French Guiana in the Amazon river
basin (Borobia et al. 1989; Da Silva 1994; Martin y Da Silva 2004; Cunha et al. 2005;
Caballero et al. 2007; Flores and Da Silva 2009; Trujillo et al. 2010; Secchi et al. 2012;
Carvajal-Castro et al. 2015; Caballero et al. 2017).
Although river dolphins are obligate aquatic mammals, external environment
variables external to this environment associated with the river basins they inhabit, such as
precipitation and atmospheric temperature, may be determining aspects of their ecology,
directly or indirectly affecting their occurrence in a particular area. Historically, the
Orinoco, Amazon and the Araguaia-Tocantins complexes have been identified as different
biogeographical and ecological units characterized by contrasting climatic conditions
(McClain and Naiman 2008; Edmond et al. 1996; WCS 2017). These deferring conditions
affect in many ways the hydric dynamics of their basins, determining the composition and
structure of the biota associated with the course of its main and tributary rivers. Despite
analyses of the potential influence that external variables may have on aquatic biota, to
date, there is no comprehensive analysis in the South American territory on the
representativeness of the distribution of river dolphins in protected areas or areas
transformed by hydroelectric plants.
New tecnological approaches based on Geographic Information Systems (GIS),
currently allow the characterisation of the availability of habitat for species in a given area
through suitability models at a regional scale. Regionally, ecological conditions such as
elevation, temperature and precipitation in combination with latitude, determine, among
others, the vegetation cover, which consequently at a local scale define aspects such as
productivity, nutrient flow and conductivity in the related water ecosystems at a local scale.
Although the relationship between regional and local variables has been studied relatively
poorly, GIS analysis opens a window of possibility to understand the distribution and
ecology of aquatic organisms such as river dolphins at the continental level.
In the present study, the results from river dolphin monitoring, based on sightings
are integrated with environmental information through niche modeling tools, with the
purpose of investigating the representativeness of the different species and subspecies in
protected areas and areas transformed by hydroelectric plants in the Amazon and Orinoco
basins.
!
4!
Material and Methods
Obtaining records and environmental variables
The environmental conditions included 19 bioclimatic variables from the Worldclim
database (Hijmans et al. 2005), in addition, radiation was obtained as an approximation of
primary productivity. To obtain this information, the values of the 12 months of the year
provided by Worldclim v.2 are averaged (Fick and Hijmans 2017). All the variables
presented a resolution of 30 arc-seconds (~1 km2 in Ecuador) (Table 1).
Table 1. Bioclimatic variables used in the potential distribution models of river dolphins
(Inia and Sotalia) in the Amazon and Orinoco river basins.!
Variables
Description
Elevation
Height in meters above sea level
Bio 1
Annual average temperature
Bio 2
Average daytime range (Mean of the month
(Max Temp - Min Temp))
Bio 3
Isothermality ((Bio 2/Bio 7) * 100)
Bio 4
Seasonality of temperature (Standard
deviation * 100)
Bio 5
Maximum temperatura of the hottest month
Bio 6
Minimum temperature of the coldest month
Bio 7
Annual temperature range (Bio 5 - Bio 6)
Bio 8
Average temperature of the wettest quarter
Bio 9
Average temperature of the driest quarter
Bio 10
Average temperature of the warmest quarter
Bio 11
Average temperature of the coldest quarter
Bio 12
Annual rainfall
Bio 13
Precipitation of the wettest month
Bio 14
Precipitation of the driest month
Bio 15
Seasonality of precipitation (Coefficient of
variation)
Bio 16
Precipitation of the wettest quarter
Bio 17
Precipitation of the driest quarter
Bio 18
Precipitation of the warmest quarter
Bio 19
Precipitation of the coldest quarter
Rad
Solar radiation
Potential distribution models
!
5!
The models of potential distribution of the subspecies and species of the genera Inia
and Sotalia were generated using the environmental variables and the georeferenced
records obtained through sightings in expeditions carried out within the framework of the
South American River Dolphin Conservation Programme (2006-2018) and the records
issued by river dolphins tagged with satellite telemetry in Colombia, Brazil and Bolivia
(Table 2, Figures 1& 2). The algorithm used was the maximum entropy (MaxEnt) (Phillips
et al. 2006). MaxEnt was chosen because it has been widely used and its best performance
has been demonstrated to estimate the potential distribution of the species more accurate
than other approaches (Graham and Hijmans 2006; Townsend et al. 2007). As MaxEnt only
works with information on the presence of species, it must generate pseudoausencias, more
appropriately called ‘background’ in order to obtain a calibration of the model (Phillips and
Dudík 2008). Therefore, 10000 randomly generated points (background) were used through
the northern zone of the Neotropic. 10 replicas were generated to perform an assembly
methodology (Araújo and New 2007) and thus obtaining a consensus model among the
replicas.
Table 2. Records of river dolphins (Inia and Sotalia) in the Amazon, Orinoco and
Tocantins-Araguaia river basins.
Taxones
Georeferenced sightings
Records issued by
dolphins marked with
satellite telemetry
I. g. geoffrensis
9.050
7.189
I. g. humboldtiana
4.452
714
I. g. boliviensis
2.388
4575
Inia araguaiaensis
974
Sotalia fluviatilis
5.954
Sotalia guianensis
303
Total
23.121
12.473
!
6!
Figure 1. Georeferenced records for genus Inia in South America.
Figure 2. Georeferenced records for genus Sotalia in South America.
!
7!
To evaluate the performance of the model, a "k fold partitioning" approach was
used, with an area under the receiver curve (AUC) as the evaluation metric, using 75% of
the occurrence data to perform modeling and 25% of the remaining occurrence data to test
the model. The AUC value is used to evaluate the discrimination ability of the model: AUC
values close to 1 are considered as a perfect prediction and values of 0.5 or less reflect that
the model is not better than a random model. These analyses were carried out using the R
software, using the dismo libraries (Hijmans et al. 2015), SDMTools (VanDerWal et al.
2014) and raster.
The consensus model generated by MaxEnt is a probability map, on which the
probability of the presence of species is shown. This probability map was used to make a
binary map representing the presence and absence of dolphins, establishing a threshold
value as a criterion, defined as the maximum sum of specificity and sensitivity (MSS).
Therefore, the probability values (probability map) that were equal to or greater than this
threshold value were considered as demostrating the presence of river dolphins (the value
of the pixel equal to 1); on the other hand, values below this threshold were considered as
species’ absence (pixel value equal to 0).
Representation of protected and transformed areas in the potential distribution of river
dolphins (Inia and Sotalia)
A statistical analysis was carried out overlapping the models of potential
distribution, with the protected areas (WDPA) (UNEP-WCMC 2010) downloaded from
(www.protectedplanet.net) and areas transformed by hydroelectric plants in Brazil, Bolivia,
Colombia, Ecuador, Peru and Venezuela (Anderson et al. 2018, Latrubesse et al. 2017).!The
number of pixels intersecting the distribution of species and protection zones were
quantified as an indirect measure of the ability of protected areas to protect dolphins.!At the
same time, the areas transformed by hydroelectric dam construction, operation and
planning in the Amazon, Orinoco and Araguaia-Tocantins basins were quantified in order
to demonstrate the impact of this type of infrastructure on river dolphin populations. For the
purpose of calculating these areas, the function of the library rgeos was used in the
statistical software R.
Results
Potential distribution models
The generated models demonstrate a high correlation, with AUC values of 0.8852
(I. g. geoffrensis), 0.889 (I. g. humboldtiana), 0.9189 (I. g. boliviensis), 0.9738 (I .
araguaiaensis), 0.948 (Sotalia fluviatilis) and 0.933 (S. guianensis). Figure 3 shows the
modelled potential distrubution of the South American river dolphin species and subspecies
(Inia and Sotalia).
!
8!
!
!
!
!
!
!
!
Figure 3. Modelled potential distrubution of the South American river dolphins (species and
subspecies) Inia and Sotalia, derived from the algoritm MaxEnt. A. I.g. geoffrensis. B. I.g.
humboldtiana. C. I.g.boliviensis. D. I. araguaiaensis. E. Sotalia fluviatilis. F. S. guianensis.
Representation of protected and transformed areas in the potential distribution of river
dolphins (Inia and Sotalia)
A
B
C
D
E
F
!
9!
Potential distribution areas for the South American river dolphins (Inia and Sotalia) and
their representation in the aquatic ecosystems in Amazon protected areas were calculated in
order to establish their representativeness for each taxa. (Table 3). In the category of
protected areas, strict figures were included, such as National Parks and regional parks, and
other figures such as sustainable reserves and Ramsar sites were included.
Table 3. Representativeness of protected areas in the distribution of South American river
dolphins (Inia and Sotalia).
Species
Total&area&
potential&
distribution&
Areas of aquatic ecosystems in conservation km2
Total&area&in&
Conservation&
km2&
Brasil
Bolivia
Colombia
Ecuador
Peru
Venezuela
I.g.geoffrensis
468.717!
69.324
!!
5.839
2.900
11.455
397
89.915 (19,2%)
I.g.humboldtiana
114.962!
!!
!!
2.151
!!
!!
10.634
12.785 (11,1%)
I.g.boliviensis
76.597
5.892
12.494
!!
!!
!!
!!
18.386 (24,0%)
I. araguaiaensis
76.182
11.503
!!
!!
!!
!!
!!
11.503 (15,1%)
S. fluviatilis
356.716!
54.892
!!
2.637
2.900
7.726
!!
68.155 (19,1%)
S. guianensis
17.473
!!
!!
!!
!!
!!
4.630
4.630 (26.5%)
Figure 4. Distribution of the genus Inia in Amazon protected areas
!
10!
Figure 5. Distribution of of the genus Sotalia in Amazon protected areas.
The areas transformed by hydroelectric plants were calculated in phases of planning,
construction and operation, and the representation of these in the potential distribution areas
of the South American river dolphins (Table 4).
Table 4. Representativeness of transformed areas by hydroelectric plants in the distribution
of South American river dolphins (Inia and Sotalia).
Species
Total&area&
potential&
distribution&
km2
Areas transformed by hydroelectric plants in
different phases km2
Operation
Constrution
Planning
I. g. geoffrensis
468.717
77.077 (16.4%)
68.995 (14.7%)
139.981 (29.9%)
I. g. humboldtiana
114.962
26.348 (22.9%)
6.302 (5.5%)
I.g. boliviensis
76.597
1.482 (1.9%)
I. araguaiaensis
76.182
41.853 (54.9%)
16.005 (21%)
36.281 (47.6%)
Sotalia fluviatilis
356.716
77.077 (21.6%)
68.995 (19.3%)
139.981 (39.2%)
Sotalia guianensis
17.473
2.704 (15.5%)
2.555 (14.6%)
!
11!
Figure 6. Spatialisation of habitats transformed by hydroelectics in the operation phase for
the genus Inia.
Figure 7.!Spatialisation of habitats transformed by hydroelectics in the constrution phase
for the genus Inia.
!
12!
Figure 8. Spatialisation of habitats transformed by hydroelectics in the planning
phase for the genus Inia.
Figure 9. Spatialisation of habitats transformed by hydroelectics in the operation phase for
the genus Sotalia.
!
13!
Figure 10. Spatialisation of habitats transformed by hydroelectics in the constrution phase
for the genus Sotalia.
Figure 11. Spatialisation of habitats transformed by hydroelectics in the planning
phase for the genus Sotalia.
Discussion
Potential distribution models
River dolphins (Inia and Sotalia) occur in the Amazon and Orinoco river basins,
from the deltas upstream, to where impassable rapids, waterfalls, lack of water and possibly
!
14!
low temperatures block their movement (Best and da Silva 1989a, b). For the genus Inia
three geographic populations have been recognised as subspecies: I. g. geoffrensis in the
Amazon basin, except for the Madeira basin in Bolivia upstream, from the Teotonio rapids,
I. g. boliviensis in the upper Madeira basin, and I. g. humboldltiana in the Orinoco basin
(Rice 1998, Reeves et al. 2011). Genetic studies reveal the relevance of these geographical
barriers in the generation of vicariant speciation processes, as is the case with Inia
araguaiaensis, a species recently described, whose populations are isolated by the
Araguaia-Tocantins hydrographic complex (Hrbek et al. 2014).
For I. g. geoffrensis, the distribution generated by the model coincide with the
distribution presented by de Best and da Silva (1989a), (1989b) and Leatherwood (1996),
who place its occurence in the rivers Xingu, Tapajós, Madeira (downstream from the rapids
of Teotonio), Purus, Juruá, Ucayali and Marañón (and Samiria tributary) that flow
generally towards the north, and the Negro river with itstributaries Caquetá (Japurá),
Apaporis, Putumayo (Iça) and Napo.
In relation to the distribution of I. g. humboldtiana, the model outcome matches
what was reported by Pilleri y Gihr (1977); Best y da Silva (1989a), (1989b); Meade and
Koehnken (1991); Velásquez-Roa et al. (2015), identifying the subspecies’ presence in the
Apuré tributaries (Portuguesa and Guanmar), Capanaparo, Cinaruco, Meta, Bita, Vichada,
Tomo, Tuparro, Guaviare (tributary Guayabero), Inírida and Atabapo (and Temi tributary)
that flow to the south and east, and the tributaries Aro, Caura, Parquaza, Ventauri (San Juan
tributary) that flow to the north and west, as well as the Casiquiare Canal, which connects
the Orinoco with the Negro river (a tributary of the Amazon), above and below the two
series of rapids in Puerto Ayacucho, which are the main barriers that are expected to
separate the dolphin populations of the Amazon and the Orinoco (Reeves et al. 2011).
Nevertheless, river dolphins have been seen crossing the first group of rapids in Puerto
Ayacucho (Atures) during the high tide (Fernando Trujillo, personal communication with
B.D. Smith).
The modelled distribution generated for Inia boliviensis coincides with what is
stated by Pilleri and Gihr (1977); Best and da Silva (1993); Aliaga-Rossel et al. (2006); Da
Silva (2009); Aliaga-Rossel (2010); Tavera et al. (2010); Cella-Ribeiro et al. (2013), and
Gravena et al. (2014), that this unique aquatic mammal is present in the sub-basins of the
rivers Abuna, Guaporé, Iténez or Guaporé basin (including Verde and Iporuporé
tributaries), Mamoré basin and its primary and secondary tributaries: Pirai, Grande, Ichilo,
Chapare, Ibaré, Tijamuchi, Apere, Yacuma and Yata, Beni (and Orton tributary); on the
Madeira river, its distribution is considered below the Tetonio rapids located upstream from
Porto Velho. According to Cella-Ribeiro et al. (2013), the Bolivian river dolphin presents
an estimated evolutionary divergence of 3.1 million years of genetic isolation generated by
the geographic barrier formed by a series of 18 rapids along a stretch of 290 kilometers on
the Madeira river. These rapids are thought to act as a barrier to the movement of dolphins
and thus restrict the distribution of I.g.boliviensis upstream of the rapids, and I.g.geoffrensis
downstream of the rapids.!
!
15!
This example is similar to the recent description of the species Inia araguaiaensis
(Hrbek et al. 2014) distributed in the Araguaia-Tocantins river hydrographic complex,
currently isolated from the Amazon river basin (Rossetti and Valeriano 2007; Goulding et
al. 2003). The two basins have been disconnected, although not completely isolated from
each other in the transition from Pliocene to Pleistocene (Rossetti and Valeriano 2007).
Currently, only a narrow channel west of the island of Marajo joins the delta of the Amazon
river with the Para river, which drains the Araguaia-Tocantins river. The connectivity
between the Araguaia-Tocantins and Amazon basins is restricted by a series of important
rapids in the lower Tocantins river, becoming an important biogeographical element to
generate speciation processes (Hrbek et al. 2014).
The results obtained in relation to the distribution of S. fluviatilis coincide with the
reports generated by Layne (1958); Obregón et al. (1988); Borobia et al. (1991); Trujillo
(1992), (1994a), (2000); Vidal et al. (1997); McGuiere and Henningsen (2007); Gómez-
Salazar et al. (2010); Carvajal-Castro et al. (2015) and Caballero et al. (2017). All report its
presence in the Amazon basin as far inland as southern Peru, eastern Ecuador and
southeastern Colombia. This presents a sympatric distribution with I. g.geoffrensis (Secchi
2012), although S. fluviatilis does not occur in the Beni/ Mamoré river basin ,in Bolivia nor
in the upper part of the Negro river (Flores y da Silva 2009). The results obtained by
modeling identy limitations for the distribution of S. fluviatilis in the Caqueta/Japura river
basin, determined by the rapid of Cordoba and reported by Trujillo (1994b); Trujillo
(1995), Galindo (1997); Trujillo et al. (2006); Gómez-Salazar et al. (2010); Pavanato et al.
(2014) and Caballero et al. (2017); in the Apaporis river, its distribution is limitated by the
rapids of Estrella and Puerco (Gómez-Salazar et al. 2010).
In relation to S. guianensis, the presence of the species in the Colombian Orinoco is
not corroborated, which suggests that its distribution is restricted to the middle basin and
delta of the Orinoco. This can be explained by the rapids of Maipures, Atures and the Dead
between the Ayacucho and Samariapo Ports in the Venezuelan Orinoco, probably
restricting the species’ distribution (Gómez-Salazar et al. 2010; Herrera-Trujillo 2012).
Representation of protected and transformed areas in the potential distribution of river
dolphins (Inia and Sotalia)
In the Amazon and Orinoco basins there are important areas under conservation
management, such as national natural parks and Ramsar sites. Among these, there are areas
of distribution of river dolphins (Trujillo et al. 2014). However, the remoteness of these
areas, the limited resources for their control and surveillance, the constant threat of
extractive economies such as gold mining, deforestation of riparian forest, commercial
fishing and directed catches, as well as the terrestrial approach of the initiatives of
conservation initiatives, make the management of aquatic ecosystems within protected
areas still limited (Mosquera-Guerra et al. 2015).
The transformation of the heterogeneous and complex aquatic ecosystems in the
Amazon through the construction of more than one hundred hydroelectric plants causes the
ecological homogenisation of these systems, changes in the flood pulses downstream of the
!
16!
dams and the retention of limiting nutrients for primary productivity in aquatic food webs
such as the N and P (Forsberg et al. 2017; Latrubesse et al. 2017; Anderson et al. 2018).
It also causes significant losses in fluvial connectivity exactly where the greatest diversity
of river dolphins and fish on the planet are found (Mosquera-Guerra et al. 2015; Anderson
et al. 2018).!These tensors have caused the transformation of extensive areas occupied by
river dolphins in South America. Arauho and Wang (2014), report the effects on
populations of I. araguaiaensis due to the presence of seven hydroelectric projects
constructed in their distribution area and for I.g. boliviensis, I.g. geoffrensis and Sotalia
fluviatilis are reported a total of three new ones.
!These effects are not only limited to the fragmentation of complex and
heterogeneous aquatic systems such as the Araguaia, Tocantins and Madeira rivers. These
ecosystems are made up of habitats types essential for feeding, reproduction and refuge of
dolphins such as confluences, tributaries, lakes and main rivers and tributaries with
different origins, Andean in the case of white waters, jungle in the black and clear rivers
influenced by the Brazil shield
Another type of cumulative negative effect at the macrobasin scale is reported by
Forsberg et al. (2017), which predicts that upon entry into operation of the Rositas dams,
Angosto Del Bala, Inambari, Tam 40, Pongo De Aguierre and Pongo De Monseriche, will
reduce the supply of sediment by 69%, phosphorus by 67% and nitrogen by 57% in the
Andean region and in the entire Amazon basin by 64, 51 and 23%, respectively. These
large reductions in the supply of sediments and nutrients will have a great impact on the
geomorphology of the channels, the fertility of the floodplains and the aquatic productivity
(Latrubesse et al. 2017).
These effects will be greater near the dams and will extend to the lowland flood
plains (Latrubesse et al. 2017). It is expected that the attenuation of the downstream flood
pulse will alter the survival, phenology and growth of the floodplain vegetation and reduce
the primary and secondary productivity of the floodplain food source for fish in its different
phases. This reduction in the biomass of these systems could potentially exacerbate the
conflicts between the river dolphins and the artisanal and commercial fisheries due to the
decrease of fishing resource in the Amazon.
When analyzing the representativeness of protected areas, it is evident that in the
case of Inia boliviensis and Inia araguiaensis it is relatively low. That is why it is advisable
to encourage the creation of aquatic ecosystem conservation figures that generate protection
mechanisms for these species. Ramsar sites with implemented management plans can be
efficient and suitable for river dolphins.
Acknowledgments
This research was conducted as part of the South America River Dolphins
Conservation Programme sponsored by Whitley Found for Nature, Foundation Segre,
WWF, Nature Serve and Colciencias. The authors thank the researchers and local and
national authorities who participated in the 28 expeditions carried out in South America and
satellite telemetry project to study the ecology of the movement of river dolphins in the
Amazon and Orinoco.
!
17!
References
Aliaga-Rossel, E., McGuire, T. L. and Hamilton, H. 2006. Distribution and encounter rates
of the river dolphin (Inia geoffrensis boliviensis) in the central Bolivian Amazon. Journal of
Cetacean Research Management 8(1): 87–92.
Aliaga-Rossel, E. and T. L. McGuire. 2010. Iniidae. pp. 535-549. In: Wallace, R.B., H.
Gómez, Z.R. Porcel & D.I. Rumiz (eds.) Distribución, Ecología y Conservación de los
Mamíferos Medianos y Grandes de Bolivia. Centro de Ecología y Difusión Simón I. Patiño,
Santa Cruz.
Anderson, E. P., Jenkins, C. N., Heilpern, S., Maldonado-Ocampo, J. A., Carvajal-Vallejos,
F. M., Encalada, A. C., Rivadeneira, J. F., Hidalgo, M., Cañas, C. M., Ortega, H., Salcedo,
N., Maldonado, M. 2018. Tedesco, Fragmentation of Andes-to-Amazon connectivity by
hydropower dams. Sci. Adv. 4, eaao1642.
Arauho, C and Wang, J. 2014. The dammed river dolphins of Brazil: impacts and
conservation. Fauna & Flora International, Oryx, 49(1), 17–24
doi:10.1017/S0030605314000362
Araujo, MB, New M. 2007. Ensemble forecasting of species distributions. Trends Ecol.
Evol. 22: 42–47.
Banguera-Hinestroza, E., Cárdenas, H., Ruiz- García, M., Marmontel, M., Gaitán, E.,
Vázquez, R., and García-Vallejo, F. 2002. Molecular identification of evolutionarily
significant un its in the Amazon River dolphin Inia sp. (Cetacea:Iniidae). Journal of
Heredity 93:312–322.
Best, R.C, da Silva VMF (1993). Inia geoffrensis. Mamm Species 426:1–8.
Best, R. C. and da Silva, V. M. F. 1989a. Amazon River dolphin, boto Inia geoffrensis (de
Blainville, 1817). In: S. H. Ridgway and R. Harrison (eds), Handbook of marine mammals,
Vol. 4: River dolphins and the larger toothed whales, pp. 1-24. Academic Press.
Best, R. C. and Da Silva, V. M. F. 1989b. Biology, status and conservation of Inia
geoffrensis in the Amazon and Orinoco river basins. In: W. F. Perrin, R. L. Brownell, K.
Zhou and Lu Jiankang (eds), Biology and conservation of the river dolphins, pp. 23-34.
IUCN Species Survival Commission.
Borobia, M., Siciliano, S., Lodi, L., and Hoek, W.1991. Distribution of the South American
dolphin Sotalia. Canadian Journal of Zoology 69 (4): 1025-1039 pp. http: / /
dx.doi.org/10.1139/z
Bivand R, Rundel C. 2014. rgeos: Interface to Geometry Engine–Open Source (GEOS). R
Package.
Caballero, S., Trujillo, F., Vianna, J.A., Barrios-Garrido, H., Montiel, M.G., Beltran-
Pedreros, S., Marmontel, M., Santos, M.C.O., Rossi-Santos, M.R., Santos, F.R. and Baker,
!
18!
C.S. 2007.Taxonomic status of the genus Sotalia: species level ranking for 'tucuxi' (Sotalia
fluviatilis) and 'costero' (Sotalia guianensis) dolphins. Marine Mammal Science 23(2): 358-
386. http://dx.doi.org/10.1111/j.1748-7692.2007.00110.x
Caballero, S., Trujillo, F., Vianna, J.A., Barrios-Garrido, H., Montiel, M.G., Beltrán
Pedreros, S., Marmontel, M., Santos, M.C.O., Rossi-Santos, M.R., Santos, F.R and Baker,
C.S. 2010. Mitochondrial DNA diversity, differentiation and phylogeography of the South
American riverine and coastal dolphins Sotalia fluviatilis and Sotalia guianensis. Latin
American Journal of Aquatic Mammals 8(1-2): 69-79.
http://dx.doi.org/10.5597/lajam00155
Caballero, S., Trujillo, F., Del Risco, A., Herrera, O., and Ferrer, A. 2017. Genetic identity
of Sotalia dolphins from the Orinoco River. Marine Mammal Science 33(4): 1214-1223.
http://dx.doi.org/10.1111/mms.12422
Carvajal-Castro, J.D., Velásquez-Roa, T., Mantilla-Meluk, H., Trujillo, F. and Mosquera-
Guerra, F. 2015. Modelación de nicho y aspectos biogeográfícos del género Sotalia
(Cetartiodactyla: Delphinidae) en los ríos Amazonas y Orinoco, Colombia. Momentos de
Ciencia. 12 (2). 121-125.
Cella-Ribeiro, A., G. Torrente-Vilara., D.B. Hungria., Oliveira M de. 2013. As corredeiras
do Rio Madeira. Peixes do Rio Madeira. pp 47–53.
Cunha, H.A., Da Silva, V.M.F., Laílson-Brito, J.J., Santos M.C.O., Flores, P.A.C., Martin,
A.R., Azevedo, A.F., Fragoso A.B.L., Zanelatto, R.C. and Solé-Cava, A.M. 2005. Riverine
and marine ecotypes of Sotalia fluviatilis are different species Marine Biology 148(2): 449-
457.http://dx.doi.org/10.1007/s00227- 005-0078-2
Da Silva V.M.F. 2009. Amazon River Dolphin (Inia geoffrensis). In: Perrin WF, Wursig B,
Thewissen JGM (eds) Encycl. Mar. Mamm. Academic Press, London, UK, pp 18–20.
Da Silva V. and A.R. Martin. 2000. The status of the boto or Amazon River dolphin Inia
geoffrensis (de Blainville, 1817): a review of available information. Paper submitted at the
meeting of the IWC Scientific Committee, 2000. IWC, Cambridge, UK.
Da Silva, V. M. F. and Best, R. C. 1994. Tucuxi Sotalia fluviatilis (Gervais, 1853). In: S. H.
Ridgway and R. Harrison (eds), Handbook of marine mammals, Volume 5 The first book
of dolphins. 43-69. Academic Press, London, UK.
Edmond J.M., Palmer, M.R., Measures, C.I., Brown, E.T. and Huh, Y. 1996. Fluvial
geochemistry of the eastern slope of the northeastern Andes and its foredeep in the drainage
of the Orinoco in Colombia and Venezuela. Geochimica et Cosmochimica Acta 60: 2949–
2976.
Fick, S.E., Hijmans, R.J. 2017. WorldClim 2: new 1‐km spatial resolution climate surfaces
for global land areas. International Journal of Climatology, 37(12), 4302-4315.
Flores, P.A.C. and Da Silva V.M.F. 2009. Tucuxi and Guiana Dolphin - Sotalia fluviatilis
and S. guianensis. 1188-1191pp. En: Perrin, W.F., Würsig, B. y Thewissen, J.G.M. (Eds)
Encyclopedia of Marine Mammals. 2nd ed. Academic Press, Amsterdam, Netherlands.
!
19!
Forsberg, B.R., Melack, J.M., Dunne, T., Barthem, R.B., Goulding, M., Paiva, R.C.D., et
al. 2017. The potential impact of new Andean dams on Amazon fluvial ecosystems. PLoS
ONE 12(8): e0182254. https://doi.org/10.1371/journal. pone.0182254
Galindo, A.1997. Ecología y abundancia de los delfines de río Inia geoffrensis y Sotalia
fluviatilis en el río Caquetá, área de influencia de la Pedrera, Colombia. B.Sc.Thesis.
Universidad del Valle. Cali. Colombia. 77 pp.
Goulding, M., Barthem, R.B., Ferreira, E.J.G. 2003. The Smithsonian Atlas of the Amazon.
Washington, DC: Smithsonian Institution Press.
Gómez Salazar, C., Portocarrero-Aya, M., Caballero, S., Bolaños-Jiménez, J., Utreras, V.,
McGuire, T., Ferrer-Pérez, A., Pool, M., and Aliaga-Rossel, E. 2010. Update on the
freshwater distribution of Sotalia in Colombia, Ecuador, Peru, Venezuela and Suriname.
LAJAM 8(1-2): 171-178 pp. E-ISSN 2236-1057 / ISSN 1676-7497
Graham, C., Hijmans, R.J. 2006. A comparison of methods for mapping species ranges and
species richness. Global Ecology and Biogeography, 578–587.
Gravena, W., Izeni, P.F., Da Silva, M.N.F., Da Silva, V.M.F., and Hrbek, T. 2014.
Looking to past and the future: were the Madeira River rapids a geographical barrier to the
boto (Cetacea: Iniidae)?. Conserv Genet. 15:619-629 pp. DOI 10.1007/S10592-014-0565-4
Herrera-Trujillo, O. 2012. Estatus de los delfines de río Sotalia sp. e Inia geoffrensis en la
cuenca del Orinoco. Tesis para optar al título de Máster Universitario en Biodiversidad en
Áreas Tropicales y su Conservación. Universidad Internacional de Melendéz-Pelayo. Quito
(Ecuador). 99 pp.
Hijmans, R.J., Cameron, S.E., Parra, J.L., Jones, P.G., Jarvis, A. 2005. Very high resolution
interpolated climate surfaces for global land areas.International journal of climatology,
25(15): 1965-1978.
Hijmans R.J. 2015. raster: Geographic Data Analysis and Modeling. R package version 2.4-
18. ver http://CRAN.R-project.org/package=raster
Hijmans, R.J., Phillips, S., Leathwick, J., Elith, J. 2015. dismo: Species Distribution
Modeling. R package version 1.0-12. ver http://CRAN.R-project.org/package=dismo
Hrbek, T., da Silva, V.M.F., Dutra, N., Gravena, W., Martin, A.R. and Farias, I.P. 2014. A
new species of river dolphin from Brazil or: how little do we know our biodiversity. PLoS
ONE 9(1): e83623. doi:10.1371/journal.pone.0083623
Latrubesse, E. M., E. Y. Arima, T. Dunne, T., .et al. 2017. Damming the rivers of the
Amazon basin. Nature 546(7658):363-369.
Layne, J. 1958. Observations on freshwater dolphins in the upper Amazon. Journal of
Mammalogy 39(1): 1-22. http:/ / dx.doi.org/10.2307/ 1376605
!
20!
Leatherwood, S. 1996. Distributional ecology and conservation status of river dolphins
(Inia geoffrensis and Sotalia fluviatilis) in portions of the Peruvian Amazon. Thesis, Texas
A&M University.
Madeira Di Beneditto, A.P. and Arruda Ramos, R.M. 2004. Biology of the marine tucuxi
dolphin (Sotalia fuviatilis) in south-eastern Brazil. J. Mar. Biol. Ass. U.K. 84,1245-1250.
Martin, A.R., Da Silva, V.M.F. and Salmon, D.L. 2004. Riverine habitat preferences of
botos (Inia geoffrensis) and tucuxis (Sotalia fluviatilis) in the Central Amazon. Marine
Mammal Science 20: 189-200.
Meade, R. H. and Koehnken, L. 1991. Distribution of the river dolphin, tonina Inia
geoffrensis, in the Orinoco river basin of Venezuela and Colombia. Interciencia 16: 300-
312.
McClain M.E. and R.J. Naiman. 2008. Andean Influences on the Biogeochemistry and
Ecology of the Amazon River. Oxford Journals. Oxford University Press. BioScience, Vol.
58, No. 4: 325-338.
McGuire, T.L., and Henningsen, T. 2007. Movement Patterns and Site Fidelity of River
Dolphins (Inia geoffrensis and Sotalia fluviatilis) in the Peruvian Amazon as Determined
by Photo-Identification. Aquatics Mammals. 33(3), 359-367 pp. DOI:
10.1578/AM.33.3.2007.359
Mosquera-Guerra, F., Trujillo, F., Diazgranados, M. C. and Mantilla-Meluk, H.. 2015.
Conservación de delfines de río (Inia geoffrensis y Sotalia fluviatilis) en los ecosistemas
acuáticos de la Amazonia y Orinoquia en Colombia. Momentos de Ciencia 12(2):77-86.
Obregón, C., Torres, F., and Trujillo, F. 1988. Colombian dolphins. Whalewatcher 22:21
pp.
Pavanato, H.J., Melo-Santos, G., Lima, D.S., Portocarrero-Aya, M., Paschoalini, M.,
Mosquera, F., Trujillo, F., Meneses, R., Marmontel, M., Maretti, C. 2016. Risks of dam
construction for South American river dolphins: a case study of the Tapajós River.
Endangered Species Research Endang Species Res. Vol. 31: 47–60, 2016 doi:
10.3354/esr00751
Pilleri, G. and Gihr, M. 1977. Observations on the Bolivian (Inia geoffrensis d'Orbigny,
1834) and the Amazonian bufeo (Inia geoffrensis de Blainville, 1817), with a description of
a new subspecies (Inia geoffrensis humboldtiana). Investigations on Cetacea 8: 11-76.
Phillips, S.J., Anderson, R.P., Schapire, R.E. 2006. Maximum entropy modeling of species
geographic distributions. Ecological Modelling. 190: 231–259.
Phillips, S.J., Dudík, M. 2008. Modeling of species distributions with Maxent: new
extensions and a comprehensive evaluation. Ecography. 31(2): 161-175.
Reeves, R.R., Jefferson, T.A., Karczmarski, L., Laidre, K., O’Corry-Crowe, G., Rojas-
Bracho, L., Secchi, E.R., Slooten, E., Smith, B.D., Wang, J.Y. and Zhou, K. 2011. Inia
!
21!
geoffrensis. The IUCN Red List of Threatened Species 2011: e.T10831A3220342.
http://dx.doi.org/10.2305/IUCN.UK.2011-1.RLTS.T10831A3220342.en. Downloaded on
30 March 2018.
Reeves, R.R. et al. 2013. Inia geoffrensis. The IUCN Red List of Threatened Species 2013:
e.T10831A3220342. http://dx.doi.org/10.2305/IUCN.UK.2011-1.RLTS.T10831A3220342.
.en. Downloaded on 30 March 2018.
Rice D.W. 1998. Marine mammals of the world: systematics and distribution. Society for
Marine Mammalogy, Special Publication Number 4 (Wartzok D, ed.), Lawrence, KS. USA.
Rossetti D de F, Valeriano M.M .2007. Evolution of the lowest Amazon basin modeled
from the integration of geological and SRTM topographic data.Catena 70: 253–265.
doi:10.1016/j.catena.2006.08.009.
Ruiz-García M, S. Caballero, M. Martínez- Agüero and J. Shostell. 2008. Molecular
differentiation among Inia geoffrensis and Inia boliviensis (Iniidae, Cetacea) by means of
nuclear intron sequences. Pages 177–223 in V.P. Koven, ed. Population genetics research
progress. Nova Science Publishers, Inc., Hauppauge, NY.
Ruiz-García, M. 2010. Changes in the demographic trends of Pink River Dolphins (Inia) at
the microgeographical level in Peruvian and Bolivian rivers and within the Upper Amazon:
Microsatellites and mtDNA analyses and insights into Inia’s origin. Pages 225–258 in M.
Ruiz-García y J. Shostell, eds. Biology, evolution, and conservation of river dolphins
within South America and Asia: Unknown dolphins in danger. Nova Science Publishers,
Inc., Hauppauge, NY.
Secchi, E. 2012. Sotalia fluviatilis. The IUCN Red List of Threatened Species 2012:
e.T190871A17583369.http://dx.doi.org/10.2305/IUCN.UK.2012.RLTS.T190871A1758336
9.en. Downloaded on 30 March 2018.
Tavera, G., ER. Aliaga-Rossel., P.A. Van Damme,, A. Crespo. 2010. Distribution and
conservation status of the Bolivian river dolphin Inia boliviensis (d’Orbigny 1832). In:
Trujillo F, Crespo E, van Damme PA, Usma JS (eds) Action Plan South Am. River
Dolphins 2010-2020. WWF, Fundación Omacha, WDS, WDCS, Solamac, Bogota´, D.C.,
Colombia, pp 99–122.
Townsend P.A., Papeş, Eaton.M. 2007. Transferability and model evaluation in ecological
niche modeling: a comparison of GARP and Maxent. Ecography, 30, 550– 560.
Tchounwou, P.B., Ayensu, W.K., Ninashivili, N., Sutton, D. 2003. Enviromental exposure
to mercury and its toxicopathologic implications for public health. Environ Toxicol.
18:149-175.
Trujillo, F. 1992. Estimación poblacional de las especies dulceacuícolas de delfines de río
Inia geoffrensis y Sotalia fluviatilis en el sistema lacustre de Tarapoto y El Correo,
!
22!
Amazonía Colombiana. Special Report. Vol. 49. Centro de Investigaciones Científicas.
Universidad Jorge Tadeo Lozano, Bogotá D.C., Colombia.
Trujillo, F. 1994a. The use of photo-identification to study the Amazon River Dolphin, Inia
geoffrensis, in the Colombian Amazon. Marine Mammal Science 10 (3): 348-353 pp.
Trujillo, F. 1994b. Informe técnico de la Expedición Inia 94. Fundación Omacha. Bogotá,
D.C. 154pp.
Trujillo, 1995. Aspectos del comportamiento y la distribución de Inia geoffrensis y Sotalia
fluviatilis en el río Caquetá, Colombia. Special Report. Vol. 71. Centro de investigaciones
Científicas, Universidad Jorge Tadeo Lozano. Bogotá D.C., Colombia.
Trujillo, F., M.C. Diazgranados and L. Fuentes. 1999. Manual para la identificación de
Mamíferos Acuáticos. Fundación Omacha. 161p.
Trujillo, F. 2000. Habitat use and social behaviour of the freshwater dolphin Inia
geoffrensis (de Blainville 1817) in the Amazon and Orinoco basins. Ph.D. Thesis.
Aberdeen University. Scotland. 157 pp.
Trujillo, F., Diazgranados, M.C., Galindo, A and Fuentes, L. 2006. Delfón gris Sotalia
fluviatilis. Pp 273-278. En: Rodriguez-M.J.V., Alberico, M., Trujillo, F y Jorgenson, J. Eds.
2006. Libro Rojo de los Mamíferos de Colombia Serie Libros Rojos de Especies
Amenazadas de Colombia. Conservación Internacional y Ministerio de Ambiente, Vivienda
y Desarrollo Territorial. Bogotá D.C. Colombia.
Trujillo, F., Crespo, E., Van Damme, P. and J.S. Usma. 2010. The action plan for south
american river dolphins 2010 – 2020. WWF, WCS, Solamac, Fundación Omacha, 1-249.
Trujillo, F., Caicedo, D., and Diazgranados, M.C (Eds.). 2014. Plan de acción nacional para
la conservación de los mamíferos acuáticos de Colombia (PAN mamíferos Colombia).
Ministerio de Ambiente y Desarrollo Sostenible, Fundación Omacha, Conservación
Internacional y WWF. Bogotá D.C. 54 p.
UNEP-WCMC. 2010. Data Standards for the World Database on Protected Areas, UNEP-
WCMC.1–9.
VanDerWal J, Falconi L, Januchowski S, Shoo L, Storlie C. 2014. SDMTools: Species
Distribution Modelling Tools: Tools for processing data associated with species
distribution modelling exercises. R package version 1.1-221. Ver http://CRAN.R-
project.org/package=SDMTools
Velásquez-Roa, T., Carvajal-Castro, J.D., Mantilla-Meluk, H., Trujillo, F and Mosquera-
Guerra, F. 2015. Caracterización ecológica de cuencas utilizadas por el delfín rosado Inia
geoffrensis en Colombia, a través de modelamiento de nicho. Momentos de Ciencia. 12(2).
Pp: 126-130. ISSN 1692-5491
!
23!
Vidal, O., Barlow, J., Hurtado, L., Torre, J., Cendon, P., and Ojeda, Z. 1997. Distribution
and abundance of the Amazon River Dolphin (Inia geoffrensis) and the Tucuxi (Sotalia
fluviatilis) in the upper Amazon River. Marine Mammal Science 13(3) 427:445pp. http:/ /
dx.doi.org/10.1111/j.1748-7692.1997.tb00650.x
WCS (Wildlife Conservation Society) . 2017. http://aguasamazonicas.org/la-iniciativa/