Adaptations of the Pleistocene Island Canid Cynot He Rium Sardous

CRA NIUM 23, 1 - 2006

Adaptations of the Pleistocene island canid Cynothe rium sardous (Sardinia, Italy) for hunting small prey

George Lyras and Alexandra van der Geer

Summary Cynothe rium sardous is a small canid that lived on the island of Sardinia-Corsica during the Pleistocene. Once on the island, the species gradually adapted, and became specialized in hunting small prey like the lagomorph Prolagus. Moreover, in order to fulfil mass-related energetic requi rements, the species had to reduce body size compared to its ancestor Xenocyon, which was larger than the grey wolf. Cynotherium carried its head much in the way foxes do, and was able to hold its body low to the ground when stalking. In addition, it could move its head laterally better than any living canid.

Samenvat ting Cynothe rium sardous is een kleine hond achtige, die leefde op het eiland Sardinië-Corsica gedu rende het Pleistoceen. Eenmaal op het eiland paste de soort zich aan en specialiseerde zich in het jagen op kleine prooi zoals de haasachtige Prolagus. Om aan de energiebehoeften, gere lateerd aan lichaamsgewicht, te voldoen, moest de soort kleiner worden, vergeleken met zijn voorouder, Xeno cyon, die groter was dan de huidige grijze wolf. Cynothe rium hield zijn hoofd ongeveer zoals vossen doen, en hield het lichaam laag bij de grond bij het besluipen van de prooi. Daarbij kon hij zijn kop verder zijwaarts bewegen dan alle nu levende hondachtigen. Intro duc ti on When we hear about insular island mammals, we immedi a tely make associ a tions with pig-sized hippo's, mini-mammoths, giant rodents, deer adapted for mountain clim bing, apart from the scientific names of several Plio-Pleistocene insular ungulates and micro - mammals. We all think of the well-established model accor ding to which mammals reached the islands by swimming or raf ting. Once there, due to limited resources and the absence of terrestrial mammalian predators, their descen- Fig 1 The studied ske leton of Cy not he rium sardous dants gradually became adapted to the island (missing parts in whi te). The background outline re - environment by a change in body plan in presents the size of its an cestor Xe no cy on ly ca o noi des several ways, e.g. by redu cing or increasing (based on data from Sotnikova, 2001). their size, fusion of limb bones, etcetera. Het be stu deer de ske let van Cy not he ri um sar dous That is in general true, but not in all cases. (ont bre ken de de len in wit). Het ach ter grond pro fiel There are, for example, cases of mammalian stelt de grootte van zijn voorouder, Xe no cy on ly ca o - predators that lived on islands. Such cases in noides , voor (gebaseerd op data uit Sotnikova, 2001). the first instance disturb the nice picture. However, their study provides us with a better The most promising place to study an insular and more complete picture of island mammal mammalian predator is provided by the islands evolution, not only because more and different of Sardinia (Italy) and Corsica (France). From taxa are involved, but also because of their rela- the Late Miocene to the Late Pleistocene, these tion with the other endemic animals on the two islands, which were for most of their island. It appears from such studies that also geological history connected with each other, mammalian predators follow the earlier obser- were repeatedly colonized by mammals vations, and thus that also they show adapta- (Sondaar & Van der Geer, 2005), and as a conse- tions to the pecu liar island environment. quence, the fossil record of Sardinia-Corsica is

51 Adaptations of the Pleistocene island canid Cynothe rium sardous (Sardinia, Italy) for hunting small prey characterized by more faunal changes and a isolated, it became a small and speci alized greater diversity than any other island with small-prey hunter (Lyras et al., in press). insular mammals (table 1). Of particular interest of all the Sardinian-Corsican extinct mammals, are the carnivores. Fossils of several carni vore Ma te ri als and Me thods groups have been recovered from Sardinia-Corsica: the hunting hyaena Here we present the results of the morphofunc - Chasmaporthetes melei, the canid Cynotherium tional analysis of a Cynothe rium skeleton that sardous, and several otters (Algarolutra majori, was excavated in Cobeddu Cave (Lanaittu Sardolutra ichnusae, Enhydrictis galictoides, Valley, Nuoro, Sardinia, Italy; see also Van der Megalenhydris barbaricina). Geer & Van der Geer, 2001). The skeleton is fairly complete (fig. 1) and in an excellent state Cynot he rium sardous fossils are found in relati- of preservation. The fact that the mate rial vely large numbers, inclu ding not only skulls belongs to one indi vidual, provides an excellent and dentition, but also postcranials. This opportu nity for morpho func ti onal analysis. For species can therefore be studied in more detail. the description of the skeleton we follow the Cynothe rium is a small canid with dental charac- nomenclature of Evans (1993). Data on the ters that are typical for hyper car ni vorous canids myology of the living canid species have been (i.e., canids that include signi fi cant quanti ties of taken from Langguth (1969) and Evans (1993). meat in their diet and prey mostly on large To check these data and to see the nature of the animals). Its cranial characters, however, indi- different muscles and tendons, we dissected a cate that it was not able to hunt large prey. A red fox (Vulpes vulpes) and a domestic dog recent phylogenetic analysis by Lyras et al. (in (Canis fami li aris). press), led to the conclusion that Cynothe rium originates from Xenocyon lycao noides , a In order to investigate the functional signifi - large-sized dog, related either to some extinct cance of the observed features we followed the wolves (Wang et al., 2005) or to the Cape form-function correlation method of analysis. hunting dog, Lycaon pictus (Martinez-Navarro & This method extrapolates the correlation that is Rook, 2003). Xenocyon reached Sardinia-Corsica observed between anatomical forms and certain at the end of the early Pleistocene and once functions or behaviours of living animals to the

Fig 2 Greatest width of the distal end of the humerus of Xe no cy on ly ca o noi des from Stránska Skála, Czech Repu - blik, and Undermassfeld, Germany (data from Sotnikova, 2001), com pa red with Cynot he ri um spp. from Sardinia: C. sp. NMB Ty-5362 from Capo Figari (data and photograph from van der Made, 1999), C. sardous from Dra go na- ra Cave (data from Malatesta, 1970) and from Corbeddu Cave (CB 84-8022).

Groot ste breed te van het on de rui tein de van het op perarm been van Xe no cy on ly ca o noi des van Stránska Skála, Tsjechië, en Undermassfeld, Duitsland (data uit Sotnikova, 2001), vergeleken met Cynot he ri um spp. van Sardinië: C. sp. NMB Ty-5362 van Capo Figari (data en foto uit Van der Made, 1999) en C. sardous van de grot Dragonara (data uit Malatesta, 1970) en de grot Corbeddu (CB 84-8022).

52 CRA NIUM 23, 1 - 2006 fossil species (Radinsky, 1987). In order to do et al., 2005). A stratig rap hi cally younger locality so, we compared the morphology of the is Dragonara Cave, Sassari, from which material Corbeddu skeleton with specimens from most belonging to several Cynothe rium , indivi duals living canid species (see table 2). has been excavated (Malatesta, 1970). The youngest material comes from Corbeddu Cave Material of the living and of the fossil species near Oliena, Nuoro (hall II and hall IV, see Van was studied at and belongs to the following der Geer & Van der Geer, 2001). The skeleton institutes: AMNH(M): American Museum of from hall II, under study here, is the youngest Natural History, Deparment of Mammamology Cynothe rium specimen (Klein Hofmeijer, 1996) New York, USA; CB: Corbeddu Cave, Sardinia, known till now. Its postcranial is of a smaller Italy; FMNH: Field Museum of Natural History, size than any specimen from Dragonara (fig. 2). Chicago, USA; MNHN: Muséum Nati onal The latter specimens are in turn smaller than d’Histoire Naturelle, Paris, France; NMB: the humerus from Capo Figari. Natur his to ri sches Museum Basel, Swit zer land; NNML: Nati o naal Natuur his to risch Museum Although the data are limited, it is tempting to (Naturalis), Leiden, the Netherlands. suggest that they provide us with an indication of size reduc tion of Cynothe rium through time. The phenomenon of size reduction becomes Size re duc ti on even more evident when we compare Cynothe - rium with its probable mainland ancestor. As Findings of Cynothe rium in Corsica are limited shown in figure 2, the humeri of Xenocyon (data (Robert & Vigne, 2001), but Sardinia has many from Sotnikova, 2001) are considerably larger localities with fossil remains of that canid. The than those of Cynothe rium . oldest remains found till now come from breccia deposits on Capo Figari, Sassari (Van In theory, body mass can be estimated with the der Made, 1999) and from fissure fillings at use of a number of dental and skeletal measure- Monte Tuttavista near Orosei, Nuoro (Abbazzi ments (Damuth & MacFadden, 1990). In dwarf forms, however, dental measurements are not reliable in such estimations, as their rela tion with body proportions differs essentially from those of their mainland relatives (e.g. Sondaar, 1977; Lister & Davies, 2003). Thus, the body mass of Cynotherium can only be estimated based on postcranial elements. The method based on the circumference of the femur at the midshaft, developed by Anyonge (1993), gives the best estimation of body mass in living canids. Applying this method results in a body mass for Cynotherium from Corbedu Cave of approximately 10 Kg, which is the size of an average jackal. Reduction in body size is a common phenomenon in island ungulates, combined, amongst others, with changed interlimb ratios and the tendency of distal elements to fuse. This phenomenon coincides with a simultaneous Fig 3 Comparison of the skulls of Xeno cy on lyca o noi - absence of large mammalian predators and des (a, b and c) from Venta Micena (Spain) and Cy- nothe ri um sardous (d, e and f) from Corbeddu cave. limited food resources (Sondaar, 1977). The drawings of Xeno cy on are ba sed on the illustrati - Cynotherium, on the other hand, is not an ungu- ons by Palmqvist et al. (1999). late but a predator, and thus may have followed a different evolutionary path on the island. Ver ge lij king van de sche dels van Xe no cy on ly ca o noi - What seems to have followed the ungulate des (a, b en c) van Venta Mi cena (Spanje) en Cynot he - pattern is its size reduction. It is known that the rium sardous (d, e en f) uit de grot Corbeddu, size of a carnivore is closely related to that of its Sar di nië. De te ke ning en van Xe no cy on zijn ge baseerd prey (Carbone et al., 1999). According to them, op de illustraties van Palmqvist et al. (1999). in order to fulfil mass-related energetic require -

53 Adaptations of the Pleistocene island canid Cynothe rium sardous (Sardinia, Italy) for hunting small prey ments, large-sized carnivores prey on large also continues with an aponeurosis on the animals (near predator’s mass) while smaller dorsal nuchal line; its other insertion is situ ated carnivores feed on small prey (less than half of on the spinous processes of the first thoracic predator’s mass). As we noted in Lyras et al. (in vertebra. Its function is raising the head and press), the cranial characters of Cynotherium entire neck. (weak zygomatic arch, low sagittal crest, thin The ventral surface of the basioccipital of mandible; fig. 3) suggest that it was not able to Cynothe rium is broader than in similar sized hunt big game. From the diagrams of Carbone living canids and the two rugose fossae for neck et al. (1999), it appears that carnivores smaller muscle attachment are deep and wide (Lyras et than 21 kg hunt prey smaller than half their al., in press). These fossae are the scars for the own size. The body mass of 10 kg for Mm rectus capitis ventralis, rectus capitis lateralis Cynotherium falls well within the range of and longus capitis. Their prin cipal action is Carbone et al.'s (1999) small prey hunters. flexion of the atlanto-occipital joint; the latter also draws the neck downwards. Lo co mo tor adap ta ti ons The overall picture indi cates that Cynothe rium had stronger neck movements than any of the Most morp hological features that can be living canids. This includes rota tion, extension observed in Cynothe rium can be found also in and flexion of the atlanto-occipital joint and one or more living canids; in other words, they raising and lowering of the head. fall within the morp hospace of the living canids. As there is no significant substantial In addition to the stronger neck musculature, diversity in loco motor patterns present in the the neck as a whole was held lower than in living canids (Van Valkenburgh, 1985; Canis, as already noted and figured by Mala- Andersson, 2003), the pattern in Cynothe rium testa (1970). Furthermore, it appears that C3 could not have been much different from those already bears a prominent spinous process, as seen in the living canids. There are, however, in Vulpes, whereas this is in general the case for some unique morphological features in the neck C4 in Canis and for C5 in Cuon. and anterior limb of Cynothe rium that are not Anteri or limb seen in other canids. These features are the only ones that, in our view, could shed some light on The very tip of the atlas is sharp and prominent particular adapta tions of Cynothe rium. in Cynothe rium (fig. 4). It is the insertion point for the tendon of the M. omotrans vers arius. This Neck mobi li ty and posi ti on muscle draws the limb forward; it is only a flat Apart from the unusual combination of cranial narrow muscle in Canis and Vulpes, but it and dental characters, the skull of Cynothe rium appeared to have been stronger and more has another pecu liarity: its mastoid process is important in Cynothe rium . The other insertion much enlarged and is situ ated laterally, being point of this muscles lies on the scapular spine more or less a continuation of the nuchal crest just proximal to the acro mion (figs. 4 and 5). (Lyras et al., in press). The mastoid process is This area is enlarged in Cynothe rium , though the place of insertion for several muscles. Most this may equally be due to a larger insertion of the surface of the process is occu pied by the area for the M. deltoideus (see below). insertion of the M. obliquus capitis cranialis, and The large, high and rugose scar for the M. deltoi- lateral to this area are the insertions for M. sple- deus at the caudal surface of the posterior end of nius, and two other neck muscles. the deltoid ridge of the humerus indicates that The M. obliquus capitis cranialis conti nues its this muscle was important; the same can be insertion further upward on the dorsal nuchal inferred from its insertion on the scapula. Its line. The function of this muscle is to extend the action is lifting of the humerus and flexion of atlanto-occipital joint, while its caudal coun ter- the shoulder (fig. 5). Another shoulder flexor is part rotates the atlas, and thus the head on the the M. teres minor. The scar for this muscle axis. The anterior parts of the atlas wings are insertion is well-developed in Cynothe rium . reduced (fig. 4), therefore, the fibres of the The well-developed attachment areas for the complemen tary muscle, the M. obliquus capitis triceps on the scapula, humerus and ulna indi- caudalis, were oriented more ventrolaterally in cate that this muscle was important in Cynothe - Cynothe rium , which is suggestive of a stronger rium, more so than in any other canid (fig. 5). Its rotati onal component. The M. splenius insertion

54 CRA NIUM 23, 1 - 2006 function was assisted by an equally well-devel- living canids. If we take into account that the oped M. anconeus, which is also an extensor of elbow joint of Cynothe rium has a higher the elbow. In view of the pronounced tubercles, maximal flexion than observed in Canis, Cuon the two main extensors of the elbow are and Vulpes (Lyras et al., in press), it is clear that stronger in Cynothe rium than in Canis and Cuon. the elbow extensors needed to work more for the same end result. To sum up, flexion of the shoulder and extension of the elbow was very powerful in Cynothe - The powerful shoulder flexion and elbow exten- rium. The forward motion of the anterior limb sion, together with the relatively flexed elbow, may also have been more developed than in

Fig 4 Postcra ni al elements of Cynot he rium , Ly ca on and Canis .a:Cy not he ri um sardous CB 84-8022 humerus, (a1) caudal, (a2) dorsal view; Lyca on pictus AMNH(M) 81853 humerus, (b1) dorsal and (b2) caudal view; Cynot he ri um sardous CB 84-8022 atlas, (c1) dorsal and (c2) ventral view; d: Lyca on pictus AMNH(M) 81853 atlas ventral view; e: Canis simen sis AMNH(M) 81081 atlas dorsal view; f: Lyca on pictus AMNH(M) 85154 scapu la late ral view; g: Cy- nothe ri um sardous CB 84-8022 scapula lateral view.

Post cra ni a le ele men ten van Cynot he rium , Ly ca on en Canis .a:Cy not he ri um sardous CB 84-8022 opperarmbeen, (a1) ach ter aan zicht, (a2) bo ven aan zicht; Lyca on pictus AMNH(M) 81853 op per arm been, (b1) bo ven aan zicht and (b2) ach ter aan zicht; Cynot he ri um sardous CB 84-8022 atlas, (c1) boven aan z icht en (c2) onder aan zicht; d: Lyca on pictus AMNH(M) 81853 atlas onder aan zicht; e: Canis simen sis AMNH(M) 81081 atlas bovenaanzicht; f: Lyca on pictus AMNH(M) 85154 schou der blad, zij-aan zicht; g: Cynot he ri um sardous CB 84-8022 schouderblad, zij-aanzicht.

55 Adaptations of the Pleistocene island canid Cynothe rium sardous (Sardinia, Italy) for hunting small prey

Fig 5 a: Di rections of pull of the mus cles of the neck (so lid line) and an terior limb (dotted lines) of Cynot he ri um sardous. 1: M. sterno cep ha li cus ; 2: M. obli quus ca pi tis; 3: M. omotrans vers ari us ; 4: M. triceps caput late ra le ; 5: M. tri ceps ca - put logum; 6: M. te res minor ; 7: M. del toi deus. b: Dorsal view of the el bow of a dis sected fox. 1: M. ancon eus ; 2: M. triceps caput medi a le . a: rich tingen van trekkracht van spieren van de nek (doorgetrokken lijn) en van de voorpoot (gestippelde lijnen) van Cynot he ri um sardous . b: de el leboog van een ontlede vos gezien van boven. Voor de spiernamen, zie overeenkomstige nummers in het Engelstalig on derschrift. points to stalking behavior with a lowered body Inter pre ta ti on and a lowered neck and head. The increased extension power of the elbow joint and flexion of the humero-scapular joint of Cynothe rium fits a stalker. When we take the lowered neck and the more flexed elbow into account, the picture arises of a canid which with

56 CRA NIUM 23, 1 - 2006

Fig 6 La ter al view of the Si me ni an wolf, Canis simen sis , eng aged in stal king lo co mo ti on whi le appro aching its prey. Drawn from a frag ment of a BBC film.

Zij-aan zicht van de Ethi o pi sche wolf, Canis simen sis , bezig met het besluipen van een prooi. Nagetekend naar een fragment uit een BBC film. its body and neck held low to the ground animals and the same fast lateral head move- followed his prey. The strong flexors of the ments as living canids do, but to a much higher shoulder point to powerful, sudden forward degree. Cynot he rium sardous was a small-sized thrusts to grab the prey. The increased power of highly specialized hunter of small, swift prey the neck musculature indicates fast or strong (fig.7). movements of the neck, which is needed in cases where the prey escapes with random movements or in zigzag-patterns. The dentition Acknow ledgments of Cynothe rium , together with its cranial features, indicates specia li za tion on small prey. The authors wish to thank John de Vos (NNML), Richard Tedford (AMNH), Marina At present, canids that hunt small terrestrial Sotnikova (Geological Institute, Russian animals use stalking to approach their prey. Academy of Sciences, Moscow) and Vera Eisen- This is well-known for foxes (Ewer, 1973), but mann (MNHN) for the useful discussions we has also been reported for the Simenian wolf had with them. We also thank Chris Smeenk (Sillero-Zubiri & Gottelli, 1995, see also fig. 6) and occasionally coyotes (Fox, 1969). Most canids catch birds from time to time (Ewer, 1973), but a canid which mainly fed on birds was the Falkland wolf Dusi cyon australis (Mivart, 1890), a recently extinct island canid. This species had strongly developed mastoid processes comparable to those of Cynothe rium . As it became extinct before the turn of the 19th century (Nowak & Paradiso, 1999) unfortuna- tely almost nothing is known about its hunting behavior. Fig 7 Cynot he ri um , which origi na tes from the lar ge, wolf-like Xeno cy on , be ca me spe ci a li zed in hun ting As already noted above, none of the living small prey instead of large animals. That is the sub- canids has a truly specialized loco motion or ject of this cartoon: two Xeno cy on , whi le hun ting an neck mobility. This means that when we ungulate, realize that the island is full of small Pro la - observe that some of the anatomical features of gus and de cide to adapt. the anterior limb and neck of Cynothe rium are beyond the degree seen in the living canids, it is Cynot he ri um , een af stammeling van de gro te, likely that its behaviour was more specialized wolf-achti ge Xeno cy on , speci a li seer de zich in het ja- than that of any living canid. Based on the gen op klei ne prooidieren in plaats van gro te prooi. skeleton of Cynot he rium sardous, we conclude Dat is het onderwerp van deze cartoon: twee Xeno cy - that this species performed a similar stalking ons, ja gend op een hoefdier, re aliseren zich dat het with lowered body while hunting small, swift eiland vol is met kleine Pro la gus en besluiten zich aan te passen.

57 Adaptations of the Pleistocene island canid Cynothe rium sardous (Sardinia, Italy) for hunting small prey and Duncan Reeder (NNML), Bruce Table 1. Fauna list of Pleistocene of Sardinia. Cava 6, 7 and 10 Patterson and Bill Stanley (FMNH), Jean are fissu re depo sits at Monte Tutta vis ta near Orosei, Nuo ro Spence (AMNH), Michel Tranier and Géral- (after Sondaar, 1998; added are species names for Chas ma port- dine Pothet (MNHN) for their assistance hetes melei and Sus sondaari ). during the study of collec tions under their care. Finally, we thank Hans Brinkerink Faunalijst van het Pleistoceen van Sardinia. Cava 6, 7 en 10 (Vista Natura, Baarn, the Netherlands) for zijn spleet op vul ling en in de berg Mone Tut ta vis ta dicht bij Orosei, Nu oro (naar Son daar, 1998; toe gevoegd zijn de soort- making a cast of the Cynothe rium skeleton. na men voor Chasma port he tes melei en Sus sondaari ). This project has been co-financed within Op. Education by the ESF (European Social Fund) and National Recourses. Pleis to ce ne Ear ly Middle Late

Cava 7 Addresses of the authors Cava 6 Cava 10 and Cor - George Lyras bed du Museum of Palae ontology and Geology Cynot he ri um sp. X ? Department of Geology National and Kapodis trian University of Cynot he ri um sardous X X Athens Ma ca ca ma jo ri X X Greece e-mail [email protected] Talpa tyrrheni ca X X

Alexandra van der Geer Nesi o ti tes simi lis X X X [email protected] avander [email protected] Prola gus figa ri X Prola gus sardus X X

Le po ri dae X

Tyrrheni co la son daari X

Tyrrhenico la hense li X X

Rhagapo de mus minor X X

Rhagamys ortho don X

Chasma port he tes melei X

Muste li dae sp.1 X

Muste li dae sp.2 X X

Neso go ral melo nii X

Neso go ral sp.2 X

Sus sondaari X

“Mega lo ce ros” cazi o ti X

Ca pri nae X

58 CRA NIUM 23, 1 - 2006

Table 2. List of ex tant ca nid ske letons used in this stu dy. Note: Dusi cy on austra lis is re presented only by skulls, no post cra ni als have been preserved.

Lijst van skeletten van te genwoordige hond achtigen die gebruikt zijn in deze stu die. Opmer king: van Dusi cy on austra lis zijn al leen schedels be waard ge bleven.

Spe cies Num ber Spe cies Num ber

NNML cat b, NNML cat a, NNML NNML cat a, Alopex lago pus Cuon alpi nus cat b, NNML cat j, NNML 19797, NNML 945 NNML 3329

AMNH(M) 100095, FMNH NNML 19111, Atelo cy nus micro tus Dusi cy on austra lis 60675, FMNH NNML 19112 121286

AMNH(M) 81853, NNML cat a, AMNH(M) 164162, NNML cat b, Canis adustus Lycaon pictus NNML cat a, NNML NNML cat g, cat b, FMNH 12781, NNML cat j FMNH 12813

NNML 1814, NNML 3217, Canis aureus Nyctereutes procyonoides NNML cat a NNML 39176, FMNH 105740

NNML 1232, Canis latrans Otocyon megalotis MNHN 1960-480 NNML 3400

NNML cat a, Canis lupus Pseudalopex culpaeus NNML cat a FMNH 160107

Canis lupus hodophylax NNML 39182 Pseudalopex sechurae NNML 2381

NNML cat a, NNML NNML cat a, 3224, AMNH(M) Canis mesomelas Speothos venaticus NNML 2461 167846, FMNH 121544

MNHN c.q. MNHN 1967-474, Canis sinensis Urocyon cinereoargenteus 1880-1228, FMNH AMNH(M) 81081 129298

NNML 9881, NNML NNML cat b, Cerdocyon thous Vulpes vulpes 10934, NNML 9775, FMNH 70758 NNML 8889

Chryso cy on brachyu rus FMNH 150739 Vulpes zerda AMNH(M) 90156

Chryso cy on brachyu rus AMNH(M) 133941 Vulpes zerda MNHN 2000-666

59 Adaptations of the Pleistocene island canid Cynothe rium sardous (Sardinia, Italy) for hunting small prey

Re fe ren ces

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