M Mustelid Communication Communication in mustelids is closely related and dependent on the species’ respective social Christina A. S. Mumm1 and Mirjam Knörnschild system and habitat (Buesching and Stankowich 1,2,3 2017). The social and spatial organization of 1Animal Behavior Lab, Free University of Berlin, mustelids is highly variable between species, but Berlin, Germany there may be also variation in different 2Smithsonian Tropical Research Institute, populations of the same species (Kruuk 2006; Ancón, Panamá Newman et al. 2011). Divergence of a pure 3Museum für Naturkunde – Leibniz Institute for solitary lifestyle is induced by environmental fac- Evolution and Biodiversity Science, tors such as habitat, geographical distribution, Berlin, Germany ecomorphology, resource dispersion, or season (Johnson et al. 2000). The socio-spatial organiza- tion of mustelids is based on the “classical” system of one male territory encompassing Synonyms distinct female territories. Several variations of and exceptions from this basic pattern exist as Acoustic communication; Badgers; Communica- well (Kruuk 2006). tion; Martens; Modality; Multimodal; Mustelids; As knowledge on mustelids accumulates, sev- Olfaction; Otters; Scent; Sensory channel; Signal; eral species which formerly have been described Social organization; Sociality; Vision; Vocaliza- as purely solitary turn out to be more social, or tion; Weasels have at least a much more flexible social lifestyle (e.g., Newman et al. 2011). Each communication channel has its advan- Introduction tages and disadvantages. One may be more useful than the others, depending on when, where, and Mustelids are a large family, with 60 species which information needs to be transmitted. Com- found in Eurasia, Africa, Asia, North and South pared to acoustic signals, chemical markings per- America (Koepfli et al. 2008). As shown in sist for an elongated time, thereby allowing Table 1, the family contains eight subfamilies, conspecifics or co-occurring species to gather with many fascinating species like the European information even in the physical absence of badger (Meles meles), the sea otter (Enhydra the sender. In a habitat where olfactory cues can- lutris), and the wolverine (Gulo gulo), to name not be deposited, acoustic communication may be but a few. favored (Fig. 1). This is especially true for the # Springer International Publishing AG 2018 J. Vonk, T. K. Shackelford (eds.), Encyclopedia of Animal Cognition and Behavior, https://doi.org/10.1007/978-3-319-47829-6_1191-1 2 Mustelid Communication Mustelid Communication, Fig. 1 Mustelid communication in three sensory channels. Olfactory communication: two examples of scent-marking postures; acoustic communication: spectrograms of vocalizations showing frequency over time; visual communication: throat patterns likely used for visual (Picture credits a, c, d, f, and g by Christina Mumm, b and j by Pia Weidenmüller (written license), e by Hannah Heither (written license), h by user 422737 on pixabay (CC0 license), i by user Ellis Lawrence on flickr (CC BY-SA 2.0 license). Recordings: c by Christina Mumm, d by Tierstimmenarchiv Berlin) Mustelid Communication 3 Mustelid Communication, Table 1 Subfamilies and number of genera and species in the family of mustelids. Each genus is shown with one exemplary species. The number of all genera and species is given in the subheadings for each subfamily (genera/species). All subfamilies are depicted with one representative species. Taxonomy following Koepfli et al. 2008 Subfamily: Galictinae (4/6) Galictis Greater grison (G. vitatta) Ictonyx Striped polecat (I. striatus) Poecilogale African striped weasel (P. albinucha) Vormela Marbled polecat (V. peregusna) Marbled polecat Subfamily: Helictidinae (1/5) Melogale Chinese ferret-badger (M. moschata) Chinese ferret-badger Subfamily: Lutrinae (6/13) Aonyx Asian small-clawed otter (A. cinerea) Enydra Sea otter (E. lutris) Lontra North American river otter (L. canadensis) Lutra European otter (L. lutra) Lutrogale Smooth-coated otter (L. perspicillata) Pteronura Giant otter (P. brasiliensis) Eurasian otter Subfamily: Martinae (3/10) Eira Tayra (E. barbara) Gulo Wolverine (G. gulo) Martes European pine marten (M. martes) European pine marten Subfamily: Melinae (2/4) Arctonyx Hog badger (A. collaris) Meles European badger (M. meles) European badger (continued) 4 Mustelid Communication Mustelid Communication, Table 1 (continued) Subfamily: Mellivorinae (1/1) Mellivora Honey badger (M. capensis) Honey badger Subfamily: Mustelinae (3/20) Lyncodon Patagonian weasel (L. patagonicus) Mustela Stoat (M. erminea) Neovison American mink (N. vison) Stoat Subfamily: Taxidiinae (1/1) Taxidea American badger (T. taxus) American badger Picture credits: Marbled polecat by Volker Röhl on Wikipedia; Chinese ferret-badger by Chien-Jen Wang; Eurasian otter by Gellinger on Pixabay; European pine marten by Maxmann on Pixabay; European badger by Chris Parfitt on Flickr; Honey badger by Derek Keats on Flickr; Stoat by Jo Garbutt on Flickr; American badger by Northwest Power and Conservation Council on Flickr subfamily of otters. With the exception of sea detailed information, the animals may vary fre- otters (see below), they show a semiaquatic quency and location of markings. lifestyle and only use scent-marking on land Territorial scent-marking can be done by the (Kruuk 2006). deposition of feces, urine, or a combination of both (i.e., excremental marking, see Fig. 1a), as well as through gland secretion (i.e., secretional Olfactory Communication marking). Secretion may be produced in anal, ventral, foot, or subcaudal glands and applied by Olfactory communication has a multipurpose body rubbing, anal drag, or scratching (see function. It is regarded as the most important Fig. 1b). Gland secretions can also be used to communication channel for mustelids, probably mark conspecifics. In some species, territorial because all mustelids are territorial and scent- scent-marking is enhanced by visually marking mark their territories to announce ownership. It the surroundings, e.g., by scratching the ground further allows mustelids to signal information on or trees (e.g., Hutchings and White 2000; reproductive state, as well as information on iden- Leuchtenberger and Mourão 2009). Otter spraints tity and food resources. To exchange more consist of feces, anal gland secretion, and often of Mustelid Communication 5 a third compound, the “jelly.” Not much is known in badgers. The use of different latrine sites about this slimy substance, the only available depends on population density and season. In information is that it “is secreted somewhere in high-density populations, European badgers the intestine itself” (Kruuk 2006). Chemical com- form social groups, whereas in low-density position varies among feces, urine, and gland populations, groups mainly consist of a pair and secretion, thereby allowing for the transmission their offspring. Larger groups have more latrines of differential information (Clapperton 1989). inside their home range, but space is limited in Deposition of scent is not random in time and high-density populations, so that these groups space (Kilshaw et al. 2009). For scent-marking, usually occupy smaller areas. They invest mainly the animals select prominent places, on stones, in boundary latrines; however, their function in under bridges, or on other landmarks, which are territorial defense is not fully understood frequently revisited (Hutchings and White 2000). (Buesching et al. 2016). Nevertheless, European Otters may even chose marking places which badgers can discriminate between feces from are relatively far and visit them repeatedly neighboring and alien groups and react more (Kruuk 2006). Furthermore, European badgers aggressively towards the scent of alien groups. do not only carefully select marking places, but Smaller badger groups in low-density populations also the positioning of scent inside each place may use larger areas, but their ability of marking is (Stewart et al. 2002). As badger territories contain limited by the number of individuals. Therefore, more than one latrine site, the animals systemati- small groups may neglect boundary latrines and cally revisit several places to refresh their scent the use of hinterland latrines peaks during mating marks (Kilshaw et al. 2009). time. Latrine use is even abandoned seasonally, Social species like European badgers and when home range size is enlarged. This shows a giant otters (Pteronura brasiliensis) make use of trade-off between group size and the area a group communal latrines – enlarged areas for the depo- is able to defend, either in terms of limited space sition of excremental and secretional marks from or in terms of limited number of individuals mark- all group members. Giant otters regularly clear ing the latrines (Buesching et al. 2016). Popula- off the vegetation at communal latrines. Otters tion density also affects the main information in general show an elaborate marking behavior provided at latrines: they likely function to signal because the deposition of excremental and territoriality in high-density populations, but secretional marks can be accompanied by a reproductive state and other individual informa- scent-marking “dance.” Giant otters, as well as tion in low-density populations (Zhou et al. 2015). spotted-necked otters (Lutra maculicollis), and In low-density populations of Japanese badgers North American river otters (Lontra canadensis) (Meles anakuma), information provided at both show dance-like