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Summary
Discriminating amongst individuals is a disquisitional social behavior in humans and many other animals [
,
,
] and is ofttimes required for offspring and mate recognition, territorial or coalitional behaviors, signaler reliability assessment, and social hierarchies [
4
- Medvin One thousand.B.
- Stoddard P.K.
- Beecher G.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
v
- Hare J.F.
Juvenile Richardson'due south footing squirrels, Spermophilus richardsonii, discriminate amongst private alarm callers.
- Crossref
- PubMed
- Google Scholar
,
,
7
- Hare J.F.
- Atkins B.A.
The squirrel that cried wolf: Reliability detection by juvenile Richardson's ground squirrels (Spermophilus richardsonii).
- Crossref
- Scopus (115)
- Google Scholar
,
,
9
- Krams I.
- Krama T.
- Igaune K.
Alarm calls of wintering great tits Parus major: Warning of mate, reciprocal altruism or a message to the predator?.
- Crossref
- Scopus (41)
- Google Scholar
]. Being individually discriminated is more than hard in larger groups, and large group size may select for increased individuality–signature data–in social signals, to facilitate discrimination [
4
- Medvin M.B.
- Stoddard P.Thou.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and befouled swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
,
,
,
]. Small-scale studies propose that more social species have greater individuality in their social signals, such as contact calls [
four
- Medvin M.B.
- Stoddard P.1000.
- Beecher K.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
]. However, this relationship has not been evaluated in a broader-calibration evolutionary context or in social signals other than contact calls. Information technology is not yet known whether social group size may be viewed every bit a general evolutionary driver of individuality. Here nosotros show a strong positive evolutionary link between social group size in sciurid rodents and individuality in their social alert calls. Social grouping size explained over 88% of the variation in vocal individuality in phylogenetic independent contrasts. Species living in larger groups, but non in more complex groups, had more signature information in their calls. Our results propose that social grouping size may promote the evolution of individual signatures and that the sociality-individuality relationship may be a general phenomenon in nature.
Highlights
- Individual recognition is of import for social behaviors in humans and other animals
- Private recognition is harder in larger groups, simply increased individuality helps
- In evolutionary analyses, social group size predicts individuality in social signals
- Social group size promotes the evolution of individualistic signatures
Results
Discriminating among individuals is of import for social animals [
,
,
seven
- Hare J.F.
- Atkins B.A.
The squirrel that cried wolf: Reliability detection by juvenile Richardson'south ground squirrels (Spermophilus richardsonii).
- Crossref
- Scopus (115)
- Google Scholar
,
], and it oftentimes benefits individuals not only to discriminate amidst or between others but also to be discriminated past others [
,
,
4
- Medvin M.B.
- Stoddard P.G.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
v
- Hare J.F.
Juvenile Richardson's footing squirrels, Spermophilus richardsonii, discriminate amongst individual alarm callers.
- Crossref
- PubMed
- Google Scholar
,
,
,
9
- Krams I.
- Krama T.
- Igaune K.
Warning calls of wintering great tits Parus major: Warning of mate, reciprocal altruism or a message to the predator?.
- Crossref
- Scopus (41)
- Google Scholar
]. Social animals feel selective pressure level to clearly signal their individual identity to others, a job that becomes more difficult as the number of signalers increases [
,
]. When discrimination at the individual level is of import to a species' social behaviors, the amount of individualistic data in their signals must be sufficient to permit discrimination amongst the individuals that routinely interact socially using these signals. We therefore wait that individuality should evolve with the size of a species' typical social group (offset proposed in [
], see as well [
4
- Medvin 1000.B.
- Stoddard P.K.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative assay of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
,
,
]). Previous comparisons across pairs or modest numbers of species take indicated a relationship between communal crèche size and individuality in parent-offspring contact calls [
4
- Medvin M.B.
- Stoddard P.Yard.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
], simply a broader-scale evolutionary relationship between social grouping size and individuality has not yet been tested. Here we present the first phylogenetically controlled independent-contrasts study of the human relationship betwixt individuality and social group size, and the first to examine social signals other than contact calls.
To test the hypothesis, we calculated the corporeality of individuality contained in a social signal, vocal alarm calls, and compared this across eight species of sciurid rodents that vary in social group size and social construction complexity (Figure i). Footing-dwelling sciurids have individualistic alarm calls, are recognized past group mates based on these calls, and stand to reap fitness benefits from being recognized ([
five
- Hare J.F.
Juvenile Richardson'south ground squirrels, Spermophilus richardsonii, discriminate amongst private alarm callers.
- Crossref
- PubMed
- Google Scholar
,
7
- Hare J.F.
- Atkins B.A.
The squirrel that cried wolf: Reliability detection past juvenile Richardson'south ground squirrels (Spermophilus richardsonii).
- Crossref
- Scopus (115)
- Google Scholar
,
,
,
]; see besides the Supplemental Information bachelor online).
Individuality occurs when interindividual variation in a signal exceeds intraindividual variation in that betoken. This individuality can be quantified using information theory [
]. We used Beecher'south information statistic, Hs [
4
- Medvin One thousand.B.
- Stoddard P.K.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
,
,
,
17
- Pollard K.A.
- Blumstein D.T.
- Griffin South.C.
Pre-screening audio-visual and other natural signatures for utilise in noninvasive private identification.
- Crossref
- Scopus (21)
- Google Scholar
], to quantify the vocal individuality present in the alarm calls and compared this with group size data taken from the literature. As predicted, individuality was positively related to social grouping size (R 2 = 0.888, y = 0.572x + 1.628, p < 0.001, north = viii; Effigy 2A ) in the raw data.
Figure 2 Social Group Size versus Vocal Individuality for Eight Species of Basis-Home Sciurids
Show full caption
(A) Group size versus individuality (raw data; due north = 8). Data points are labeled by species: C. leucurus (CYLE), C. ludovicianus (CYLU), Thou. flaviventris (MAFL), M. olympus (MAOL), S. beecheyi (SPBE), S. beldingi (SPBL), S. richardsonii (SPRI), S. tridecemlineatus (SPTR).
(B) Group size versus individuality (independent-contrasts data, positivized; n = 7).
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Nosotros used phylogenetic contained contrasts and regression to control for phylogeny and identify the correlated evolution betwixt individuality and group size. As predicted, contrasts of social group size were significantly correlated with contrasts of individuality (R 2 = 0.887, y = 0.519x, p < 0.001, n = 7 independent contrasts; Figure 2B).
Amongst species living in stable social groups, we expected grouping size to be the major predictor of evolutionary changes in individuality. However, animate being species differ not but in their grouping size but also in how their groups are structured, and we wanted to control for this. Social structure complexity may be quantified in a variety of means [
,
,
,
21
- Michener G.R.
Kin identification, matriarchies, and the evolution of sociality in ground-dwelling sciurids.
- Google Scholar
] and may influence individuality. The social reproductive complexity of paper wasps influences facial individuality [
]. In sciurids, social complication influences the size of song repertoires [
], simply its influence on individuality, if any, is unknown. Nosotros therefore controlled for three measures of social structure complexity [
,
,
21
- Michener G.R.
Kin identification, matriarchies, and the evolution of sociality in ground-dwelling house sciurids.
- Google Scholar
] in our study. Group size and social structure complexity were non intercorrelated in our data (see below) and are not necessarily correlated in nature [
].
We regressed individuality separately against social group size, Armitage's sociality index [
], Michener's social grade [
21
- Michener G.R.
Kin identification, matriarchies, and the evolution of sociality in ground-dwelling sciurids.
- Google Scholar
], and Blumstein and Armitage'south social complexity index [
]. In the raw data, individuality was unrelated to social course (R two = 0.148, y = −0.246x + six.856, p = 0.347), sociality alphabetize (R 2 = 0.375, y = −0.473x + 7.536, p = 0.107), or social complexity index (R two = 0.177, y = −0.901x + six.796, p = 0.300) (north = eight in all cases). We performed multiple regressions with backward emptying to determine whether each alphabetize of social structure complexity interacted with social group size to influence individuality and/or explained additional variation. The social complication indexes and whatsoever interactions containing them were not meaning and were dropped from each model until but social grouping size remained.
We performed the aforementioned regressions on independent contrast information to command for effects of phylogeny. Individuality was non evolutionarily correlated with whatever social complication index (Armitage'due south sociality index [
]: R 2 = 0.224, y = −0.378x, p = 0.236; Michener'southward social form [
21
- Michener Thou.R.
Kin identification, matriarchies, and the development of sociality in basis-dwelling sciurids.
- Google Scholar
]: R 2 = 0.095, y = −0.191x, p = 0.457; Blumstein and Armitage's social complexity alphabetize [
]: R 2 = 0.076, y = −0.669x, p = 0.509) (n = 7 contained contrasts in all cases). As with the raw data, indexes of social structure complication dropped out of multiple regressions in which they were entered, leaving only social group size.
Give-and-take
Animals often do good by discriminating individuals [
,
,
7
- Hare J.F.
- Atkins B.A.
The squirrel that cried wolf: Reliability detection by juvenile Richardson'south footing squirrels (Spermophilus richardsonii).
- Crossref
- Scopus (115)
- Google Scholar
,
] and by signaling their private identity to others [
,
5
- Hare J.F.
Juvenile Richardson'south ground squirrels, Spermophilus richardsonii, discriminate among individual alarm callers.
- Crossref
- PubMed
- Google Scholar
,
]. For example, territory holders, socially ranked individuals, parents, offspring, mates, coalition partners, and alarm signalers benefit past being recognized by their neighbors, grouping members, kin, mates, or companions [
,
four
- Medvin M.B.
- Stoddard P.K.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative assay of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
five
- Hare J.F.
Juvenile Richardson's basis squirrels, Spermophilus richardsonii, discriminate among individual alarm callers.
- Crossref
- PubMed
- Google Scholar
,
,
,
9
- Krams I.
- Krama T.
- Igaune K.
Alarm calls of wintering great tits Parus major: Alert of mate, reciprocal altruism or a message to the predator?.
- Crossref
- Scopus (41)
- Google Scholar
]. This recognition utilizes individualistic signatures (individuality) in animals' social signals. Beingness discriminated from other conspecifics is more hard in larger groups because in that location are more than distracters and targets to exist sifted through. A greater degree of individuality is thus required for successful bigotry in larger groups [
4
- Medvin M.B.
- Stoddard P.K.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
,
,
,
] and should evolve when individually discriminating animals increase their group size on an evolutionary timescale. Our results support this expectation. In both the raw data and independent contrasts, typical social group size explained over 88% (p < 0.001) of the variation in vocal individual distinctiveness across species. It thus seems likely that selection for individuality is stronger in species that must routinely interact with more individuals.
Individuality in Other Signals and Other Taxa
Our study's hypothesis [
4
- Medvin M.B.
- Stoddard P.Grand.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and befouled swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
,
,
,
] predicts that the number of individuals that must be discriminated (group size) should positively touch the amount of individuality in signals used for discrimination, provided that signalers do good from being discriminated. These conditions are met in a diversity of signal contexts [
,
,
4
- Medvin Yard.B.
- Stoddard P.K.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
5
- Hare J.F.
Juvenile Richardson'due south ground squirrels, Spermophilus richardsonii, discriminate among individual alarm callers.
- Crossref
- PubMed
- Google Scholar
,
,
,
ix
- Krams I.
- Krama T.
- Igaune K.
Alert calls of wintering slap-up tits Parus major: Warning of mate, reciprocal altruism or a message to the predator?.
- Crossref
- Scopus (41)
- Google Scholar
]. Individuality has been found in various social signals, and the degree of individuality has been plant to chronicle to social group size in avian and chiropteran contact calls [
4
- Medvin M.B.
- Stoddard P.Thousand.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
] and in rodent warning calls (this written report). We wait a positive evolutionary relationship betwixt group size and individuality to be common in other social communicative signals, and in other taxa as well.
Our group size-individuality hypothesis is relevant to taxa in which social groups are relatively stable and individual discrimination among grouping members is important. Some animal species have a bureaucracy of nested group levels (e.thousand., fission-fusion) or may exhibit unlike types of social interaction (e.thou., solitary animals with contiguous range borders). Animals may use different social signals to communicate at these different levels. For a given social bespeak, the relevant level of group should provide the selective force per unit area for individuality in those signals. For instance, the size of a communicating "group" of lone-living animals would likely predict the extent of individualistic information in their territorial marks. The caste of individuality in a given signal could furthermore provide clues to the social group level at which the indicate is typically used. The human relationship should hold even if simply i or a few receivers must discriminate among multiple signalers: for example, the number of customer fish visiting the aforementioned cleaner station may predict the individuality in their bodily markings. Such relationships remain to be tested.
We do not wait the grouping size-individuality human relationship to hold for all animal taxa. For animals in relatively fluid or impersonal aggregations (e.1000., some ungulate herds, fish schools, or insect swarms), individual interaction is less important and group membership is unstable. In such cases, it may be less important to discriminate individuals. Eusocial species also may not benefit as much from discrimination at the individual level; group membership bigotry may be more critical. The relationship between group size and individuality is less probable to hold for such situations, but this remains to exist tested.
Sociality every bit a Driver of Individuality
Signaling or attending to private identity is important in a wide range of social chatty encounters in many taxa, and it hinges on the social importance of attention to underlying individual differences such as behavioral tendencies, past or continuing condition, or past actions (e.k., [
,
]). Signalers should indicate their individual identity whenever they do good by having their behavioral tendencies, personal history, etc. known by others [
]. Such a socioecological environs exists for a range of animal species, including humans and other primates, marine mammals, social rodents, social carnivores, equines, elephants, and many birds. We wait that the individuality-sociality human relationship may exist a widespread miracle in nature.
Larger social groups crave more intricate individual signatures for effective discrimination, and our results implicate sociality as a major predictor of individuality. The bigger the crowd, the more information technology takes to stand out.
Experimental Procedures
Data Collection
Eight species of footing-home sciurid rodents were selected to represent a range of grouping sizes and social complexity levels, as well every bit phylogenetic diverseness (Figure 1; Table ane). All eight species live in social groups and produce vocal alarm calls in response to predatory stimuli. Refer to Supplemental Information for details on why individuality is of import in alarm signals. Brute procedures conformed to all regulatory standards and were canonical by the animal care and use committees of the Academy of California, Los Angeles; University of California, Santa Barbara; University of Montana; and University of Manitoba. We captured animals using alive traps baited with peanut butter, horse feed, and/or fruit. If the animal vocalized in the trap when the researcher approached, these calls were recorded. The animals were so weighed, sexed, and individually marked with numbered ear tags and/or pilus dye, then released at the capture site. Warning calls were elicited past man presence (run across [
]). The researcher walked toward the focal animal and recorded calls that the animal produced in response to being approached. When possible, nosotros recorded calls from individuals within traps because this ensured high recording quality. Some species practice not normally alarm phone call in alive traps; these species were given individualistic dye markings, released, and later approached on foot to arm-twist and record calls (Supplemental Information).
Table i Location and Recording Information for Each Study Species
| Species | Recorded by | Year | Site | Location | Approximate Coordinates |
|---|---|---|---|---|---|
| C. leucurus | K.A.P. | 2006 | Arapaho National Wildlife Refuge | Jackson County, CO | twoscore° 37′ N, 106° 16′ W |
| C. ludovicianus | Chiliad.A.P. | 2006 | UL Curve National Wild animals Refuge | Phillips County, MT | 47° 56′ N, 107° 52′ West |
| M. flaviventris | D.T.B. | 2001, 2002, 2003 | Rocky Mountain Biological Laboratory | Gunnison County, CO | 38° 55′ N, 106° 60′ W |
| Yard. olympus | S.C. Griffin | 2004, 2005 | Olympic National Park | Clallam County, WA | 47° 49′ N, 123° thirteen′ West |
| South. beecheyi | K.A.P. | 2006, 2008 | Sedgwick Reserve (Academy of California Natural Reserve System) | Santa Barbara County, CA | 34° 42′ N, 120° two′ W |
| S. beldingi | G.A.P. | 2007 | Rock Creek Coulee/Lower Equus caballus Corral | Mono County, CA | 37° 25′ N, 119° 0′ W |
| Southward. richardsonii | J.F. Hare | 2006 | Assiniboine Park Zoo belongings | Winnipeg, MB | 49° 52′ N, 97° 14′ W |
| S. tridecemlineatus | K.A.P. | 2005 | Shortgrass Steppe Long-Term Ecological Research Station | Weld County, CO | 40° 44′ N, 104° 44′ Westward |
- Open up tabular array in a new tab
Compared to other aspects of sociality (e.g., [
,
21
- Michener Chiliad.R.
Kin identification, matriarchies, and the development of sociality in footing-domicile sciurids.
- Google Scholar
,
,
]), we expected social group size to have the largest outcome on the demand for individuality in social signals such as warning vocalizations (encounter as well [
]). The number of individuals that alive together, and hence the number of individuals that must be routinely discriminated, is the major cistron affecting the difficulty of bigotry tasks and the corporeality of individualistic information that an fauna must produce to be discriminated [
4
- Medvin M.B.
- Stoddard P.K.
- Beecher Thousand.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
]. Individuals within the social grouping are those well-nigh often in shut proximity and those heard most often, and are presumed to be intended recipients of signals (such every bit alarm calls) given by group members. Species-typical social group size (Tabular array two) was calculated from demographic data (Supplemental Information). Social groups are defined by common cooperation and/or amicability (burrow sharing, alert calling, aid during chases, and/or cooperative defense of territories and immature). Boundaries between such social groups are easily identified by increased agonism, territoriality, and/or lack of influence on alarm calling rates. Social groups are labeled by different terms in unlike taxa (Tabular array two; Supplemental Information).
Table 2 Individuality, Group Size, and Social Complexity Indices for Eight Sciurid Species
| Species | Sociality Alphabetize [ xx
Sociality as a life-history tactic of ground squirrels.
| Social Form [ 21
Kin identification, matriarchies, and the evolution of sociality in ground-dwelling house sciurids.
| Social Complication [ eighteen
Does sociality bulldoze the evolution of communicative complexity? A comparative test with ground-dwelling sciurid alarm calls.
| Hs | Group Size | Type of Group | Group Size Sources |
|---|---|---|---|---|---|---|---|
| C. leucurus | iii | 2 | 0.84 | v.05 | half-dozen.42 | association | [ 30
Some comparisons of black-tailed and white-tailed prairie dogs in north-key Colorado.
31
Observations on reproduction in the white-tailed prairie-dog (Cynomys leucurus).
32
Breeding biology of the white-tailed prairie dog, Cynomys leucurus, in Wyoming.
|
| C. ludovicianus | 5 | five | 1.12 | 4.89 | 6.19 | coterie | [ 30
Some comparisons of black-tailed and white-tailed prairie dogs in north-fundamental Colorado.
33
The Black-Tailed Prairie Dog: Social Life of a Burrowing Mammal.
|
| M. flaviventris | 4 | 4 | one.06 | five.79 | vi.35 | matriline | [ 19
Life history consequences of social complication: A comparative study of ground-dwelling sciurids.
34
Social and population dynamics of yellow-bellied marmots: Results from long-term research.
35
A examination of the social cohesion hypothesis: Interactive female marmots remain at habitation.
36 Olson, 50.E. (2009). Male person social behavior in a facultatively social rodent, the yellow-bellied marmot (Marmota flaviventris). PhD thesis, Academy of California, Los Angeles, Los Angeles, California.
|
| M. olympus | four | five | 1.46 | 6.45 | 8.84 | colony | [ 37
The social biology of the Olympic marmot.
|
| Southward. beecheyi | 2 | 2.5 | 0.26 | vi.26 | vii.70 | family | [ 38
A population study of the Beechy footing squirrel in central California.
39
Reproductive behaviour and multiple paternity of California footing squirrels.
twoscore
Home range, population construction, and spatial organisation of California ground squirrels.
|
| South. beldingi | 2 | 2 | 0.40 | 7.76 | x.23 | kin group | [ 41
Demography of Belding's basis squirrels.
|
| South. richardsonii | 2 | two | 0.39 | half-dozen.00 | 7.63 | family unit | [ 42
Variation in litter size: A test of hypotheses in Richardson's ground squirrels.
43
Reproduction of Richardson's footing squirrel (Spermophilus richardsonii) in southern Saskatchewan.
|
| S. tridecemlineatus | 2 | 1.v | 0.l | half-dozen.74 | 9.39 | family | [ 44
A life history report of thirteen-lined ground squirrels in southern Wisconsin.
45
Notes on a population of the striped ground squirrel (Citellus tridecemlineatus) in an abandoned field in southeastern Michigan.
46
Annual wheel, population dynamics and adaptive behavior of Citellus tridecemlineatus.
47
Breeding habits and early on life of the thirteen-striped footing squirrel, Citellus tridecemlineatus (Mitchill).
|
Hsouth is Beecher'southward information statistic.
- Open tabular array in a new tab
Acoustic Processing and Measurements
Nosotros measured 20 high-quality calls each from ten individuals per species, for a total of 1600 calls. To command for potential bout-specific phone call traits, we used calls from at least 2 distinct calling bouts per individual, with no single tour contributing more than 10 calls. We considered bouts singled-out if they were separated by at to the lowest degree 20 minutes of noncalling during which the fauna resumed normal, nonvigilant behavior (e.g., feeding, grooming). Bouts were typically separated by a few days.
We either recorded straight to digital files or digitized call recordings to 16-bit 48 or 44.1 kHz .aif files, then normalized each call to 95% maximum amplitude. Using consistent settings, nosotros made a series of measurements in the time, frequency, and relative amplitude domains (Supplemental Information).
Data Analysis
Nosotros standardized the audio-visual data according to Beecher [
] before computing principal components (Supplemental Information). We calculated Beecher's data statistic (Hsouth, [
,
]) for each species from the main components, using all F values that were pregnant at the α = 0.05 level. Beecher'southward statistic is a standard measure used to quantify individuality [
4
- Medvin M.B.
- Stoddard P.K.
- Beecher Yard.D.
Signals for parent-offspring recognition: A comparative assay of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
,
,
17
- Pollard K.A.
- Blumstein D.T.
- Griffin S.C.
Pre-screening acoustic and other natural signatures for use in noninvasive private identification.
- Crossref
- Scopus (21)
- Google Scholar
]. Information technology expresses a indicate'southward ability to reduce a receiver's initial uncertainty about the identity of the signaler down to a minimum level (the inside-individual error) and can quantify individuality beyond disparate characteristics and modalities. With consistent sampling, the data statistic can be directly compared across species or signal types [
iv
- Medvin M.B.
- Stoddard P.Thou.
- Beecher M.D.
Signals for parent-offspring recognition: A comparative analysis of the begging calls of cliff swallows and barn swallows.
- Crossref
- Scopus (68)
- Google Scholar
,
17
- Pollard G.A.
- Blumstein D.T.
- Griffin Due south.C.
Pre-screening audio-visual and other natural signatures for use in noninvasive individual identification.
- Crossref
- Scopus (21)
- Google Scholar
]. The more bits of individualistic data in a betoken, the easier it is to discriminate individuals, and the more total individuals can be reliably discriminated [
,
].
We calculated independent contrasts (Supplemental Information) using phylogenetic tree hypotheses from Harrison et al. [
25
- Harrison R.G.
- Bogdanowicz S.Grand.
- Hoffmann R.Southward.
- Yensen E.
- Sherman P.Due west.
Phylogeny and evolutionary history of the basis squirrels (Rodentia: Marmotinae).
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] and Herron et al. [
]. We chose Felsenstein'south method [
] considering it is a standard phylogenetic tool used in evolutionary studies of morphological, behavioral, and social traits, including grouping size (see [
28
- Beauchamp Thousand.
- Fernández-Juricic E.
Is there a human relationship betwixt forebrain size and group size in birds?.
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,
]).
Acknowledgments
We thank Suzanne Griffin for recordings of Olympic marmots and James Hare for recordings of Richardson's ground squirrels. We thank Olivier Munos for measurements of xanthous-bellied marmot calls. For admission to sites and animals, nosotros thank Abby Benson, Dean Biggins, David Eads, John Hoogland, Jill Mateo, Dan Salkeld, Arapaho National Wildlife Refuge, Assiniboine Park Zoo, Olympic National Park, Stone Creek and the Sierra Nevada Aquatic Research Laboratory, Rocky Mountain Biological Laboratory, Sedgwick Reserve, Shortgrass Steppe Long-Term Ecological Research Station, UL Bend National Wildlife Refuge, and their personnel. We give thanks our laboratory and field assistants: Dalal Farsakh, Erik Tanaka, Nina Beri, Anh Thu Tran, Aya Satoh, Nina Mahoney, Yeshika Sharma, Asal Askarinam, John Mina, Aaron Nichols, Facundo Bermudez, Pontea Niaki, Christine Guenzi, Hao Li, Kathy Constabulary, Rebecca Lee, Shabnum Patel, Liz Devietti, Rashad Mansour, Sahar Askarinam, Grace Li, and Cory Pender. We thank Peter Nonacs, Greg Grether, Joan Silk, Ben Rossi, and reviewers for comments on previous versions of this manuscript. K.A.P. was funded past the American Philosophical Society, the American Society of Mammalogists, the Animal Behavior Society, the Explorers Lodge, Mildred Mathias/UC Reserves, the National Science Foundation's Graduate Research Fellowship Program, Sigma Xi, a UCLA George Bartholomew Inquiry Fellowship, a UCLA Holmes O. Miller Fellowship, and a UCLA Quality of Graduate Instruction Fellowship.
Supplemental Information
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Article Info
Publication History
Published online: February 17, 2011
Accepted: January 20, 2011
Received in revised class: December 9, 2010
Received: September 29, 2010
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DOI: https://doi.org/10.1016/j.cub.2011.01.051
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© 2011 Elsevier Ltd. Published by Elsevier Inc.
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