Inter-individual association levels correlate to behavioral diversity in coastal bottlenose dolphins (Tursiops truncatus) from the Southwestern Gulf of Mexico

Abstract

Introduction: Societies in mammal species are influenced by intrinsic and extrinsic factors that affect the temporality of the associations among individuals. Coastal bottlenose dolphins generally live in small fluid aggregations variable in composition, but the nature of their associations is commonly unknown. Our goal was to determine if school size was influenced by dolphins’ behavior, and if individuals associated to develop particular activities within the coastal waters of Alvarado, Mexico.
Methods: In total, 80 boat-based surveys were conducted (2002 - 2003 and 2006 - 2009), where group size, behavior, and photo-identification data were collected. From 237 sightings and 2,021 dolphins the mean school size was 8.5 animals (s. d. = 8.6), but individuals and pairs were observed more frequently (33 %).
Results: Temporal differences in school sizes and behavior conveyed with habitat seasonality (P < 0.05), but were inconsistent across years; thus short-term factors such as marine traffic and fisheries intensity were deemed important. Dolphins were commonly feeding (29 %) corresponding to groups of 4 - 6 animals, whereas solitary individuals typically showed evasion (P < 0.01).
Discussion and Conclusions: Association coefficients computed for 89 of the 232 identifiable dolphins proved not random only in 6 % of the 3,915 combinations (P < 0.05), and these coefficients were positively correlated to the diversity of activities developed by each dyad (P < 0.01), thus the nature of their associations in most cases (93 %) became richer with time. The community is likely formed by very small units that frequently exchange members; this may be due to large food availability and low predator abundance in the area, but also to avoid detection and threats posed by local artisanal fisheries.

Full text

THERYA, 2015, Vol. 6(2): 337-350 DOI: 10.12933/therya-15-270, ISSN 2007-3364
Las asociaciones entre individuos se correlacionan
con la diversidad de las conductas en delnes costeros
(Tursiops truncatus) del Sureste del Golfo de México
Inter-individual association levels correlate
to behavioral diversity in coastal bottlenose
dolphins (Tursiops truncatus) from the
Southwestern Gulf of Mexico
Marah García-Vital1, Eduardo Morteo1, 2*, Ibiza Marnez-Serrano3, Alberto Delgado-Estrella4 y Carmen Bazúa-Durán5
1Instituto de Ciencias Marinas y Pesquerías, Universidad Veracruzana. Calle Hidalgo Núm. 617, Col. Río Jamapa, CP 94290, Boca del
Río, Veracruz, México. E-mail: maravital@gmail.com (MGV).
2Instituto de Investigaciones Biológicas, Universidad Veracruzana. Calle Dr. Luis Castelazo Ayala s/n, km 2.5 Carr. Xalapa-Veracruz,
Col. Industrial Ánimas, CP 91190, Xalapa, Veracruz, México. E-mail: eduardo.morteo@gmail.com (EM).
3Facultad de Biología, Universidad Veracruzana. Museo de Zoología. Facultad de Biología, Universidad Veracruzana. Circ. Gonzalo
Aguirre Beltran s/n Zona Universitaria. CP 91090. Xalapa, Veracruz, México. E-mail: ibimartinez@uv.mx (IMS).
4Facultad de Ciencias Naturales, Universidad Autónoma del Carmen. Calle 56 Núm. 4, Esq. Av. Concordia, Col. Benito Juárez, 24180,
Cd. del Carmen, Campeche, México. E-mail: delgadoestrella@gmail.com (ADE).
5Facultad de Ciencias, Universidad Nacional Autónoma de México. Circuito exterior s/n, Ciudad Universitaria, 04510 México, D. F.,
México. E-mail: bazua@unam.mx (CBD).
*Corresponding author
Introduction: Societies in mammal species are inuenced by intrinsic and extrinsic factors that aect the temporality
of the associations among individuals. Coastal bottlenose dolphins generally live in small uid aggregations variable in
composition, but the nature of their associations is commonly unknown. Our goal was to determine if school size was
inuenced by dolphins’ behavior, and if individuals associated to develop particular activities within the coastal waters
of Alvarado, Mexico.
Methods: In total, 80 boat-based surveys were conducted (2002 - 2003 and 2006 - 2009), where group size, behavior,
and photo-identication data were collected. From 237 sightings and 2,021 dolphins the mean school size was 8.5
animals (s. d. = 8.6), but individuals and pairs were observed more frequently (33 %).
Results: Temporal dierences in school sizes and behavior conveyed with habitat seasonality (P < 0.05), but were
inconsistent across years; thus short-term factors such as marine trac and sheries intensity were deemed important.
Dolphins were commonly feeding (29 %) corresponding to groups of 4 - 6 animals, whereas solitary individuals typically
showed evasion (P < 0.01).
Discussion and Conclusions: Association coecients computed for 89 of the 232 identiable dolphins proved not
random only in 6 % of the 3,915 combinations (P < 0.05), and these coecients were positively correlated to the diversity
of activities developed by each dyad (P < 0.01), thus the nature of their associations in most cases (93 %) became richer
with time. The community is likely formed by very small units that frequently exchange members; this may be due to
large food availability and low predator abundance in the area, but also to avoid detection and threats posed by local
artisanal sheries.
Key words: Avoidance; behavior; group size; membership; socialization; sheries.

338 THERYA Vol.6(2): 337-350
DOLPHIN BEHAVIOR AND SOCIOLOGY
Introduction
Societies in mammal species are dynamic and complex (e. g. carnivores, primates, cetaceans),
in which many of their members may interact and associate with other known or unknown
individuals, forming distinctive assemblages (Goodall 1986). The structure of such social units is
often aected by age, sex, reproductive status, hierarchy and kin selection (Beddia 2007); all these
aspects may modify the behavior of individuals (Bräger 1993) and determine how the animals
spend their time in certain areas, thus producing dierent patterns of residency, seasonality or
migration that change their associations and the social structure of the population (Scott and
Chivers 1990).
Dolphin societies or communities (sensu Wells et al. 1987) are generally composed by individuals
that inhabit the same general area and have frequent interactions with each other (Goodall 1986);
thus marine mammalogists often use this term to describe assemblages of individuals of the same
species, instead of assemblages of dierent species as referred in texts of classic ecological theory
(Roughgarden 1989). Such aggregations are loose and uent, where individual and genetic
exchange may occur over time within the limits of the community (Wells et al. 1987).
Individual interactions are inuenced by intrinsic factors such as the presence of recurrent
associations (Connor et al. 2001), which in turn may be determined by extrinsic factors such as
habitat variability (e. g. food abundance and availability, as well as natural and human-related
threats (Lusseau et al. 2006; Quintana-Rizo 2006; Morteo and Hernández 2007; Morteo et al. 2012).
Social and ecological pressures may dictate the temporality of the associations at dierent scales
(McDonald and Carr 1989; Quintana-Rizzo 2006; Morteo et al. 2014). For instance, some species
are known to maintain strong permanent and even multilevel associations such as killer whales
(Orcinus orca) and sperm whales (Physeter microcephalus; Bräger et al. 1994; Whitehead et al. 2012),
whereas other like the spinner dolphin (Stenella longirostris) are very loose (Chilvers and Corkeron
2002).
The social structure of coastal bottlenose dolphins (Tursiops truncatus) has been characterized
as ssion-fusion, involving small groups that constantly exchange individuals forming a wide range
of social bonds (Goodall 1986; Wells et al. 1987; Connor et al. 1992). Several communities of the
coastal form of this species are known to present segregation related to sex, age and reproductive
status, thus the nature of their associations is highly variable (Wells et al. 1987; Wells 1991; Connor
et al. 1992; Smolker et al. 1992; Connor et al. 2000; Quintana-Rizzo and Wells 2001; Morteo et al.
2014).
Living in social groups facilitates feeding, protecting against harassment, predation, enhances
reproductive output, and promotes communication and learning (Bräger et al. 1994). Individuals
may join or leave a group in response to the gains or losses of participating with the partners
involved within a given social unit (Wrangham et al. 1993). Thus recording the activities of animal
groups is useful to establish behavioral patterns in a specic habitat (Bräger 1993; Steiner 2011),
but also to determine the current state of individuals within a community (Beddia 2007).
Coastal bottlenose dolphins inhabiting the waters o Alvarado in the state of Veracruz have
been studied intermittently since 1993 (García 1995) and reliable data on individual identities has
been collected since 2002 (Del Castillo 2010; Morteo et al. 2014). Many of these dolphins are
known to associate and develop a range of activities, but little is known on how these animals
interact and form groups, and if they do it with dierent purposes (Morteo 2011; García-Vital
2012). The goal of this study is to establish if school size is determined by the activities of the
dolphins, and if individuals associate with specic partners to develop particular sets of activities.

www.mastozoologiamexicana.org 339
García-Vital et al.
Materials and methods
Study area. The Alvarado region is a shallow (< 20 m depth) open coastal habitat in the
southwestern Gulf of Mexico, strongly inuenced by river discharges Figure 1; habitat
modication is the major threat to the area (Del Castillo 2010; Morteo 2011; García-Vital 2012).
It is the third largest coastal lagoon system in Mexico, and according to the National Institute
of Fisheries (INAPESCA), it’s the most important shrimp shing ground in the state of Veracruz,
which takes place year-around depending on weather and market demands. Sea surface
temperature ranges from 20 to 32.5 °C, with an annual average of 27 °C. The regional climate is
tropical with three seasons: a Dry season (March-June) with a signicant reduction in average
precipitation; a Rainy season (July-October), in which runo from rivers into the adjacent lagoon
and mangrove forest causes high organic matter and nutrient input into coastal waters; and a
Windy season (November-February, locally known as “Nortes”), featuring strong winds (up to
80 km h-1) associated with the incursion of northern cold fronts, which may last several days.
Fisheries are relevant to coastal dolphin populations due to the frequent adverse interactions
between dolphins and local sheries, causing incidental mortality of dolphins in nets (Morteo
2011; Morteo et al. 2012). Around 2000 shermen were active in the area, most of which (75 -
85 %) operate in the lagoon and the rest operate in open waters. No ocial data is available on
marine trac or shing eort and port facilities are dedicated to shing, therefore there is no
alternative commercial seagoing activity.
Surveys. Photographic line-transect surveys were conducted at least twice per month from
October 2002 to September 2003 and from May 2006 to April 2009. The extent of the surveyed
area was dependent on the duration of daily operations, and intended to maximize the chance
Figure 1. Study area and survey trajectories (bold lines). Dashed lines show depth contours every 5 m.

340 THERYA Vol.6(2): 337-350
DOLPHIN BEHAVIOR AND SOCIOLOGY
of encountering coastal dolphins, based on their habitat preferences (Fazioli et al. 2006). Surveys
were carried out at constant speed (15 - 18 km h-1) always in low swell conditions (sea state
Beaufort ≤ 3, where wind speed < 15 km h-1) on board of a 7 m outboard berglass ski (40 / 60
hp). When dolphins were sighted, the survey was paused to allow habituation (10 min) while
their location was recorded using a GPS (Garmin eTrex Legend). Dolphins were then observed
until their behavior was determined ad libitum (Altmann 1974) and classied into one of the most
commonly used categories which were: 1) Feeding, 2) Socializing, 3) Traveling, 4) Avoiding, 5)
Resting, and 6) Undetermined, following the literature (see Shane 1990; Bräger 1993; Chilvers and
Corkeron 2002; Constantine et al. 2004; Steiner 2011). Subsequently, the group was approached
with caution to avoid disturbing them, at the time that group size was estimated; we used an
inclusive denition of group, consisting in all dolphins observed in apparent association, moving
in the same direction and often, but not always, engaged in the same activity (Bräger et al. 1994);
however, some groups included only one dolphin (Fazioli et al. 2006). Dolphins were followed
until all dorsal ns were photographed or until they were lost from sight; we used SLR cameras,
both analogical (Canon EOS Rebel 2000 with lm Kodak Tri-X-pan ISO 400) and digital (Canon
Rebel XT and Nikon D50) with 70 - 300 mm lenses. The survey was then resumed and the search
proceeded until the study area was completed.
Group size and behavior. All sighting records from 2002 - 2003 and 2006 - 2009 (2002 - 2009
henceforth) were arranged into histograms to determine the most common types of aggregations
and behaviors within the area. Average group size was computed for comparison, whereas
dierences among months, seasons and years were examined for both group size and behavior
frequencies using non-parametric tests (α = 0.05). Also, a correspondence analysis was performed
to explore the relation between group size and the behavioral categories using Statistica 7.0.
Photographic identication. Individual dolphins were identied by the marking patterns on
their dorsal ns (Würsig and Jeerson 1990; Morteo 2011). Markings such as tooth rakes, scars,
pigmentation marks, supercial wounds, and epiphytic organisms are temporary, thus these were
considered unidentiable and were excluded. Only dolphins with permanent and conspicuous
markings were included, only if they were sighted in ve or more survey days (Bräger et al. 1994;
Félix 1997; Bejder et al. 1998; Rogers et al. 2004); thus, analyses were performed only for dolphins
with a certain degree of residency.
Association patterns and behavioral diversity. Photographic data were also used to compute
half-weight (i. e. controlled for sighting frequencies) coecients of association (COA) for each
dyad (pair of individuals; e. g. Smolker et al. 1992; Bräger et al. 1994; Félix 1997; Quintana-Rizzo
and Wells 2001; Rogers et al. 2004) using SOCPROG 2.4 (Whitehead 2009). COA values range from
zero for dolphins that are never seen together, to one for a pair that is always seen together. COA
values were categorized as infrequent (0.0 - 0.2), casual (0.2 - 0.4), fair, (0.4 - 0.6), moderate (0.6
- 0.8) or strong (0.8 - 1.0; Smolker et al. 1992; Quintana-Rizzo and Wells 2001). Also, to prevent
the occurrence of articial dyads (i. e. individuals photographed together by chance due to our
inclusive denition of group), a preferred/avoided partner assessment was developed through a
permutation test (Smolker et al. 1992; Bejder et al. 1998; Gero et al. 2005).
The recorded behaviors were used to calculate a measure of diversity for the activities
developed by all paired individuals; therefore, the Shannon-Wiener index was used as a proxy
for each dyad; the latter was standardized to meet the COA range values (0 - 1) and renamed as
activity diversity index (ADI). Since dolphin sightings containing more than two individuals would
use the same behavioral records for the computation of the ADI in their respective dyads, the
result would overestimate the contribution of each pair to the index (Hurlbert 1984); therefore, we
randomly removed photographic data from individual dolphins until the statistical distribution of
the ADI and the COA values stabilized. We then computed new COA and ADI in order to reduce

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García-Vital et al.
the bias from the pooling fallacy (Machlis et al. 1985). Finally, the resulting COA and ADI matrices
were analyzed using a Mantel’s one tailed test, which measures the correlation between the values
within two symmetrical matrices containing the data from all the possible combinations between
individuals, throughout a randomization process (α = 0.05, Spearman correlation, Monte Carlo
method and 10,000 permutations).
Results
Survey eort and group size. The Alvarado area was sampled on 80 photographic surveys
accounting for 237 groups and 2,021 sighted dolphins. Average school size was 8.5 (s. d. = 8.6) but
single animals (16.7 %) and pairs (16.3 %) accounted for almost a third of the recorded sightings
(Figure 2). Groups were signicantly larger in specic months (June and April, P < 0.01, Table 1),
seasons (dry, P < 0.01, Table 1) and years (2002-3, 2007-8, P < 0.01, Table 1) across the study period.
Table 1. Group size (numbers) and prevalent behaviors of bottlenose dolphins observed
within dierent periods in the study area (n = 237, * = P < 0.01).
Period Mean (s.d.) Group size Behavior
2002-2003 10.3 (1.1) * Apr*, Jun* Feeding*, Resting*
2006-2007 7.6 (0.6) - Feeding*
2007-2008 9.6 (0.9) * Sep* Socializing*
2008-2009 8.2 (0.7) Nov*, Feb* -
Rainy 8.4 (0.7) Sep* Feeding*
Windy 7.6 (0.8) Nov*, Feb* -
Dry 9.3 (0.7) * Apr*, Jun* -
0
2
4
6
8
10
12
14
16
18
1
2
3
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
>20
Proportion %
Group size
Figure 2. Group size proportion for bottlenose dolphins sighted in the study area during 2002 - 2009 (n = 237).
Behavior and group size. Behavioral records existed for 220 of the 237 sightings, where feeding was
the most common activity in the area (29 %, χ2 = 15.22, P < 0.05), but this was true only for years
2002-2003 and 2006-2007 (P < 0.05, Table 1). Seasonal dierences were observed only for the
rainy months, were feeding was also signicantly more frequent (χ2 = 23.05, P < 0.05).
Group size was arranged into class intervals according to its frequency in order to homogenize
sample sizes (Figure 2). Correspondence between group size and behavior was signicant within
the studied period (χ2 = 69.47, P < 0.01) where the rst two dimensions explained 76.1 % of the

342 THERYA Vol.6(2): 337-350
DOLPHIN BEHAVIOR AND SOCIOLOGY
variance, and only three behavioral categories were associated to group size: dolphin pairs were
mostly resting, whereas feeding aggregations were formed of four to six dolphins, and solitary
individuals commonly showed evasion (Figure 3).
Association patterns and behavioral diversity. A total of 232 dierent dolphins were individually
identied, but only 89 were sighted at least ve times, and only one was recognized on up to
47 occasions. Association values (COA) changed annually (Figure 4), as the composition of the
members within the community was very uid, and only eight individuals were consistently
identied over the years. The combination of these 89 individuals resulted in 3,915 possible dyads
and 67.2 % of these were photographed at least once; however, the permutation test showed that
only 237 pairs (6 %) were non-random (P < 0.05). From the latter, visual inspection of COA values
showed that 11 % avoided each other (0.0 - 0.2), or met infrequently (0.2 - 0.4), whereas 68 % were
moderately associated (0.4 - 0.6), and the remaining were either close (7 %; 0.6 - 0.8) or very close
(0.8 - 1.0) partners (3 %; Figure 5).
The diversity of activities (ADI) for each dyad showed a wide range of values, and its statistical
distribution stabilized upon the random elimination of half of the records from the 237 signicant
pairs (Figure 5). Only 5 % of the dyads had very high diversity (0.8 - 1.0), 9 % had high diversity
(0.6 - 0.8), 31 % were fairly diverse (0.4 - 0.6), 34 % developed a low diversity of activities (0.2 - 0.4),
and 20.3 % of the dyads were selective (0.0 - 0.2, all null values corresponded to avoided partners,
i. e. COA = 0; Figure 5).
The Mantel’s test showed signicant correlation between the COA and ADI matrices (r(AB) =
0.98, P < 0.01), thus the diversity of activities developed by each dyad increased signicantly with
their level of associations; however, a few instances deviated signicantly from this pattern. For
instance, 7.2 % of the dyads had moderate to very high associations but developed a low or very
low diversity of activities (low-right corner in Figure 6), specically feeding and socializing.
Figure 3. Correspondence analysis between activities and group class (size) for bottlenose dolphins sighted in the study area
during 2002 - 2009. F = Feeding; T = Travel; E = Evasion S = Social; R = Rest (n = 220).

www.mastozoologiamexicana.org 343
García-Vital et al.
Discussion
Group size. Aggregations in animals are widely variable in size and composition, and their study
is closely linked to the criteria used to dene their limits (Shane et al. 1986; Wells et al. 1987; Shane
1990; Wells 1991; Smolker et al. 1992; Bejder et al. 1998; Whitehead 1999; Connor et al. 2000;
Mareike 2003; Morteo 2011). The number of individuals in a group has been related to advantages
while overcoming selective pressures, including but not limited to protection against harassment
and predation, improving food acquisition and energy eciency, thus leading to enhanced
reproductive output (Wells et al. 1980; Shane et al. 1986; Mareike 2003).
Mean group size is often helpful to represent the optimal number of individuals for a community
within a specic habitat (Würsig 1979), and the average group size found here (8.5 ± 8.6 s.d.) was
similar to other coastal locations across the Gulf of Mexico, the Caribbean and the Atlantic (Shane
et al. 1986; Wells et al. 1987; Scott and Chivers 1990; Delgado 2002; García 1995; Morteo and
Hernández 2007; Hernández-Candelario 2009; Martínez-Serrano et al. 2011). However, since the
statistical distribution was skewed towards smaller aggregations (i. e. one third of the sightings
were composed of solitary individuals and pairs) despite our inclusive denition (Figure 2), this
value seems inconclusive. Morteo et al. (2014) anticipated this for bottlenose dolphins within
the same study area, and determined that many of these small aggregations were likely single
sexed. This strongly suggests that the community may be divided into small social units that join
in common activities (Campbell et al. 2002).
Behavior. The general behavioral pattern suggested for most bottlenose dolphin communities
is: Travelling > Socialization or Feeding > Resting > Evasion (Morteo 2002). Feeding has been
reported as a common activity in many dolphin communities (Shane 1990; Bräger 1993; Morteo
Figure 4. Annual sociograms for dolphins identied at least 5 times in the study area during 2002 - 2009. Line thickness shows
the values for coecients of association between dyads. Individuals with COA < 0.6 were removed (n = 27, n = 89), and also diagrams
and line thickness were scaled dierently to t the gure.

344 THERYA Vol.6(2): 337-350
DOLPHIN BEHAVIOR AND SOCIOLOGY
2002; Beddia 2007; Steiner 2011) and was the single most frequent activity (29%) within the
studied period, especially in the rainy months (15.5 %). Maze and Würsig (1999) hypothesized
that dolphins may use environmental cues such as continental water runos to congregate over
shallow coastal areas close to rivers and lagoons, taking advantage of increased prey populations;
in fact, coastal bottlenose dolphins in this and in neighboring locations are known to experience
seasonal expansions in their distribution during this time of the year (Martínez-Serrano et al. 2011;
Medellín-Ortiz 2012). Although the latter may seem to enhance the possibilities of interacting with
other dolphins in larger aggregations, Morteo (2011) suggested that as food seems abundant and
predators are scarce in this study area, large groups might not necessarily imply an advantage for
foraging or protection. The statistical distribution for group size is consistent with this hypothesis
and it also explains the correspondence between the feeding records and the groups composed
by 4 - 6 dolphins (Figure 3), thus strengthening the notion of a small-unit based social structure.
Following the same hypothesis, solitary individuals were signicantly associated to evasive
behavior (17 % of recorded sightings, Figure 3), which may be due to the frequent and antagonistic
interactions between dolphins and artisanal sheries using gillnets in this location. Resident
Figure 5. Proportion of non-random dyads by association (COA) and behavioral diversity (ADI) class in the study area during
2002 - 2009 (n = 237).
0
0.1
0.2
0.3
0.4
0.5
0.6
0.7
0.8
Very Low
(0.0-0.2)
Low (0.2-
0.4)
Moderate
(0.4-0.6)
High (0.6-
0.8)
Very high
(0.8-1.0)
Index value
Class
COA
ADI
Figure 6. Mantel’s correlation test (r = 0.98, P < 0.01) between association coecients (COA) and diversity of activities (ADI) for
non-random dyads identied in the study area during 2002 - 2009 (n = 237). Half of the individual data was randomly removed.

www.mastozoologiamexicana.org 345
García-Vital et al.
individuals face the inherent risk of being harassed, entangled or even killed, thus individuals rather
than groups are more likely to evade detection by shermen due to the extremely low visibility
below the sea surface (Morteo et al. 2012). The latter also supports the argument that individuals
in general gain larger benets by aggregating in small groups (Figure 2) within this heavily shed
area, and in extreme cases points to a “selsh strategy” as a mean for auto-preservation.
Habitat seasonality agreed with temporal dierences found in both group size and behavior
(Table 1; Del Castillo 2010; Martínez-Serrano et al. 2011; Morteo 2011; Medellín-Ortiz 2012);
however, short-term habitat variability also seems to play an important role, since both of these
variables were inconsistent throughout the studied period (Table 1). Morteo et al. (2012) pointed
out the importance of boat trac and shing activities in the area, which seem to inuence the
presence and distribution of dolphins on a daily basis or even instantaneously (Constantine et al.
2004; Lusseau et al. 2006; Hernandez-Candelario 2009). Although many cetaceans seem to have
habituated to certain levels of marine trac, other evidence suggest that it may cause severe
alterations in their behavior and in this case may promote the separation of group members,
probably altering the social bonds within the community (Constantine et al. 2004; Morteo et al.
2012; Morteo et al. 2014).
Association patterns and behavioral diversity. A social organization is dened by the relations
and interactions among individuals within the sampled population (Chapman et al. 1995).
Societies are believed to gain from all the variety in their associations (Dunbar 1989); however,
even in ssion-fusion societies, partners could be selected to maximize eciency or benets while
joining to develop their activities (Gero et al. 2005). In this study, up to 94 % of all possible paired
associations were non-signicant, rearming the uid nature of the community; however, 78 %
of the non-random pairs exhibited moderate to high membership (Figure 4). The congregation of
prey within a limited area could help explain the random encounters for many of these dolphins
(Morteo et al. 2012); on the other hand, the close associations for a small part of the community
is likely the result of male alliances, and female bands (sensu Connor et al. 2000), by means of the
sexual segregation occurring in the area (Morteo et al. 2014).
We found that dolphin interactions seem to modulate group size (Figure 2), and these are
inherently associated to their own activities (Connor et al. 2001; Figure 3 and 5). The positive
correlation between the level of association and the diversity of activities (Figure 6) was expected,
as the context of the interactions between dyad members should become richer over time (Gero et
al. 2005); however, the fact that 17 of the 237 signicant dyads involved partnerships with specic
purposes suggests that only a handful of these individuals nd larger benets from exploiting
the specic abilities of selected partners. Lusseau et al. (2006) evidenced the importance of roles
for specic individuals within a social network, where previous knowledge of group members
and reduced number of participants helped to coordinate actions and facilitated cooperation;
however such interactions were rare and deserve further attention.
Knowledge of the social structure of dolphin communities is important for assessing their
ecological and evolutionary patterns; and behavioral data helps to understand how the animals
adapt to the habitat and its selective pressures. However, both aspects are rarely combined to
determine the nature of their associations, and how these are modied by their life conditions,
including natural and anthropogenic factors. Human developments cause that coastal bottlenose
dolphins face increasing disruptions within their core areas of distribution, therefore these aspects
should be considered while developing and applying conservation and management strategies
and policies.

346 THERYA Vol.6(2): 337-350
DOLPHIN BEHAVIOR AND SOCIOLOGY
Acknowledgments
This research is part of the lead author’s MSc thesis at the Universidad Veracruzana, Mexico,
where she obtained fellowship from CONACyT. J. Montano, I. Hernández, V. Del Castillo and N.
Medellín were involved in surveys and data collection. Fieldwork was carried with authorization
from SEMARNAT permits SGPA/DGVS/00351/06 (E. Morteo) and SGPA/DGVS/00870/07, 02788/07,
01344/08 and 01649/08 (C. Bazúa). This work was supported by the following grants: PROMEP
Apoyo a Nuevo PTC (E. Morteo) and CAMyCRA (E. Morteo, and H. Pérez-España), CONACyT grant
45468 (E. Velarde) and the Marine Mammal Laboratory of Universidad Veracruzana and Acuario de
Veracruz, A.C.
Resumen
Introducción: Las sociedades en mamíferos están inuenciadas por factores intrínsecos y extrínsecos afectando
la temporalidad de las asociaciones entre individuos. Los tursiones costeros generalmente viven en agregaciones
pequeñas y uidas de composición variable, pero comúnmente se desconoce la naturaleza de sus asociaciones.
Nuestro objetivo fue determinar si el tamaño de grupo se relaciona con el comportamiento de los delnes y si los
individuos se asocian para desarrollar actividades particulares en las costas de Alvarado, México.
Metodología: Se realizaron 80 navegaciones (2002 - 2003 y 2006 - 2009) para registrar la conducta, el tamaño
de grupo y para foto-identicar a los delnes. De 237 avistamientos, con 2,021 delnes observados, se obtuvo un
promedio de 8.5 (d. e. = 8.6) animales por grupo, donde las parejas y los individuos fueron más frecuentes (33 %).
Resultados: Las diferencias en tamaño de grupo y comportamiento concordaron con la estacionalidad del
hábitat (P < 0.05), siendo inconsistentes entre años, por lo que los factores de corto plazo (i e. tránsito marino e
intensidad de pesca) fueron importantes. La alimentación fue común (29 %) y correspondió con grupos de 4 – 6
animales, mientras los individuos solitarios típicamente mostraron evasión (P < 0.01).
Discusión y conclusiones: Los coecientes de asociación calculados para 89 de 232 delnes identicables fueron
no aleatorios en sólo 6 % de las 3,915 combinaciones (P < 0.05); dichos valores se correlacionaron positivamente con
la diversidad de actividades de cada pareja (P < 0.01) y la naturaleza de sus asociaciones se volvió más rica con el
tiempo en el 93 % de los casos. Se cree que esta comunidad se compone de pequeñas unidades que intercambian
miembros; esto puede deberse a una alta abundancia de alimento y bajo número de depredadores en el área, pero
también a una estrategia para evitar la detección y las amenazas derivadas de la pesca artesanal.
Palabras clave: Comportamiento; evasión; membresía; pesquerías; socialización; tamaño de grupo.
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Summited: February 13, 2015
Review: April 8, 2015
Accepted: April 21, 2015
Associated editor: Juan Pablo Gallo

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