Beitr. Paläont., 31:391-396, Wien 2009

The Late Miocene Mammal Faunas of the M ytilinii Basin, Samos Island, Greece: New Collection 15. Carnivore Guild Structure

by

Doris Nagel^ & George D. Koufos* 2)

N a g e l , D. & K o u f o s , G.D., 2009. The Late Miocene Mammal Faunas of the Mytilinii Basin, Samos Island, Greece: New Collection. 15. Carnivore Guild Structure. — Beitr. Palaont., 31:391-396, Wien.

Abstract Mytilinii-Becken bestätigt. Eine weitere A rt kam hinzu, Protictitherium, sowie neue Informationen zu den bereits The carnivore community of previous collections from the bekannten Formen (siehe Kapitel 5, dieser Band). Drei Upper Miocene of Samos contained 16 taxa, and most of Parameter werden für die Klassifizierung der Raubtier- them were found during recent field works in the My­ Guild verwendet - Nahrungspräferenz, Lokomotionstyp tilinii Basin, as well. The new material added one taxon, und Gewichtsklasse. Die Ergebnisse sind gemeinsam mit Protictitherium, as well as new information concerning the der Artenliste aufgeführt. Dieselbe Analyse wurde zu Ver­ already known taxa (see chapter 5, this volume). Three gleichszwecken am Raubtiermaterial von Pikermi durch­ parameters are used to classify the carnivoran guild from geführt. Beide Resultate sind in Form von 3D-Grafiken Samos - diet class, locomotion pattern and body size. The dargestellt. Evaluierungen von Paläoguilds können als results are given together with the taxon list. The same Proxies für paläoökologische Interpretationen von ehema­ analyses were done on the fossil carnivore material from ligen Habitaten verwendet werden. Dies sind die ersten Pikermi for comparison. Both results are visualised in a Guild-Untersuchungen des mediterranen Raumes. three dimensional graph. Evaluations of palaeo-guilds can be used as proxies for palaeoecological interpretations of Schlüsselworte: Oberes Miozän, Samos, Griechenland, former habitats. These are the first guild structure analyses Säugetiere, Carnivora, Paläoguilds. from the Mediterranean area.

Keywords: Late Miocene, Samos, Greece, Mammalia, 1. Introduction Carnivora, Guild Structure. The late Miocene carnivoran assemblage of Samos has been studied by several authors(P i l g r i m , 1931; S o l o u - Zusammenfassung n i a s , 1981; K o u f o s & M e l e n t i s , 1982; K o u f o s , this volume-a). The old collections from Samos, housed at Aus früheren Aufsammlungen aus dem Obermiozän von various museums include a great number of carnivore’s Samos konnten insgesamt 16 Raubtier-Taxa identifiziert taxa (S o l o u n i a s , 1981; B e r n o r et al., 1996). In our new werden. Die meisten wurden durch neuere Funde aus dem collection the carnivores are relatively few and they mainly come from the fossiliferous sites of the locality Mytilinii-1 (MTL), situated in the Adrianos ravine (K o u f o s , this Prof. Doris N a g e l , University of Vienna, Department of volume-b; K o s t o p o u l o s et al., this volume). The faunal Palaeontology, Althanstrasse 14, A-1090 Wien, Austria, e- list in the N OW database (Neogene Old World database) mail: [email protected] is based on the determination ofB e r n o r et al. (1996) and preliminary ecomorphological assignments are already 2) Prof. George D. K o u f o s , Aristotle University of Thessaloniki. indicated. Department of Geology, Laboratory of Geology and Palaeon­ The present guild analysis is based on the list of the NOW tology, GR-54124 Thessaloniki, Greece, e-mail: koufos@geo. database and on the list of the new collection. The list of auth.gr NOW is based on the old collections from Samos which come from various fossiliferous sites of different age; most Corresponding Author: Department of Palaeontology, Uni­ of them are without locality indications and the material versity of Vienna, Althanstrasse 14, A-1090 Wien, Austria, is mixed (K o u f o s , this volume-b). Thus, the authors give e-mail: [email protected] the guild structure based on the NOW faunal list (old col­ 392 Beitr. Palaont., 31, Wien, 2009

lection) and on the new collection. The carnivores of the and hypocarnivorous (V a l k e n b u r g h , 1988). The hypocar- new collection originate mainly from the locality MTL nivores are a quite diverse group since they include omni­ which is dated to middle Turolian (MN 12) and more vores as well as durophagous taxa. Although insectivorous precisely to 7.1-7.0 Ma by magnetostratigraphy(K o u f o s carnivores could be seen as hypocarnivorous,F r i s c i a et al. et ah, this volume-b). (2007) defined them by their pointed teeth and so “insec­ Several studies concerning carnivorous guilds are known tivorous” can be added to the diet class system. combining two parameters, usually body mass and either Locomotor Pattern. No new information is given for the locomotor pattern or food preferences (V a n V a l k e n - carnivores from Samos in this volume, but the taxa are b u r g h , 1992,1999; W e r d e l i n , 1996; V a n V a l k e n b u r g h all known from earlier descriptions (W e r d e l i n , 1996; et al., 2004; W e s l e y -H u n t , 2005). New approaches G i n s b u r g , 1961; G i n s b u r g , 1999) and from other sites as combine all three paramentersM o r l o , 1999; N a g e l & well. Therefore it is possible to assign them to different lo­ M o r l o , 2000, 2003; M o r l o & G u n n e l l , 2005, 2006; comotor patterns. Qualitative characters were described by M o r l o & N a g e l , 2007; N a g e l et ah, 2005; S t e f e n et ah, following authors: B a r n e t & N a p i e r (1953), G i n s b u r g 2005; G u n n e l l & M o r l o , 2006; M o r l o et al. in press). (1961), H e i n r i c h & R o s e (1997), J e n k i n s & C a m a z i n e Three parameters are used to evaluate the ecomorphologi- (1977), L a b o r d e (1987), T a y l o r (1974, 1976, 1989), cal space of the different taxa: body mass, diet type, and B e r t r a m & B i e w i e n e r (1990) and A n d e r s s o n (2004). locomotor pattern. Different postcranial features were used to define locomo­ Body Mass. Several studies have documented the quantita­ tion preference including: scapular outline, shape and size tive relationship between body mass and carnassial tooth of humeral head, size of distal humeral epicondyle, length size (T h a c k e r a y & K i e s e r , 1992; V i r a n t a & A n d r e w s , and orientation of olecranon, shape and size of humeral 1995; L e g e n d r e & R o t h , 1988; V a n V a l k e n b u r g h , and radial notches and size of distal radioulnar articular 1990). Another approach is to take limb bone measure­ process at the ulna, hand posture, shape of astragalus ments to estimate the body mass of carnivores(G i n g e r i c h , and calcaneum, and foot posture. W e separate arboreal, 1990; A n y o n g e , 1993; H e i n r i c h & B i k n e v i c i u s , 1998; scansorial, cursorial, generalized terrestrial, semifosso- C hristiansen , 1999). Since no limb bones are described rial, and semiaquatic taxa. The results are given in 3-D from the new carnivore collection from Samos (K o u f o s , visualizations of guild structure. this volume-a), we chose to use carnassial tooth size as a proxy for body mass in both the new taxa, as well as for taxa not described in this volume. In previous studies, the 2. The Carnivore Guild from Samos MN 12 use of body mass classes proved to be more reliable as an ecomorphological indicator, instead of absolute body mass The following taxa have been recognized in the new col­ data, since there are always an individual species variation lection from Samos: Parataxidea maraghana, P lioviver- as well as methodological differences. Therefore we used rops orbignyi, Hyaenictitherium wongii, Adcrocuta eximia, following body mass classes: 0 -1 kg, 1-3 kg, 3-10 kg, Metailurusparvulus and Machairodus giganteus (K o u f o s , 10-30 kg, 30 -100 kg, >100 kg (M o r l o , 1999). this volume-a). Additionally S o l o u n i a s (1981), B e r n o r Diet. In general, four diet classes can be distinguished in et al. (1996) and the N OW database listed: Ursavus cf. carnivorans: bone/meat, hypercarnivorous, carnivorous depereti, Indarctos atticus, Promelespalaeattica, Promephitis

Family Genus Species Body Mass Locomotor Class Diet Class Ursidae Ursavus cf. depereti 30-100 kg ambulatorial terrestrial hypocarnivorous Ursidae Indarctos atticus >100kg generalized terrestrial carnivorous Mustelidae Promeles palaeattica 3 - 10 kg ambulatorial terrestrial carnivorous Mustelidae Promephitis larteti 3 - 10 kg unknown hypocarnivorous Mustelidae Parataxidea maraghana 3 - 10 kg unknown hypocarnivorous Hyaenidae Plioviverrops orbignyi 3 - 10 kg generalized terrestrial insectivorous Hyaenidae Protictitheriu m crassum 3 - 10 kg scansorial insectivorous Hyaenidae Ictitherium viverrinum 10-30 kg cursorial carnivorous 9 Hyaenidae Hyaenictitherium wongi 30-100 kg generalized terrestrial carnivorous 10 Hyaenidae Lycyaena chaeretis 30-100 kg cursorial meat/bone 11 Hyaenidae Belbus beaumonti 30-100 kg cursorial carnivorous 12 Hyaenidae Adcrocuta eximia 30-100 kg generalized terrestrial meat/bone 13 Felidae Felis attica 3 - 10 kg scansorial hypercarnivorous 14 Felidae Metailurus major 30-100 kg scansorial hyp ere ar nivorous 15 Felidae Metailurus parvulus 30-100 kg scansorial hypercarnivorous 16 Felidae Machairodus giganteus > 100 kg generalized terrestrial hypercarnivorous

Table 1: Assignment of the carnivorous taxa from Samos, Late Miocene (MN 12) to body mass, diet and locomotion class. Nagel, D. & Koufos, G.D., Carnivore Guild Structure. 393

Samos, Greece, MN 12

bone/meat bone/meat hypercarnivorous hypercarnivorous carnivorous carnivorous unknown unknown hypocarnivorous hypocarnivorous semifossorial semifossorial insectivorous insectivorous cursorial cursorial

ambulatorical terrestrial ambulatorical terrestrial

generalized terrestrial generalized terrestrial

scansorial scansorial

aboreal aboreal

Figure 1: 1 - U rsavus cf. deperetr, 2 - Indarctos a tticu s; 3 - Promeles palaeattica\ 4 - Promephitis larteti', 5 - Parataxidea maraghana\ 6 - Plioviverrops orbignyv, 7 - Protictitherium crassum\ 8 - Ictitherium viverrinunv, 9 - Hyaenictitherium wongi; 10 - Lycyaena chaeretis\ 11 - Belbus beaumonti\ 12 - Adcrocuta eximia\ 13 - Felis attica; 14 - Metailurusparvulus', 15 - M etailurus m ajor; 16 - Machairodus giganteus. A - graph with all known carnivores from Samos ; B - only carnivores from recent excavation from Samos published byK o u f o s (this volume), in bold.

larteti, Ictitherium viverrinum, Lycyaena chaeretis, Belbus 2006), probably in Maragha (B e r n o r et ah, 1996), and beaumonti, Felis attica, Metailurus major. the smaller P. macedonicus (S c h m i d t -K i t t l e r , 1995) The characterization of all mentioned taxa is given in Tab. in Maramena, Greece. This medium-sized mustelid had 1, but the authors are well aware that the material was a carnivorous tendency, with the trigonid and talonid not collected from one but from several sites on Samos of almost equal size and a short M l. The postcranial and their stratigraphic position is questionable. Thus, for elements, described in more detail byR o u s s i a k i s (2002) comparative reasons, in one graph the result for the recent from Pikermi, indicate a plantigrade animal; it is therefore carnivore list following K o u f o s (this volume) is depicted, assigned to the ambulatorial terrestrial locomotion type. and in the other the result for all carnivores known from The genus Promephitis is known from Europe to Asia with Samos (Fig. 1). several species. The dental morphology is rather primitive The hyaenids were characterized byW e r d e l i n & S o l o u - in some aspects, such as a subequal paracone and metacone n i a s (1996) and according to their work: Protictitherium is on M l, or extra roots on m l (W a n g &. Qui, 2004). The insectivorous and scansorial (semi-aboreal), Plioviverrops latter is often seen in aquatic mammals as well. There is no is evaluated as insectivorous and generalized terrestrial, indication that Promephitis was in anyway a semi-aquatic Hyaenictitherium and Ictitherium as carnivorous and gen­ form, but it probably preferred a wider range of food eralized terrestrial, although the latter was described from and is therefore judged as hypocarnivorous. Promephitis Lothagam, at least partly, as a cursorial species (W e r d e ­ larteti from Samos was medium-sized compared to other l i n , 2003). Belbus and Lycaena are seen as cursorial meat Promephitis species. Parataxidea was first described by eaters, the latter as a meat/bone eater andAdcrocuta as a Z d a n s k y (1924) from China and is of Eurasian distribu­ specialized bone cracker with stout limbs (= generalized tion. Its dental structure is enforced by cingula, the first terrestrial). upper molar is broad, the P4 is low and broad, as well; a The feline forms are all hypercarnivores, very easily identi­ detailed description is given in K o u f o s (this volume-a). fied by the sectorial shape with a highly reduced talonid The morphology of the skull of the type genus induced on the carnassial. A ll have limb proportions close to the Z d a n s k y (1924) to speculate about a semi-aquatic life­ extant felids, but maybe M etailurus is not as digitigrade style for Parataxidea but unfortunately no evaluation of (G i n s b u r g , 1999). postcranial material is known so far. The teeth confirm W hile Indarctos has almost identical limb proportions as the preference of a larger variety of food, even including modern Ursus, Ursavus (P i l g r i m , 1931; G i n s b u r g 1999) molluscs or crabs (Z d a n s k y , 1924). is at least close to the same form of locomotion and both are considered as ambulatorial terrestrial. The low crown height as well as the elongated teeth with reduced shearing 3. Discussion and Comparison function are typical for hypocarnivorous forms. Promelespalaeattica (W e i t h o f e r , 1888) was found on Sa­ The carnivore guild from Samos, all described species mos, in Perivolaki, Pikermi (Greece) and Dorn-Durkheim, taken together, consisted of 16 taxa. Compared to recent Germany (W e i t h o f e r , 1888; M o r l o , 1999; K o u f o s , assemblages, this is a very good record, since extant car- 394 Beitr. Paläont., 31, Wien, 2009

bone/meat bone/meat hypercarnivorous hypercarnivorous carnivorous carnivorous 'unknown unknown hypocarnivorous hypocarnivorous semifossorial 'semifossorial Insectivorou insectivorou: cursorial cursorial 1 -3kg 1 -3 kg ambulatorical terrestrial ambulatorical terrestrial 3-10kg ^ 3-10kg generalized terrestrial generalized terrestrial 10-30kg^ 10-3 Okg scansorial 30-100kg scansorial 30-100kg' a bo real <100 kg a bo real <100 kg

Samos, Greece, MN 12 Pikermi, Greece, MN 12

Figure 2: Carnivores from Samos. 1 U rsavus cf. deperetr, 2 - Indarctos atticus', 3 - Promeles palaeatticar, 4 - Promephitis lartetv, 5 - Parataxidea maraghana\ 6 - Plioviverrops orbignyv, 7 — Protictitherium crassum; 8 - Ictitherium viverrinunr, 9 - Hyaenictitherium wongi; 1 0 - L ycyaena chaeretis\ 1 1 - Belbus beaumontr, 1 2 - Adcrocuta eximicr, 1 3 - F elis atticcr, 1 4 - M etailurus par vulus\ 1 5 - M etailurus major, 1 6 - Machairodus giganteus.

Additional elements in the Pikermi fauna: 17 - Simocyon primigenius\ 18 - Sinictis pentelicv, 19 - Martes woodwardi] 20 - Hyaenictis graeca\ 21 - Paramachairodus orientalis. nivore communities tend to have between 15 and 25 taxa. Protictitherium crassum and Belbus beaumonti. Known from Judging from the locomotor type, the carnivore guild from Pikermi but not from Samos are: Simocyon primigenius, Samos contained no semifossorial or arboreal taxa. To an­ Sinictispentelici, Maries woodwardi, Hyaenictis graeca and swer the question if any of the smaller taxa were adapted to Paramachairodus orientalis. a semi-aquatic lifestyle, further investigation is necessary, The Pikermi guild results are similar to the Samos guild it and cannot be answered in this paper. Concerning the (Fig. 2). Again the smaller forms < 10kg are missing. No locomotion, the main part of the carnivoran community arboreal or semi-fossorial forms are known and all diet belongs to the generalists, to the scansorial and cursorial classes are occupied. In Pikermi, three bone/meat eater forms. Smaller forms, the ones below three kilograms, are and three larger (30-100 kg) hypercarnivorous taxa were missing in the body mass evaluation. It seems possible that present, while in Samos only one bone/meat eater and two some were not preserved or are difficult to identify, if only larger hypercarnivorous taxa were found. fragmentary material was found. However, carnivores over Wooded environments seem to favour larger carnivores 100 kilograms were present (.Indarctos and Machairodus). from three kilograms up, but no extreme forms over 100 A ll diet classes are represented in the Samos guild. Insec­ kg. It comes as no surprise that these environments hold tivorous forms {Proticitherium and Ictitherium) were found, many scansorial forms, but also a lot of hypercarnivorous as well as taxa with a mixed diet, with a more carnivorous taxa. Savannah-like habitats are characterized by carni­ tendency, and the hypercarnivorous class, filled by the vores with a larger variety in different body mass classes felids Felis attica, Metailurus major and M. parvulus and (below 1 kg and over 100 kg) and a larger variety in food by Machairodus. One large bone/meat eater was present, preferences (insectivorous and bone/meat eater). Adcrocuta eximia. We compared the carnivoran guild from Samos with the one from Pikermi. Again, the analysis is based on mate­ 4. Results rial from several collections, summarized in the NOW database. Samos holds sixteen well defined taxa and The carnivore guild from Samos presents a very good Pikermi has twenty taxa, but three are questionable and record of Late Miocene predators in the Mediterranean only identified at genus level. Since no further informa­ area. Although information about locomotor assignment tion about food preferences, locomotion or body size was in some taxa is still missing, a general interpretation is available, these three taxa were omitted from the evalua­ possible. The carnivore community from Samos neither tion (? Enhydriodon sp., tPlesiogulo sp., Felis sp.). Although fits a heavily wooded environment like an equatorial the two carnivore communities are similar in size, the tropical rain forest (e.g. Guyana), nor a typical savannah taxa are not the same. Known from Samos, but not from like equatorial Serengeti today. The larger taxa, as well as Pikermi are: Ursavus cf. depereti, Parataxidea maraghana, the few cursorial forms, indicate areas with open habitats Nagel, D. 8 c Koufos, G.D., Carnivore Guild Structure. 395

but, at the same time, the generalists indicate the presence G in sburg, L., 1999. Order Carnivora. — [in:] Röss- of wooded environments as well. These environmental ner, G. 8c H eissig, K. (eds.). The Miocene. Land conditions fit in quite well with the proposed open bush- Mammals of Europe: 109-168. - Dr. Friedrich Pfeil land with a thick grass floor landscape for the Turolian Verlag. - München. of Samos (K oufos et ah, this volume-a). The Pikermi G unnell, G.F. 8c M orlo, M ., 2006. Comparison of carnivoran community is quite similar and mirrors the carnivore guild structure across the Paleocene-Eocene results from Samos. boundary in North America. — Climate and Biota of These are the first guild investigations for the Mediter­ the Early Paleogene Conference, Bilbao 12th to 18th ranean area of the Late Miocene. It will be interesting to June 2006, Abstracts:87. compare these in future with other sites in this very special H einrich, R.E. 8 c B ikn eviciu s, A.R., 1998. 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