Complex Patterns Of Male Alliance Formation In A Dolphin Social Network

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Chose TWO of the 4 articles attached below. For the summary follow the instructions below please thanks!

you have to define the scientific jargon into your own words so that a non-science person could understand your write up. Each summary should take up two FULL pages with 1-inch margins all around, 12pt font that is New Roman Times, and line spacing should be single spaced. The summary should be concise, no quoting directly from the article, and no outside citations. This will require you to read and understand the entire paper. You will have to define terms you are not familiar with and then you will have to take a long and complex paper and break it down to its essential content and significance


Format:


Name

Date


Full Citation of article as described form citation guidelines (follow formatting!!)


Body of your summary:

You should use the following subheadings and directions to write your summary.


Introduction:

Discuss the basic biological concepts of the article or the question(s) being asked by the study. Then briefly discuss what is known about the concepts the authors are writing about. This section would be critical to a reader who was not familiar with the scope of the study you are writing about.


Methods:

How was the research done? What were the methods used? This section should not get to technical or detailed since many readers might not be familiar with techniques in the area the authors are experts on. However, it should be very clear what was done and the assumptions made by the authors in terms the reader can understand.


Results:

Briefly and concisely report what was discovered in the study. This section should be short and to the point and do not try to itemize everything that the authors report inyou have to define the scientific jargon into your own words so that a non-science person could understand your write up. Each summary should take up two FULL pages with 1-inch margins all around, 12pt font that is New Roman Times, and line spacing should be single spaced. The summary should be concise, no quoting directly from the article, and no outside citations. This will require you to read and understand the entire paper. You will have to define terms you are not familiar with and then you will have to take a long and complex paper and break it down to its essential content and significance

Discussion and Conclusions:

How do the authors present how their results alter or change the field or concept discussed in the introduction? What is new about the results? What remains to be done on the topic and future steps?

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Journal of Mammalogy, 93(1):239–250, 2012 Complex patterns of male alliance formation in a dolphin social network JOANNA WISZNIEWSKI,* CULUM BROWN, AND LUCIANA M. MÖLLER * Correspondent: jo.wiszniewski@gmail.com The formation and maintenance of alliances is regarded as one of the most socially complex male mating strategies in mammals. The prevalence and complexity of these cooperative relationships, however, varies considerably among species as well as within and between populations living in different ecological and social environments. We assessed patterns of alliance formation for Indo-Pacific bottlenose dolphins, Tursiops aduncus, in Port Stephens, New South Wales, Australia, to investigate the stability of these alliances, the structure of associations, as well as variation in schooling patterns among males. Our results showed that association patterns among males within this population showed considerable variability. Males either formed strong and enduring alliances that lasted for at least 8 years with minimal partner switching, or less stable partnerships within a much larger male social network. Male alliances with the strongest levels of association within a given time period were significantly more likely to maintain their relationships over the long term compared with alliances with lower levels of association. Males in stable alliances also associated in significantly smaller schools than males who formed less stable alliance partnerships. Finally, we found that alliances consisting of more related males did not persist longer than alliances between unrelated individuals. Our study suggests that intrapopulation variation in male alliance formation in dolphins likely reflects different mating strategies adopted as individual responses to their complex fission–fusion social environment. Key words: alliance stability, bottlenose dolphin, coalition, cooperation, relatedness, social behavior, Tursiops E 2012 American Society of Mammalogists DOI: 10.1644/10-MAMM-A-366.1 The formation of coalitions and alliances is regarded as one of the most socially complex male mating strategies in mammals. Coalitions are generally defined as individuals opportunistically cooperating with others during competitive or aggressive interactions, whereas alliances refer to groups of individuals that maintain long-term cooperative relationships with each other (De Waal and Harcourt 1992). The formation of male coalitions and alliances for the purpose of reproduction has been an intriguing area of research given that in some cases, males cooperate to obtain a nondivisible resource (i.e., successful fertilization; van Hooff and van Schaik 1994). The degree of complexity of coalitionary behavior has been related to the presence of 3 traits: mutual tolerance, cooperation, and preference for particular partners on the basis of factors such as age, competitive ability, familiarity, and relatedness (Olson and Blumstein 2009). Only a few primate and delphinid species have been shown to possess all 3 traits, whereas several other species display less complexity with only 1 or 2 typical coalitionary behaviors (reviewed in Olson and Blumstein 2009). Cooperative partnerships among males may evolve through 3 possible interdependent pathways: kin selection, where individuals obtain indirect fitness benefits by cooperating with kin (Hamilton 1964); mutualism, where individuals gain immediate benefits from cooperation (Dugatkin 1997); or reciprocal altruism, in which the reciprocation of altruistic acts over time results in overall net benefits to all cooperating individuals (Trivers 1971). Mutual tolerance and collaboration among males can increase an individual’s reproductive success through a variety of mechanisms, including increased success in mate guarding (bottlenose dolphins, Tursiops—Connor et al. 1992a; lions, Panthera leo—Packer et al. 1991; chimpanzees, Pan troglodytes—Watts 1998), increased success in aggressive contests with other males (baboons, Papio cynocephalus—Noë 1994; lions—Grinnell et al. 1995), larger home ranges, which may increase their encounter rates with females (bottlenose www.mammalogy.org 239 Downloaded from https://academic.oup.com/jmammal/article-abstract/93/1/239/896437 by University of Indianapolis user on 05 April 2019 Department of Biological Sciences, Macquarie University, NSW 2109, Australia (JW, CB) School of Biological Sciences, Flinders University, Adelaide, SA 5001, Australia (LMM) 240 JOURNAL OF MAMMALOGY variability appear to differ among populations. For instance, Owen et al. (2002) showed that males displaying solitary behavior in 1 population were in a transitional phase rather than using alternative mating strategies. In contrast, the males adopting different types of alliance strategies in another population (Connor et al. 1992a, 2011) have been successful in obtaining paternities (Krützen et al. 2004a). Similarly, the influence of relatedness in alliance partner choice and mating patterns also appears to differ among populations. For example, in populations where stable alliances are formed among closely related males (Krützen et al. 2003; Parsons et al. 2003), individuals that increase the reproductive output of their partners can obtain inclusive fitness benefits even if they are not successful at siring offspring themselves. Consequently, male reproductive success is often highly skewed within these types of alliances (Krützen et al. 2004a). In other populations, however, patterns of alliance formation cannot be explained by kin selection (Möller et al. 2001; Owen 2003) and several lines of evidences suggest that preferences for non-kin partners are genuine. First, bottlenose dolphins are capable of recognizing maternal kin by their unique signature whistles (Sayigh et al. 1999). Second, Owen (2003) showed that although close relatives were generally available at the time when a male was forming an alliance partnership, males more often chose nonrelatives as alliance partners. Finally, levels of kinship among partners can vary depending on alliance strategy (Krützen et al. 2003). Similar intraspecific differences in kinship structure have been observed in lions, where alliances of 2 to 3 males were generally not related, but larger alliances were (Packer et al. 1991). Thus, in the absence of kin selection, cooperation among males can be explained by either mutualism or reciprocal altruism (Grinnell et al. 1995; Möller et al. 2001; van Hooff and van Schaik 1994; Watts 2002). Our study aimed to assess the long-term stability of male alliances in an embayment population of bottlenose dolphins (T. aduncus) in Port Stephens, Australia. The population displayed characteristics of a ‘‘fission–fusion’’ society and was socially and spatially segregated into 2 distinct communities (Wiszniewski et al. 2009). The core areas of the eastern and western communities coincide with a distinct change in habitat type (marine and estuarine, respectively; Fig. 1) and differences in social behavior have been observed between these 2 dolphin communities (Wiszniewski et al. 2009). School sizes were similar between the 2 communities (6 and 5.6 for the eastern and western communities, respectively); however, the average size of schools containing both eastern and western dolphins was more than double the size found for each community (Wiszniewski et al. 2009). Although the population was not strongly assorted by sex (Wiszniewski et al. 2010b), social relationships between males and females appear to be highly distinctive on the basis of short-term association patterns. Males often formed strong bonds with 1 to 3 other males (Möller et al. 2001), whereas females generally associated in a loose social network with a larger number of females (Möller et al. 2006). During the breeding season, male alliances were frequently observed cooperatively Downloaded from https://academic.oup.com/jmammal/article-abstract/93/1/239/896437 by University of Indianapolis user on 05 April 2019 dolphins—Owen et al. 2002), greater ability to defend or acquire territories (cheetahs, Acinonyx jubatus—Caro 1994), increase in social rank (chimpanzees—Goodall 1986), greater foraging efficiency (river otters, Lontra canadensis—Blundell et al. 2004; prairie dogs, Cynomys gunnisoni—Verdolin 2007), and reduced predation risk (squirrels, Xerus inauris—Waterman 1997). Theory predicts that alliances form only under a restricted range of social conditions, many of which are dependent on environmental variables. For example, alliances may be expected to form in populations when male competition is high due to limited availability of receptive females, when the benefits of group living (e.g., increased mating opportunities) outweigh the costs (e.g., increased feeding competition), or when alliances are effective in outcompeting single roving males (Whitehead and Connor 2005). Consequently, the prevalence and complexity of alliance formation is expected to vary considerably among species as well as within and between populations living in different ecological and social environments (Gehrt et al. 2008; Kappelar 2000; Whitehead and Connor 2005). Intrapopulation differences in social tactics among males (i.e., alternative reproductive behaviors) may result from 2 different pathways. First, individuals may adopt fixed strategies that remain constant over time (e.g., horses, Equus caballus—Feh 1999). More commonly, however, variation reflects conditional strategies, where males switch between mating tactics depending on their physical condition, age, or social rank in an attempt to increase their chances of reproductive success (e.g., three-spined stickleback, Gasterosteus aculeatus—De Fraipoint et al. 1993; and lance-tailed manakins, Chiroxiphia lanceolata—DuVal 2007; chimpanzees, Nishida 1983; baboons—Noë 1994). In bottlenose dolphins, genus Tursiops, patterns and complexity of alliance formation vary greatly among populations and appear to be correlated with population density and degree of sexual dimorphism. For instance, in 1 high-density embayment population, almost all males were found to associate in alliances and complex hierarchical patterns of alliance formation were also observed (Connor et al. 1992a, 2011). These higher-level alliances include second-order alliances, consisting of 2 or more 1st-order alliances, and 3rd-order alliances, which could consist of a combination of 1st- or 2nd-order (or both) alliances (Connor et al. 2011). Since males in this population are only slightly larger than females, cooperation in alliance partnerships may be required to successfully monopolize receptive females (Connor et al. 2000). In contrast, there has been no evidence of alliances in some low-density populations where either males are substantially larger than females and appear to use more solitary reproductive strategies (Wilson 1995) or where males associate in large, stable mixed-sex groups (Lusseau et al. 2003). In populations where male alliances are common, the range of alliance sizes and the stability of these alliances can vary considerably (Connor et al. 1992a; Möller et al. 2001; Parsons et al. 2003; Wells 1991). Explanations for such intrapopulation Vol. 93, No. 1 February 2012 WISZNIEWSKI ET AL.—MALE ALLIANCE FORMATION IN DOLPHINS 241 herding individual females for up to weeks at a time (Möller et al. 2001). We defined alliances using both association criteria and behavioral observations of 2 or more males herding a single female. Since the strength of association can vary among alliances (Möller et al. 2001) we investigated whether association strength is a good predictor of alliance stability. We also assessed whether strong associations existed between different alliances (2nd-order alliances, in sensu Connor et al. 1992a). We investigated potential differences in social patterns among males and compared alliance structure and complexity with other well-studied dolphin populations (e.g., Sarasota Bay, Florida and Shark Bay, Western Australia) to help increase our understanding of the social and ecological factors driving the evolution of complex male behaviors in dolphins and other social species (Connor et al. 2006; Olson and Blumstein 2009). Finally, we used a long-term behavioral and genetic data set to assess whether the longevity of alliances was significantly affected by the level of kinship among males. MATERIALS AND METHODS Data collection and restrictions.—The study was conducted within the Port Stephens embayment of approximately 166 km2 in surface area, located 200 km north of Sydney on the New South Wales coast of Australia (32u429S, 152u069E). The eastern basin of Port Stephens is typically a marine environment, with a strong tidal influx of coastal waters, sandy substrate, and large areas of sea grass, whereas the deeper western port is predominated by estuarine processes, including turbid, freshwater outflow from rivers and a muddy benthic habitat (Fig. 1). The bottlenose dolphin population is relatively small, consisting of around 90 individuals that were considered residents on the basis of photoidentification between 1998 and 2000 (Möller et al. 2002). The population is also genetically distinct from communities ranging on the adjacent coastline, with male-biased dispersal and a directional bias in migration from Port Stephens (Möller et al. 2007; Wiszniewski et al. 2010a). Data on dolphin school membership was obtained from transect surveys conducted between 1998 and 2008 using standard photoidentification techniques (Möller et al. 2006; for further details see Wiszniewski et al. 2009). A school was defined as all individuals within a 100-m radius (Irvine et al. 1981), and if traveling, the animals were heading in the same direction (Möller et al. 2006; Shane 1990). For each sighting, the latitude and longitude from a global positioning system, time, school size, number of calves, and predominant behavior were recorded. Sizes of schools were estimated by at least 2 trained observers. Estimates were later adjusted if the number of uniquely recognized dolphins exceeded the number of adults estimated in the field. To ensure reliability and independence of data, schools were excluded from the analysis if a minimum of 75% of the estimated school size was not reliably photographed; a fusion event occurred during photoidentification; or, if an individual was re-encountered during the survey, the second school was removed from the analysis. From a total of 945 schools, 113 did not meet the restriction criteria and were removed from the data set. Analyses were then conducted on data from the remaining 832 schools. Samples for genetic analysis were collected between 1999 and 2008 using the PAXARMS biopsy system (Timaru, New Zealand; Krützen et al. 2002), as described in Möller and Beheregaray (2001) and Wiszniewski et al. (2010a). Photoidentification and biopsy sampling surveys were conducted under licenses from New South Wales (NSW) Department of Environment and Climate Change (DECC; license no. S10763) and the NSW Marine Parks Authority (MPA; permit no. PSGLMP 2008/003). The research was also under approval by Macquarie University Animal Ethics Committee (AEC reference no. 2007/013) and met guidelines approved by the American Society of Mammalogists (Sikes et al. 2011). Males were identified using either genetic methods as described in Möller et al. (2001) or in the absence of genetic data, an individual was considered a male if the following 3 criteria were met: the individual was sighted more than 12 times over the study period, which was the median number of sightings over all individuals (X̄ 5 17.4, SE 5 1.1, max 5 Downloaded from https://academic.oup.com/jmammal/article-abstract/93/1/239/896437 by University of Indianapolis user on 05 April 2019 FIG. 1.—Habitat map of the Port Stephens embayment in New South Wales, Australia. 242 JOURNAL OF MAMMALOGY 38 males that were sighted 12 or more times over the study period and used the modularity matrix clustering technique described by Newman (2006) and Lusseau et al. (2008) to identify male group structure. In brief, the modularity matrix is the association index (i.e., weight) between 2 individuals minus the expected weight if associations were randomly distributed in the population. We calculated the expected weight by permuting associations within daily sampling periods using 10,000 permutations and 1,000 flips per permutation. This approach was used since it can account for the number of observations and associations of dolphins in each sampling period (Whitehead 2008). The eigenvector of the dominant eigenvalue of the modularity matrix is then used to successively split the matrix into 2 clusters. This divisive procedure is then iterated on all resulting clusters. The most parsimonious division in the network is subsequently determined by the division that maximizes the modularity coefficient, Q. Modularity is calculated by subtracting the expected proportion of total associations within groups from the observed proportion. Thus, Q ranges from 0, indicating randomly assigned groups, to 1, indicating no intergroup associations. The HWI matrix and resulting group structure determined by Qmax was visualized using the springembedding method (Kamada and Kawai 1989) in NETDRAW (Borgatti 2002). The spring-embedded algorithm is simply an iterative procedure that arranges nodes in such a way that those with the highest association levels are closer together (Gajer and Kobourov 2002). Thus, nodes with the greatest density of links are often more centrally located, whereas those with few links are placed on the periphery of the network. Stability of male associations.—We used the male alliances defined in the four 2-year time periods to assign each individual a value between 1 and 4 on the basis of the number of 2-year time periods that the male had the same alliance partner (procedure adapted from Mitani 2009; Silk et al. 2006). Thus, males whose partner was the same in all 4 time periods had a stability index of 4. Males that were allied in 1 or no time periods were grouped together in category 1. To determine whether associations among males predicted their association levels in future years, we followed the procedure of Mitani (2009) and conducted a series of Mantel correlation tests (Hemelrijk 1990; Mantel 1967; Schnell et al. 1985) that compared pairwise HWI values calculated in 1 time period with values calculated in subsequent years. For each comparison, only males present in both time periods were included in the analysis. Significance for all correlations was assessed using 10,000 random permutations. To assess whether alliance stability is related to strength of association, 2 statistical analyses were conducted. First, we correlated mean association strength within alliances with alliance duration using the Pearson’s correlation coefficient. Second, we used lagged association rate (LAR) analysis (Whitehead 1995), where LAR is the probability of finding any given pair of individuals together after different time lags. The LAR was calculated for males within each category (i.e., Downloaded from https://academic.oup.com/jmammal/article-abstract/93/1/239/896437 by University of Indianapolis user on 05 April 2019 65); the individual was considered an adult in 2002 or earlier (on the basis of size) and never seen with a dependent calf; and the individual’s top associate was a genetically sexed male. We found 9 individuals that met all 3 criteria. Although there is some uncertainty in this classification, these restriction criteria indicate a high probability that these individuals were males (Santos and Rosso 2008; Shane 2004). For example, of the 64 females identified in the population, only 2 had a top associate that was a genetically sexed mal ...
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Wiszniewski, J., Brown, C., & Möller, L. M. (2012). Complex patterns of male alliance
formation in a dolphin social network. Journal of Mammalogy, 93(1), 239-250.
Introduction
The article presents results of a study conducted by Joanna Wiszniewski, Culum Brown
and Luciana M. Moller on “Complex patterns of male alliance formation in a dolphin social
network.” Mammals often form coalitions and alliances as strategies for male mating. This has
been an area of interest in research as it entails male mammals cooperating by forming coalitions
and alliances in order to attain successful fertilization; a resource that is not divisible. The
formation of coalitions and alliances is a complex process that depends on numerous traits such
as cooperation, mutual tolerance, as well preference for given partners based on compatibility,
age, and relatedness among other factors. Coalition refers to individuals cooperating with each
other to get an opportunity in competitive. Alliances on the other hand entail individuals forming
long-lasting relationships with one another.
Methods
The PAXARMS biopsy system was used to collect samples for genetic analysis between
the period of 1999 and 2008. Genetic methods were used in identification of males or in the
absence of genetic data. The half-weight index was used to examine the patterns of association
such as strength of the association among dyads. Social network analysis was carried out using
data set pooled for 8 year period to further examine the patterns of male association. The male

alliances established in the four 2-year time periods was employed to assign each male a number
between 1 and 4 based on the number of times during the 2-year time periods a male maintained
the same alliance partner. Hence, males with the same partner throughout the total 4 periods
were assigned a stability index of 4. Mantel correlation tests were utilized to determine whether
existing associations were a prediction of future levels of association. In order to examine
whether there is a correlation between alliance stability and strength of association, two tests
were conducted: correlations was done on mean association strengths and with the use of
Pearson’s correlation coefficient. Secondly, the lagged association rate (LAR) analysis was
conducted whereby LAR represented the probability of two individuals been in the same alliance
during different time lags.
The PERM program was used to examine the role of genetic relatedness in the formation
of male associations. The school size of the alliances formed was also examined on various
alliances with varied stability. School sizes were also grouped based on the level of male stability
in order to test the hypothesis that larger schools constituted of less stable associations.
Results
The results of the study indicated that formation of alliances was predominant among the
bottlenose dolphin population in the P...

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