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Integration or Modularity in the Mandible of Canids (Carnivora: Canidae): a Geometric Morphometric Approach

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Abstract

Understanding the interplay between morphological integration and modularity is considered an important topic in the study of the evolution of the form of complex structures. The mandible is a complex structure that can be shaped by diverse factors such as ontogeny, ecology, and evolutionary history. In canids, this is particularly interesting because they have a large diversity in feeding behavior and hunting strategy. Here, we employed geometric morphometric techniques to evaluate the balance between integration and modularity in 1011 mandibles of a sample of extinct and extant canids. The results show that allometric scaling seems to have little influence in determining the mandibular shape of canids. Some divergence associated with ecology was observed, especially for highly specialized taxa (hypercarnivores and insectivores). Finally, macroevolutionary patterns were more integrated than intraspecific patterns, suggesting that correlational selection might play a strong role in the evolution of mandibular form and function. We found no evidence of an evolutionary line of least resistance in shaping mandible disparity.

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Data Availability

The datasets generated during and/or analyzed during the current study are available from the corresponding author on reasonable request.

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Acknowledgments

We thank to N. Toledo and S. Vizcaíno for inviting us to participate in this tribute to L.B. Radinsky within the framework of the Symposium: El paradigma de correlación forma-función en mastozoología: un tributo a Leonard Radinsky (1937–1985), which took place during the XXXI Jornadas Argentinas de Mastozoología, in La Rioja, Argentina, 25 October, 2018. For permission to access material under their care and the attention in mammal collections we thank Robert Voss, Eileen Westwig (AMNH), Sergio Bogan, Yolanda Davies (CFA), Rubén Barquez, Mónica Díaz (CML), Bruce Patterson (FMNH), Mauro Lucherini, Estela Luengos (GECM), Mauro Schiaffini, Gabriel Martin (LIEB), Andrés Pautasso (MFAZV), Pablo Teta, David Flores, Sergio Lucero (MACN), Diego Verzi, Itatí Olivares (MLP), Mario de Vivo, Juliana Gualda (MZUSP), and Kristofer Helgen, Darrin Lunde (NMNH). This is a contribution to PICT 2014-1930, 2015-2389, 2015-0966, 2016-3151, 2016-3682, PUE 0125 and DEB 1350474 (NSF grant to Liam Revell).

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Authors and Affiliations

  1. Unidad Ejecutora Lillo, Consejo Nacional de Investigaciones Científicas y Técnicas–Fundación Miguel Lillo, Miguel Lillo 251, 4000, San Miguel de Tucumán, Argentina

    Valentina Segura

  2. Consejo Nacional de Investigaciones Científicas y Técnicas (CONICET), Buenos Aires, Argentina

    Valentina Segura, Guillermo H. Cassini, Francisco J. Prevosti & Fabio Andrade Machado

  3. División Mastozoología, Museo Argentino de Ciencias Naturales “Bernardino Rivadavia”, Av. Ángel Gallardo 470, C1405DJR, Ciudad Autónoma de Buenos Aires, Argentina

    Guillermo H. Cassini & Fabio Andrade Machado

  4. Departamento de Ciencias Básicas, Universidad Nacional de Luján, Ruta 5 y Av. Constitución s/n, Luján, 6700, Buenos Aires, Argentina

    Guillermo H. Cassini

  5. Museo de Ciencias Antropológicas y Naturales, Universidad Nacional de La Rioja (UNLaR), Av. Luis M. de la Fuente s/n, 5300, La Rioja, Argentina

    Francisco J. Prevosti

  6. Department of Biology, University of Massachusetts, 100 William T. Morrissey Blvd, Boston, MA, 02125, USA

    Fabio Andrade Machado

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  1. Valentina Segura
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Correspondence to Valentina Segura.

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Appendix 1. List of specimens used in this study

Appendix 1. List of specimens used in this study

Atelocynus microtis (N = 23). AMNH: 76031; 76579; 95284; 95285; 98639; 100095. FMNH: 5249; 57836; 60674; 60675; 60676; 93955; 98080; 98081; 110949; 121286. MZUSP: 4320; 19750; 19751; 19752; 19753; 19754. NMNH: 361013.

Canis aureus (N = 1). MLP: 1035–1031.

Canis latrans (N = 1). MACN: 25.123.

Canis lupus (N = 5). AMNH: 18215. MACN 3.76; 23.15; 35.210. MLP: 1031.

Canis nehringi (N = 1). MACN-PV: 500.

Cerdocyon thous (N = 109). CFA: 3697; 3875; 4265; 4419; 4496; 4511; 4512; 4661; 4663; 4664; 4717; 5048; 5197; 5278; 5283; 5313; 5375; 6000; 6071; 6128; 6129. CML: 588; 3719; 3756; 3827; 4083; 4083; 4692; 5964; 5966; 5967; 6213; 6214; 6340. MACN: 4.213; 20.32; 24.85; 24.127; 25.119; 25.159; 29.839; 30.344; 30.345; 32.261; 32.262; 32.75; 33.6; 34.676; 36.191; 36.481; 39.460; 43.26; 44.11; 45.34; 45.40; 47.116; 47.189; 47.190; 47.191; 47.192; 47.193; 47.402; 48.3; 48.5; 48.6; 48.7; 48.10; 49.221; 49.367; 50.40; 50.43; 50.45; 50.57; 50.59; 50.60; 50.61; 50.62; 50.63; 50.64; 52.54; 52.63; 52.64; 13051; 14,322; 14681; 15741; 16189; 20316; 20454; 20456; 20815; 20816; 20817; 21228; 23180; 23669; 23670; 23726; 23727; 24045; 24046; 24207; 24208. MFA-ZV: 228; 1204. MLP: 20.IX.49.13; 16.X.01.7; 31.XII.02.77; 1322. MZUSP: 9687.

Chrysocyon brachyurus (N = 82). AMNH: 36962; 71179; 120999; 133940; 133941; 135274. CFA: 12826; 12827. CML: 1376; 6133; 6352. FMNH: 28311; 28312; 28313; 44534; 46003; 54406; 96003; 101848; 125401; 127434; 134483; 137425; 150739. MACN: 3.71; 3.73; 4.32; 4.303; 24.4; 30.29; 30.231; 53.49; 13466; 19146; 20646; 23456; 23984; 24043; 24201. MFA-ZV 517; 524; 581; 651; 652; 919; 1166. MLP: 2.IV.02.4; 5.X.99.1; 31.XII.02.88; 6; 92; 564; 695; 1684; 1686. MZUSP: 525; 3025; 3338; 9420; 19733; 19736; 29870; 31981; 32039; 32042; 32043; 32056; 32199; 32505; 32629. NMNH: 196975; 258614; 261022; 261023; 270371; 271567; 314863; 521007; 534807; 534970; 588,223; 588425.

Lycalopex culpaeus (N = 107). CFA: 2129; 6451. CML: 5067; 5068; 5069; 5070; 5071; 5970; 5974; 6343; 6344. LIEB: 791; 793. MACN: 3.68; 4.41; 7.42; 24.119; 25.128; 27.131; 30.69; 31.58; 31.59; 33.67; 33.68; 33.69; 38.39; 41.55; 15024; 15033; 15037; 15040; 15044; 15045; 15049; 15050; 15055; 15062; 15063; 15064; 15073; 15078; 15081; 15082; 15083; 15089; 15093; 15096; 15101; 15106; 15112; 15119; 15121; 15124; 15127; 15138; 15140; 15149; 15151; 15154; 15158; 15163; 15168; 15172; 15173; 15177; 15181; 15882; 15190; 15194; 15196; 15208; 15212; 15220; 15223; 15224; 15226; 15227; 15228; 15229; 15232; 15233; 15240; 15243; 15246; 15248; 19221; 19222; 20813; 21899; 23072; 23076; 23077; 23093; 23095; 23103; 23108; 23119; 23123; 23125; 23148; 23719; 23720; 23721; 23,915; 24210. MLP: 1264; 1266.

Lycalopex fulvipes (N = 2). FMNH: 23814; 23815.

Lycalopex griseus (N = 127). AMNH: 17440a; 17440b; 17441a. CFA: 2175; 4197; 5291; 5649; 5650; 5777; 5782; 10243. CML: 837; 838; 1177; 1178; 1427; 1489; 3714; 4967; 6189; 6190; 6192. FMNH: 154639; 154640. LIEB: 794; 809. MACN: 4.253; 23.20; 24.50; 24.52; 24.53; 24.54; 24.56; 24.57; 24.59; 24.62; 24.63; 24.64; 24.66; 24.68; 24.69; 24.71; 24.74; 24.75; 24.76; 24.79; 24.80; 24.81; 223; 225; 226; 13781; 14540; 14902; 15020; 15185; 15186; 15187; 15189; 15262; 15263; 15264; 15265; 15269; 16321; 16322; 16325; 20205; 20206; 20207; 20208; 20276; 20277; 20278; 20814; 20829; 23150; 23468; 23662; 23,663; 2664; 23668; 23718; 23728; 23729; 23730; 23910; 24206; 29.895; 50.419; 50.420; 50.432; 50.490; 51.170. MLP: 5.III.36.12; 5.III.36.27; 2.IV.60.1; 4.VIII.98.4; 240; 441; 559; 696; 701; 712. NMNH: 92139; 92140; 92141; 92142; 92143; 92144; 92145; 92146; 92147; 92149; 92150; 92151; 92152; 92169; 92173; 92174; 92175; 92176; 92177; 92178; 92179; 482163; 482164.

Lycalopex gymnocercus (N = 355). AMNH: 41502; 41503; 41504; 41505; 41506; 41507; 41508; 41509; 41510. CFA: 3255; 3698; 3962; 4256; 4406; 4416; 4417; 4659; 8312; 8313; 8588; 8589; 8590; 8591; 10887; 11062; 11063. CML: 192; 495; 545; 645; 834; 836; 895; 908; 909; 959; 1179; 1526; 1526; 3072; 4081; 4082; 5143; 5473; 5474; 5479; 5480; 5772; 6342. GECM: 24; 34; 40; 51; 57; 65; 67; 75; 76; 85; 100; 108; 112; 119; 121; 129; 139; 149; 152; 153; 179; 217Bis; 220Bis; 227Bis. MACN: 4.271; 20.33; 26.28; 20.35; 23.33; 23.34; 23.36; 23.37; 23.38; 24.48; 24.49; 24.133; 24.134; 24.140; 24.141; 24.142; 24.143; 24.144; 24.145; 24.146; 24.147; 24.148; 24.149; 24.151; 24.152; 24.154; 24.156; 24.162; 24.169; 24.170; 26.129; 26.162; 26.163; 27.53; 28.182; 29.35; 30.150; 30.210; 30.211; 30.212; 32.252; 32.263; 33.177; 33.266; 33.268; 34.317; 35.241; 36.178; 36.479; 36.480; 37.82; 38.243; 39.191; 39.194; 41.220; 41.221; 44.17; 48.266; 49.134; 49.139; 49.148; 49.149; 49.159; 49.160; 49.167; 50.56; 50.443; 50.491; 50.492; 50.494; 50.495; 50.497; 50.498; 50.500; 50.501; 50.502; 50.503; 50.504; 50.505; 51.81; 53.2; 54.133; 246; 285; 293; 13299; 13313; 13327; 13331; 13337; 14319; 14323; 14386; 14409; 15363; 15364; 15387; 15388; 15389; 15390; 15601; 15692; 15742; 15748; 15749; 15750; 15751; 15752; 15754; 15757; 15758; 15760; 15761; 15762; 15764; 15765; 1766; 157769; 15771; 15783; 15784; 15785; 15787; 15788; 15791; 15792; 15794; 15795; 15796; 15797; 15800; 15818; 15820; 15831; 15833; 15834; 15838; 15854; 15859; 15862; 15863; 15864; 15865; 15866; 15867; 15868; 15869; 15870; 15871; 15873; 15875; 15879; 15882; 15888; 15892; 15894; 15895; 15896; 15898; 15901; 15902; 15906; 15908; 15909; 15917; 15932; 15933; 15934; 15938; 15941; 15958; 15963; 15964; 15966; 15970; 15973; 15979; 15981; 15982; 15986; 15987; 15992; 15998; 15999; 16000; 16001; 16006; 16009; 16010; 16013; 16014; 16015; 16024; 16025; 16026; 16027; 16030; 16031; 16032; 16035; 16036; 16037; 16038; 16039; 16040; 16041; 16046; 16047; 16048; 16049; 16050; 16055; 16059; 16062; 16063; 16066; 16068; 16074; 16077; 16079; 16080; 16083; 16085; 16088; 16094; 16096; 16097; 16099; 16100; 16101; 16102; 16103; 16104; 16105; 16106; 16107; 16108; 16110; 16111; 16115; 16117; 16118; 16120; 16122; 16123; 16130; 16131; 16139; 16143; 16145; 16149; 16151; 22936; 23153; 23154; 23155; 23156; 23157; 23158; 23290; 23920; 24203; 24204; 24205; 24208; 24209; 24259; 24265; 24282. MLP: 16.III.99.16; 13.IV.99.3; 13.IV.99.13; 13.IV.99.14; 13.IV.99.36; 26.V.95.9; 4.VIII.98.9; 30.XII.02.65; 710. NMNH: 172789; 172790; 236366; 331065.

Lycalopex sechurae (N = 35). AMNH: 100091; 100100; 133926; 133927; 133928; 133929; 133937; 2091; 349; 36457; 391; 46525; 46526; 46527; 46528; 46529; 46530; 46531; 46532; 46533; 63709; 70091. FMNH: 19971; 19972; 20747; 53911; 80953; 80954; 80955; 80956; 80957; 80958; 80959; 80960; 80961; 80962; 80963; 80964; 80965; 80966; 80967; 80968; 80969.

Lycalopex vetulus (N = 31). MLP: 1258. MZUSP: 1011; 1012; 1014; 1015; 1016; 1018; 1075; 1076; 1084; 12040; 13611; 3046; 3047; 3047; 3048; 3049; 3050; 825. NMNH 121171; 121172; 181150; 545109.

Speothos venaticus (N = 34). AMNH: 136285; 167846; 175306; 184688; 37472; 76035; 76805; 76806; 98558; 98559; 98560; 98640. FMNH: 121544; 125402; 60290; 87861. MACN: 50.67; 16510. MZUSP:19743; 19744. NMNH: 253504; 270165; 270171; 270368; 270369; 270370; 307650; 314048; 395841; 398030; 521045; 538307; 544414; 582465.

Urocyon cinereoargenteus (N = 37). AMNH: 255645; 255648; 254470; 8197; 243449; 100301; 243095; 120989; 184105; 184122; 184012; 183939; 184002; 184065; 183979; 184098; 184094; 184077; 184091; 184126; 184083; 184121; 184014; 184087; 184013; 184009; 183991; 183956; 183942; 183943; 183960; 183953; 183995; 183954; 185512; 184064; 184007.

Vulpes lagopus (N = 1). MACN: 4.1.

Vulpes vulpes (N = 54). FMNH: 106726; 107271; 140172; 140176; 74472; 74987; 74988; 74989; 75644; 75645; 75646; 77130; 77136; 78650; 78651; 80827; 80829; 80836; 80837; 80839; 80840; 84697; 84698; 85216; 85217; 85218; 86820; 89369; 89370; 89371; 89372; 89587; 89710; 89712; 89963; 90361; 90473; 90474; 91605; 91725; 91726; 91731; 91741; 92727; 95863; 95865; 95867; 95870; 95872; 95873; 98733; 98734; 98735; 98736.

Vulpes zerda (N = 2). CML: 3731. MACN: 3.14.

Theriodictis platensis (N = 1). MLP-PV: 96-IX-1-1.

Otocyon megalotis (N = 1). MACN: 26.115.

Lycaon pictus (N = 1). MACN: 38.249.

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Segura, V., Cassini, G.H., Prevosti, F.J. et al. Integration or Modularity in the Mandible of Canids (Carnivora: Canidae): a Geometric Morphometric Approach. J Mammal Evol 28, 145–157 (2021). https://doi.org/10.1007/s10914-020-09502-z

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