J Mammal Evol (2019) 26:237–251 DOI 10.1007/s10914-017-9391-z ORIGINAL PAPER Cryptoprocta spelea (Carnivora: Eupleridae): What Did It Eat and How Do We Know? Lindsay Renee Meador 1 & Laurie Rohde Godfrey1 & Jean Claude Rakotondramavo2 & Lovasoa Ranivoharimanana2 & Andrew Zamora1 & Michael Reed Sutherland3 & Mitchell T. Irwin4 Published online: 27 May 2017 # Springer Science+Business Media New York 2017 Abstract The extent to which Madagascar’s Holocene ex- carnivory by mammals, raptors, and crocodiles at different tinct lemurs fell victim to nonhuman predators is poorly un- sites and ecoregions. Our data reveal that crocodiles, raptors, derstood. Madagascar’s Holocene predator guild included and the largest of Madagascar’s mammalian predators, several now-extinct species, i.e., crocodiles, carnivorans, and C. spelea, all preyed on large lemurs. Cryptoprocta opportu- raptors. Here we focus on mammalian carnivory, specifically nistically consumed lemurs weighing up to ~85 kg. Its fore- the roles of Cryptoprocta spelea and its still-extant but limb anatomy would have facilitated predation on large- smaller-bodied sister taxon, C. ferox, the fosa. Cryptoprocta bodied prey. Social hunting may have also enhanced the abil- spelea was the largest carnivoran on Madagascar during the ity of C. spelea to capture large, arboreal primates. Quaternary. We ask whether some extinct lemurs exceeded the Cryptoprocta carnivory is well represented at cave and river- upper prey-size limits of C. spelea. We use univariate and ine sites and less prevalent at lake and marsh sites, where multivariate phylogenetic generalized least squares regression crocodylian predation dominates. models to re-evaluate the likely body mass of C. spelea. Next, we compare characteristics of the forelimb bones of C. ferox Keywords Fosa . Cryptoprocta . Extinct lemurs . and C. spelea to those of other stealth predators specializing Madagascar . Extinction . Eupleridae . Predator on small, mixed, and large-bodied prey. Finally, we examine humeri, femora, crania, and mandibles of extinct lemurs from six sites in four ecoregions of Madagascar to identify damage Introduction likely made by predators. We test the relative prevalence of Cryptoprocta spelea was the largest Holocene carnivoran on the island of Madagascar. It became extinct sometime during Electronic supplementary material The online version of this article (doi:10.1007/s10914-017-9391-z) contains supplementary material, the past 2000 years, leaving its sister taxon, C. ferox,asthe which is available to authorized users. largest of the remaining endemic Madagascan carnivorans. Male and female C. ferox overlap in body mass but show * Lindsay Renee Meador some sexual dimorphism, and there is considerable interpop- [email protected] ulation variation, so that adult body mass of C. ferox ranges from 5.5 to 9.9 kg (Goodman 2012). Past reconstructions of 1 Department of Anthropology, University of Massachusetts Amherst, the mean body mass of C. spelea have differed considerably, 240 Hicks Way, Amherst, MA 01003, USA from not much more than 10 kg (Goodman and Jungers 2014) 2 Mention Bassins sédimentaires, Evolution Conservation (BEC), BP to around 20 kg (based on regressions published by Van 906, Faculté des Sciences, Université d’Antananarivo, Valkenburgh 1990), the latter being over twice the body size 101 Antananarivo, Antananarivo, Madagascar of living C. ferox. Cryptoprocta ferox regularly kill prey that 3 Data Science Program, New College of Florida, Sarasota, FL 34243, match or exceed their body size (Goodman et al. 1997, 2004; USA Britt et al. 2001; Dollar et al. 2007), and it is likely that 4 Department of Anthropology, Northern Illinois University, Grant C. spelea would have done the same, but whether it also killed Tower South A – 507, DeKalb, IL 60115, USA much larger now-extinct lemurs has remained uncertain 238 J Mammal Evol (2019) 26:237–251 (Goodman and Jungers 2014). The largest of the extinct le- we verified the species for each of the previously-dated long murs may have weighed over 150 kg, and several others bones of Cryptoprocta. This information allowed us to evalu- weighed around 50 kg or more. Other formidable megafaunal atesympatryintimeaswellasspace. predators, including a now-extinct crocodile (Voay robustus) Long bone measurements were used to reconstruct the (Grandidier and Vaillant 1872;Brochu2007), may have com- body mass of subfossil Cryptoprocta and indices were calcu- peted with C. spelea, perhaps more successfully killing larger lated to assess forelimb function, following Meachen-Samuels lemurs. Living crocodiles routinely attack and kill large mam- and Van Valkenburgh (2009) (Table 1). Meachen-Samuels mals (Baquedano et al. 2012). Goodman and coworkers and Van Valkenburgh (2009) devised forelimb skeletal indices (Goodman 1994a, b; Goodman and Rakotozafy 1995; to distinguish felids concentrating on small prey from those Goodman and Jungers 2014; Goodman and Muldoon 2016) targeting prey of mixed sizes and those specialized to bring have also identified three species of extinct raptors (two down large prey. Following Carbone et al. (2007), we classi- Aquila,orBtrue^ eagle species, and a crowned eagle, fied Bsmall-prey specialists^ as species targeting prey smaller Stephanoaetus mahery) that would have been capable of prey- than themselves, Blarge-prey specialists^ as those targeting ing on the now-extinct lemurs. prey larger than themselves, and Bmixed-prey specialists^ as We reconstruct the role of Cryptoprocta as a possible pred- opportunists that regularly target either. Despite being distant- ator of large-bodied lemurs by combining an analysis of the ly related to cryptoprocts (family Eupleridae), felids make an size and morphology of its forelimb bones with an analysis of excellent reference population for cryptoprocts because, like predator traces on the bones of extinct lemurs. First, we estab- lish the contemporaneity of subfossil Cryptoprocta and ex- Table 1 Postcranial measurements taken and indices calculated tinct lemurs by examining the geographic distribution of Cryptoprocta and its radiocarbon records. We then ask wheth- Measurement (mm) or Index er the forelimb structure of C. spelea conforms to expectations Humeral length for small-, mixed-, or large-prey hunting by predators that Humeral midshaft circumference depend on stealth ambush methods. We use univariate and Humeral midshaft transverse diameter multivariate phylogenetic generalized least squares (PGLS) Humeral midshaft anteroposterior diameter models to reconstruct the body size of C. spelea, and test the Femoral length notion that Cryptoprocta targeted only prey animals at the low Femoral midshaft circumference end of the megafaunal range in body size by examining the Femoral midshaft transverse diameter predation traces on the bones of the extinct lemurs themselves. Femoral midshaft anteroposterior diameter We test the hypothesis that Cryptoprocta was an opportunistic Radial length hunter by examining the correspondence between the size of Ulnar length (olecranon tip to distal styloid) its selected prey and the size of individuals that we take to Humeral biepicondylar breadth represent the populations of available prey animals at partic- Humeral distal articular breadth ular sites – i.e., those that do not show predation traces. Length of the ulnar olecranon process Finally, we determine the relative prevalence of carnivoran predation on animals of different body sizes, in different Radial midshaft diameter ecoregions, and at sites of different types (marsh or lake, cave, Mediolateral diameter of distal radial articular facet and flood plain). Anteroposterior diameter of distal radial articular facet Brachial Index (BI): radial length / humeral length Humeral Robustness Index (HRI): humeral midshaft transverse diameter / humeral length Materials and Methods Humeral Epicondylar Index (HEI): humeral biepicondylar breadth / humeral length JCR collected metric data on 75 postcranial bones of subfossil Humeral Condylar Index (HCI): humeral distal articular breadth / C. spelea and C. ferox (including 17 humeri, 25 femora, 17 humeral length radii, and 16 ulnae; see Online Resource 1), as well as on Olecranon Index (OI): length of olecranon process / (ulnar length – length miscellaneous fragmentary skulls, a complete skull of a of olecranon process) C. spelea from Bevoha (uncatalogued) and a complete skull Radial Robustness Index (RRI): radial midshaft diameter / radial length of a modern C. ferox, AM 240 (AM = Académie Malgache). Radial Articular Index (RAI): mediolateral diameter distal radial articular Both of the complete skulls were previously illustrated facet / radial length (Lamberton 1939). Radiocarbon dates for subfossil Radial Distal Articular Area Index (RAA): (mediolateral diameter of the distal radial articular facet x anteroposterior diameter of the distal radial Cryptoprocta have been published, but, as pointed out by .5 articular facet) / radial length Goodman and Jungers (2014), the species identifications of radiocarbon-dated individuals have been uncertain. Therefore, Indices follow Meachen-Samuels and Van Valkenburgh (2009) J Mammal Evol (2019) 26:237–251 239 Cryptoprocta, they are generally hypercarnivorous ambush transverse humeral diameter, and midshaft humeral cir- hunters, and many are arboreal or semi-arboreal. Genetic re- cumference) that showed average R2 values above 0.85 search confirms that Cryptoprocta belongs to the Feliformia and regressed them against body mass in a