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Form and Function of the Musteloids

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Form and Function of the Musteloids chapter 3 Form and function of the musteloids Andrew C. Kitchener, Carlo Meloro, and Terrie M. Williams Fisher skull © Neil McLean, National Museums Scotland Introduction with short legs. This common body plan is adapted to three main functional types: terrestrial/fossorial With a fossil history dating back 30–32 million years (e.g. weasels, stoats, ferrets; see Chapters 10 and 15, ago (late Eocene and early Oligocene), the Muste- this volume), scansorial/arboreal (e.g. martens; see loidea are now the most successful superfamily of Chapter 12, this volume), and aquatic/semi-aquatic the Carnivora, as shown by an abundance of species (e.g. mink and otters; see Chapters 16, 17, 22, and with a global distribution (Agnarsson et al. 2010; 23, this volume), although many species are not Sato et al. 2012). Ranging in size from tiny weasels restricted to one of these functional types, for ex- (Mustela nivalis, which may weigh 30 g or less) to ample, martens (Martes spp.) may be terrestrial. Ex- huge specialized sea otters (Enhydra lutris, >30 kg), ceptions to this rule, the bear-like wolverines (Gulo they may not have evolved to be as physically large gulo, see Chapter 18, this volume) and badgers (see as members of other carnivoran families, but the ra- Chapters 13, 19, and 21, this volume), have shorter, tio of the smallest to largest is the greatest—a mas- robust bodies and relatively longer legs compared sive 1000:1 in body masses (Table 3.1). With around with those of other mustelids. However, mustelids 90 species, the Musteloidea is the most speciose are not equally successful globally. Although about superfamily (c.30% of carnivorans) and the Mus- 43% of smaller carnivoran species in the Americas telidae (>60 species; c.20% of carnivorans) the most and the Oriental region are mustelids, they com- speciose family of the Carnivora (Wilson and Mit- prise a mere 12% of smaller carnivoran species in termeier 2009; Hunter 2011; see Chapters 1 and 2, Africa (Figure 3.1). Despite colonizing Africa from this volume). the Miocene alongside viverrids and herpestids The mustelids are mostly specialized predators (Hunt 1996), the only mustelids, apart from ot- that generally have a thin, elongated body plan ters, south of the Sahara all have black-and-white Kitchener, A. C., Meloro, C., and Williams, T. M., Form and function of the musteloids. In: Biology and Conservation of Musteloids. Edited by David W. Macdonald, Christopher Newman, and Lauren A. Harrington: Oxford University Press (2017). © Oxford University Press. DOI 10.1093/oso/9780198759805.003.0003 03-Macdonald-Chap03.indd 98 31/05/17 7:03 PM FORM AND FUNCTION OF THE MUSTELOIDS 99 aposematic coloration (e.g. honey badger [Mellivora three to four species, differentiated by size and diet capensis], African striped polecats [ Ictonyx spp.], (usually fish versus crustaceans), per biogeograph- and African striped weasels [Poecilogale albinucha]), ical region (see Chapter 1, this volume). backed up by anal sacs producing noxious secre- The other musteloid families, skunks (Mephiti- tions—perhaps the mustelids arrived a little later dae; see Chapters 24 and 25, this volume), raccoons, and only those mustelids packed with chemical coatis, kinkajou (Procyonidae; see Chapters 26, 27 weapons could muscle their way in. In contrast, and 28, this volume), and red pandas (Ailuridae; in the Palaearctic all small carnivorans are mus- see Chapter 29, this volume), are rather more eclec- telids with the possible exceptions of one or two tic and comprise fewer species, although procyo- marginal herpestids and viverrids, which may re- nids show greater diversity of form than mustelids, sult from human introductions. Otters are almost if they do indeed form a monophyletic family unique among carnivorans (the only exceptions be- (Koepfli et al. 2007; Koepfli et al., Chapter 2, this ing otter civets [Cynogale bennetti], and aquatic gen- volume). In contrast to the other families, red pan- ets [Genetta piscivora], among viverrids) in being das (Ailurus fulgens) have evolved as largely spe- adapted to aquatic environments; there are usually cialized vegetarians that show many evolutionary Table 3.1 Body size of smallest and largest members of each musteloid family. Family Mustelidae Mephitidae Procyonidae Ailuridae Number of species >60 11 14 1 Largest species Enhydra lutris Conepatus leuconotus Procyon lotor Ailurus (f.) styani Body mass (g) 45,000 4500 28,000 7500 HB (mm) 1200 510 620 730 Smallest species Mustela nivalis Spilogale pygmaea Bassariscus astutus Ailurus (f.) fulgens Body mass (g) 25 130 800 4500 HB (mm) 114 190 305 510 Ratio body masses 1800 34.6 35 1.7 Ratio HBs 10.5 2.7 2 1.4 100% 90% 80% 70% 60% 50% Non-mustelids 40% Mustelids 30% 20% 10% Figure 3.1 Proportion of small 0% carnivorans that are mustelids in different Nearctic Neotropical Oriental African Palaearctic biogeographical regions. 03-Macdonald-Chap03.indd 99 31/05/17 7:03 PM 100 BIOLOGY AND CONSERVatION OF MUSTELOIDS parallels to the unrelated giant panda (Ailuropoda (Isaac 2005; Lindenfors et al. 2007). Sexual size di- melanoleuca). Procyonids and mephitids dominate morphism may be a consequence of sexual selection the Americas, but tend to be less carnivorous and in polygynous species for larger males that compete more omnivorous than sympatric mustelids. for access to oestrous females. Females may also In this chapter we review the form and func- be smaller than males so that their total energy re- tion of the musteloid carnivorans to discover the quirements are less than for males and so they can key adaptations that have helped the musteloids invest more energy in the development of young. in general and the mustelids in particular achieve Sexual dimorphism may also allow for a degree global success in terms of numbers of species and of niche separation between the sexes so that each geographical spread (see Macdonald et al., Chap- sex can prey on animals of different sizes, thereby ter 1, this volume). We begin with a discussion of minimizing competition. Clearly there is interplay primarily terrestrial and arboreal species, followed between these factors and considerations of popu- by specific adaptations for those species that live lation density, social behaviour, home-range size, partially or fully in aquatic environments. length of oestrus and energy use (Ferguson and Larivière 2004). There is not space here to discuss Sexual dimorphism further the causes and consequences of differential sexual dimorphism in musteloids but the theme is It is very apparent from external appearances that developed further in Macdonald et al., Chapter 6, there is a wide degree of sexual dimorphism in this volume, and in Noonan et al. (2016). musteloids. Although much has been written about Working from the outside in, let us take a closer the causes of sexual dimorphism in musteloids look at the adaptations of musteloids. (especially mustelids), there has been little inves- tigation as to the degree of sexual dimorphism in Hiding in plain sight—pelage coloration different musteloid groups. and markings Therefore, based on data in the literature, we es- timated sexual dimorphism in mean condylobasal Pelage has two main functions. First, pelage colora- skull lengths and mean body weights of the Melinae tion and markings enable many musteloids to re- (badgers), Mustelinae (polecats, weasels, and mar- main hidden from predators and prey. For example, tens), Lutrinae (otters), Mephitidae (skunks) and weasels and stoats (Mustela spp.) at high latitudes Procyonidae (raccoons and coatis) (Figure 3.2). The moult into white winter fur to match the snow in degree of sexual dimorphism in condylobasal skull contrast to the brown summer pelage. However, length (CBL) is less than that in body weight. This some markings may function in communication is not surprising since the former is a linear measure within and between species (see Buesching and whereas the latter is a cubic measure. However, even Stankowich, Chapter 5, this volume). Second, the so the pattern is consistent with both measures. The pelage ensures that musteloids are insulated to Mustelinae show a much higher mean sexual di- avoid excessive heat loss to (or gain from) the sur- morphism (1.71 body weight, 1.10 CBL) than other rounding environment. This is especially important musteloid groups, which tend to display similar de- for weasels and stoats in colder northern latitudes, grees of sexual dimorphism (1.3–1.38 body weight; and also otters living in the sea and exposed to cold 1.06–1.07 CBL) with the melines showing the least water for prolonged periods, owing to the high sur- sexual dimorphism (1.22 body weight, 1.05 CBL). face area to volume ratios of their elongated bodies What is striking from this preliminary analysis is (Kruuk 2006; King and Powell 2007). that the most carnivorous terrestrial musteloids, Most musteloids have a uniform coloration either the mustelines, show the highest degree of sexual all over or dorsally, with typically paler coloration dimorphism. ventrally, providing counter-shading to remain Causes of sexual dimorphism among carnivorans hidden in a wide variety of habitats. Although are often discussed and centre around the follow- most are brownish and dark, some, for example, ing themes, which may not be mutually exclusive Siberian weasel (Mustela sibirica), Malay weasel 03-Macdonald-Chap03.indd 100 31/05/17 7:03 PM FORM AND FUNCTION OF THE MUSTELOIDS 101 (a) 1.12 1.11 1.1 1.09 1.08 1.07 1.06 1.05 1.04 1.03 condylobasal skull lengths Ratio of male to female mean 1.02 1.01 Melinae Mustelinae Lutrinae Mephitidae Procyonidae (b) 1.8 1.6 n 1.4 1.2 1 0.8 body weights 0.6 0.4 Ratio of male to female mea 0.2 0 Melinae Mustelinae Lutrinae Mephitidae Procyonidae Figure 3.2 Sexual dimorphism in musteloids.
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