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Home , Dryolestes, Dryolestida, Dryolestidae, Henkelotherium, Paulchoffatiidae, Symmetrodonta

Asociación Paleontológica Argentina. Publicación Especial 7 ISSN 0328-347X VII International Symposium on Terrestrial Ecosystems: 123-126. Buenos Aires, 30-6-2001

Mammalian fauna of the Late Guimarota ecosystem

Thomas MARTIN1

Abstract. The (Kirnmeridgian) Guimarota loca lit y near Leiria in west-central Portugal has yielded more than 800 mammalian dentaries and partial skulls representing the largest sample of Late Jurassic in the world. However, despite the enormous number of specimens collected, the mam- malian fauna appears somewhat depauperate. So far, only (several genera of ; 28% of total number), Docodonta (Halda nadan exspectaius Kühne and Krusat; 24%), and (48%) represented by Paurodontidae (Henkelatherium guimaratae Krebs and Drescheratherium acutum Krebs) and (Dryalestes leiriensis Martín, Krebsoiherium Iusiianicum Martin, and Guimaroiodus inflatus Martin) have been detected. "Triconodonta" and Syrnmetrodonta, which are well represented at other Late Jurassic localities (e.g., Morrison Forrnation), are missing. To recover the rare mammalian taxa of the Guimarota ecosystem, a project was initiated to study the nearly 7000 isolated mammalian teeth that had been obtained by screenwashing. With a single exception, the same taxa are represented. Among the thousands of isolated teeth, however, 25 lower and 23 upper premolars and mo- lars of a tiny primitive have been found, of which the lower molars closely resemble the "Porto Pinheiro ". After sorting the isolated teeth, the mammalian fauna of the Guimarota ecosystem prob- ably is completely recorded. Apparently, the coastal swamp in the Lusitanian graben where the lignite formed represented a stressed environment not appropriate for some marnmalian groups. Endemism is unlikely, beca use "Triconodonta" and were present on the Iberian archipelago in Late Jurassic-Early times, where they have been recovered from fluvio-lacustrine deposits of Porto Pinheiro (Latest Jurassic) and Galve (, Barremian).

Key words. Portugal. Guimarota. Jurassic. Mesozoic mammals.

Introduction boxes, each of which contained the phosphatic re- mains recovered in one day. Analysis of the three-di- The Guimarota coal mine near Leiria in west-cen- mensional distribution of vertebrate remains in the tral Portugal is the worlds most important locality sedimentar y body is in progress (U. Gloy, pers. for Late Jurassic mammals. From 1973 to 1982 the comm.). The thousands of isolated teeth provide the mine was exclusively worked for paleontological possibility of recovering the rare faunal elements of purposes, and it has yielded more than 800dentaries the Guimarota ecosystem that are not represented by and partial skulls of , Docodonta, and dentaries or cranial remains. Multituberculata. The most important finds were the almost complete skeleton of the paurodont Composition of the mammalian fauna guimarotae Krebs, 1991 and a partial skeleton of the docodont Haldanodon exspectatus Of the total of 6877 mammalian teeth studied, Kühne and Krusat, 1972.Despite the high number of 5783could be identified at the generic level. The dis- specimens collected, the diversity of higher taxa rep- tribution of the teeth is 48.3% Dryolestida, 42% resented at the Guimarota locality is comparatively Multituberculata, 9% Docodonta, and 0.7%, "d. low. Important groups such as triconodonts and " (figure 1). Compared to the distribution of symmetrodonts are missing. dentaries and skulIs (48% Dryolestida, 28% In addition to collecting by splitting the Multituberculata, 24% Docodonta), Docodonta is un- coal, every day 100kg of lignite were dissolved in an der-represented, which can be explained taphonom- alkaline bath and screenwashed to recover organo- ically, Docodont molars have strong roots which are phosphatic remains. Isolated bones and teeth collect- somewhat thickened at their antapical ends; there- ed through 10 years were stored in about 3500small fore, they remained much longer within the jaws and did not fall out of their sockets as easíly, for example, 'Institut für Palaontologie, Freie Universitat Berlín, as multituberculate molars. The large amount of Malteserstrasse 74-100, D-12249 Berlín, Germany. milk teeth of Dryolestida shed in vivo (n= 1543,18%

©Asociación Paleontológica Argentina 0328-347X/01$00.00+50 124 T. Martin Table 1. Genera of rnarnmals recorded frarn the Guirnarata loca- Docodonta lity. Docodonta Kretzoi, 1946 Only one docodont is present at the Haldanodon Kühne and Krusat, 1972 Guimarota locality, Haldanodon exspeciatus Kühne Multituberculata Cope, 1884 and Krusat. It is represented by a partial skeleton, nu- Pau1choffatiidae Hahn, 1969 merous dentaries, and several skulls (Krusat, 1980, Paulchoffatia Kühne, 1961 (lower jaw) Lillegraven and Krusat, 1991).The postcranial skele- Meketibolodon Hahn, 1993 (lower jaw) Guimarotodon Hahn, 1969 (lower jaw) ton indicates that Haldanodon was a ground-dwelling Plesiochoffatia Hahn and Hahn, 1999 (rn2) and probably a subterranean digger (Krusat, Xenachoffatia Hahn and Hahn, 1998a (rn2) 1991). This interpretation is supported by the fact Kuehneodon Hahn, 1969 (lower and upper jaw) that the teeth of Haldanodon often are ground down to Meketichoffatia Hahn, 1993 (upper jaw) Pseudobolodon Hahn, 1977 (upper jaw) knoblike stubs, probably by sediment adhering to the Henkelodon Hahn, 1977 (upper jaw) prey. A detailed anatomical analysis of the postcra- Kielanodon Hahn, 1987 (upper jaw) nial skeleton is in progress. Bathmochoffatia Hahn and Hahn, 1998a (MI) Albionbaataridae Kielan-Jaworawska and Ensorn, 1994 Proalbionbaatar Hahn and Hahn, 1998b (MI) Multituberculata Dryolestida Prothero, 1981 Dryolestidae Marsh, 1879 Multituberculates from Guimarota represent the Marsh, 1878 oldest unequivocal record of that group (Hahn, Krebsotherium Martín, 1999 1969).As for rodents in the Tertiary and modern ter- Guimarotodus Martín, 1999 restrial ecosystems, multituberculates are the most Paurodontidae Marsh, 1887 (includes Henkelotheriidae Krebs, 1991) diverse marnmalian group in the Guimarota locality. Henkelotherium Krebs, 1991 They are represented by 227 dentaries and skull-frag- Drescheratherium Krebs, 1998 ments whereas postcranial remains have not been Zatheria McKenna, 1975 identified with certainty. Hahn (1993),on the basis of "d. Peramus" upper jaws, distinguished four genera. On the basis of lower jaws, he distinguished five genera. All gen- of all teeth of Dryolestida), easily recognized by their era represent the Paulchoffatiidae (Plagiaulacoidea), resorbed roots, indicates that almost all original hy- involving several species. At present, only in droxyapatite available entered the record. Kuehneodon can upper and lower dentitions be corre- Because no postmortem abrasion was observed on lated. Among the isolated teeth, three additional gen- the teeth or jaws, it can be assumed that they were era of Paulchoffatiidae (see Hahn and Hahn, 1998a) not transported over long distances; instead, the and a representative of the Albionbaataridae, mammals actually lived in the coal swamp of the Proalbionbaatar plagiocyrtus, have been recently Guimarota ecosystem. named (Hahn and Hahn, 1998b)(table 1).

Guim arota-Mam malia-isolated teeth n=5783

llII Doeodonta • "et. Peramus" 9% 0.7%

o Dryolestida 48.3%

o MJltitubereulata 42%

Figure 1. Distribution of isolated rnarnmalian teeth from the Guimarata locality.

A.P.A. Publicación Especial 7, 2001 Guimarota mammalian fauna 125

Figure 2. Oblique labial view of right lower jaw (IPFUB Guimarota 19) of unnamed plesiomorphous Zatheria from the Guimarota 10- cality with P, in place and alveoli for single rooted II_4,double rooted P¡, P3" and for MI_,(al! double-rooted except for single-rooted M,). The lightly stippled areas are preserved as plastic casts of bone impressions in the matrix. Scale bar = 1 mm. Drawing by M. Bulang- Lorcher,

Dryolestida murid. Among undetermined mammalian remains from the new excavations (1973-1982)an anterior jaw Two families of Dryolestida, Dryolestidae and fragment of this taxon with three premolars exists. Paurodontidae (includes Henkelotheriidae Krebs, Study of isolated teeth from screen-washing yielded 1991), are present in the Guimarota locality. additional 23 upper and 25 lower premolars and mo- Dryolestes leiriensis Martin, 1999, with a lower jaw lars. The lower molars of "d. Peramus" exhibit an length of about 3.5 cm, is the largest and most abun- oblique crest running from the metaconid to the com- dant representative of that group; it is represented by paratively large hypoconid with a small cuspule in 89 dentaries and skull fragments. It is morphologi- the middle (Martin, in press); this is also present in cally close to Dryolestes priscus Marsh, 1878 from Peramus, Arguimus, and the "Porto Pinheiro Molar" and indicates close paleobiogeographic (Krusat, 1969;Butler, 1990).No trace of an incipient relationships. Krebsotherium lusiianicum Martín, 1999, talonid basin is present. The formula of the a1though represented by numerous dentaries and lower jaw (figure 2) is four incisors, a double-rooted skull fragments, is considerably smaller (lower jaw canine, five premolars and five molars. At present, length 2.5 cm) and is characterized by a bent para- "d. Peramus" is best placed among plesiomorphous conid at the lower molars. Guimaroiodus inflatus Zatheria (sensu McKenna, 1975). Martin, 1999is represented by only a few dentaries; its molars are characterized by inflated paraconids Discussion and metaconids. The Paurodontidae is represented by two species, Following investigation of fossils of jaws, skulls, Henkeloiherium guimarotae Krebs, 1991 and and nearly 7000 isolated mammalian teeth collected Drescheratherium acuium Krebs, 1998. Besides an al- through more than ten years of excavation the mam- most complete skeleton, a series of dentaries of malian fauna of the Guimarota ecosystem is probably Henkelotlzerium has been collected. Drescheratherium is completely recorded (table 1). "Triconodonta" and smaller than Henkelotherium and characterized by a Symmetrodonta, which are well-known from other strongly enlarged upper canine (Krebs, 1998). Jurassic localities, are missing. Although the Iberian peninsula was an archipelago in Late Jurassic times, Zatheria ("e! Peramus") it is improbable that geographical isolation is the rea- son for this reduced diversity. "Triconodonta" and Kühne (1968) reported, without further descrip- Symmetrodonta are present in the Late [urassic-Early tion, an edentulous (except for p2) lower jaw and two Cretaceous (Mohr, 1989)fluvio-lacustrine deposits of broken lower molars from the Guimarota mine, Porto Pinheiro (some 70 kilometers southwest of the which he attributed to "d. Peramus", Due to the frag- Guimarota locality) (Krusat, 1989) and in the Early mentar y nature of the fossils this assignment re- Cretaceous of Galve, Spain (Krebs, 1985). These flu- mained questionable and Sigogneau-Russell (1999) vio-lacustrine beds cover the catchment area of a contradicted identification of the molars as pera- much wider hinterland than the swamp deposits of A.P.A. Publicación Especial 7, 2001 126 T. Martin the Guimarota ecosystem and therefore contain fau- Krusat, G. 1980, Iifllrlr/nQáQfl '?íwect¡¡tu$ Kühne & Krusa] 1972- nal elements with differing ecological requirements. (Mammalia, Docodonta). Contribuicao para o Conhecimento da Fauna do Kimeridgiano da Mina de Lignito Guimarota Apparently, the Guimartoa coal-swamp was a (Leiria, Portugal) IV Parte; VIII. Memórias dos Seroicos geologi- stressed biotope not suitable for all mammalian cos de Portugal, (nava Sér.) 27: 1-79. groups. The excellent preservation of bones and teeth Krusat, G, 1989. Isolated molars of a triconodont and a sym- at Guimarota makes it probable that the recorded metrodont (Mammalia) from the uppermost Jurassic of Portugal. Berliner geowissenschaftliche Abhandlungen A 106: 277- mammals lived in place and were not transported in- 289. to the area of sedimentation. The mammalian fauna Krusat, G. 1991. Functional morphology of Haldanodon exspectatus of the Guimarota locality indicates paleobiogeo- (Mammalia, Docodonta) from the Upper Jurassic of Portugal. graphical relationships with westem North America Fifth Symposium on Mesozoic Terrestrial Ecosystems and Biota. Extended abstracts. Contributions of the Paleontological (: e.g., Dryolestes, Taihiodon- Museum, University of Oslo 364: 37-38. Pelicopsis and Henkelotherium) and southem England Kühne, W.G. 1961. Eine Mammaliafauna aus dem Kimeridge (e.g., Albionbaatar and Proalbionbaatar). Portugals. Vorlaufiger Bericht. Neues Jahrbuch für Geologie und Poldontotogie. Monatshefte 7: 374-381. Kühne, W,G. 1968. Kimeridge mammals and their bearing on the References phylogeny of the Mammalia. In: E,T. Drake, (Ed.): Evolution and environment: 109-123. New Haven, London (Yale Butler, P.M. 1990. Early trends in the evolution of tribosphenic University Press). molars. Biological Reviews 65: 529-565. Kühne, W.G. and Krusat, G. 1972. Legalisierung des Taxon Cope, E.D. 1884. The Tertiary Marsupialia. American Naturalist 18: Haldanodon (Mammalia, Docodonta). Neues Jahrbuch für 686-697. Geologie und Palaontoíogie, Monatshefte 1972: 300-302. Hahn, G. 1969. Beitrage zur Fauna der Grube Guimarota Nr. 3. Lillegraven, J.A, and Krusat, G. 1991. Cranio-mandibular anato- Die Multituberculata. Palaeontographica A 133: 1-100. my of Haldanodon exspectatus (Docodonta; Mammalia) from Hahn, G. 1977. Neue Schadel-Reste von Multituberculaten the Late Jurassic of Portugal and its implications to the evolu- (Mamm.) aus dem Malm Portugals. Geologica et Palaeontologica tion of mammalian characters. Contributions lo Geology, 11: 161-186. University of Wyoming 28: 39-138. Hahn, G. 1987. Neue Beobachtungen zum Schadel- und Gebíss- Marsh, O.e. 1878. Fossil from the Jurassic of the Rocky Bau der Paulchoffatiidae (Multituberculata, Ober-Iura). Mountains. American Iournal of Science 3, 15: 459. Palaeovertebrata 17: 155-196. Marsh, O.e. 1879. Notice of new Jurassic Mammals. American Hahn, G. 1993. The systematic arrangement of the [ournal of Science 3, 18: 396-398, Paulchoffatiidae (Multituberculata) revisited. Geologica et Marsh, O.e. 1887. American Jurassic Mammals. American ioumal Palaeontologica 27: 201-214. of Science 3, 33: 326-348. Hahn, G. and Hahn, R. 1998a. Neue Beobachtungen an Martin, T. 1999. Dryolestidae (Dryolestida, Mammalia) aus dem Plagiaulacoidea (Multituberculata) des Ober-Juras 3. Der Bau Oberen Jura von Portugal. Abhandlungen der senckenbergischen der Molaren bei den Paulchoffatiidae. Berliner geowis- naturforschenden Gesellschaft 550: 1-119. senschaftliche Abhandlungen E 28: 39-84. Martin, T. in press. New stem-lineage representatives of Zatheria Hahn, G. and Hahn, R. 1998b. Neue Beobachtungen an (Mammalia) from the Late Jurassic of Portugal. [oumal of Plagiaulacoidea (Multituberculata) des Ober-juras 4. Ein Vertebrate Paleontologu. Vertreter der Albionbaataridae im Lusitanien Portugals. McKenna, M.e. 1975. Toward a phylogenetic classification of the Berliner geowissenschaftliche Abhandlungen E 28: 85-89. Mammalia. In: w.P. Luckett and ES. Szalay (eds.), Phylogeny of Hahn, G. and Hahn, R. 1999. Nomenklatorische Notiz: Namens- the Primates: 21-46. New York, Plenum Press. Ánderung bei Multituberculata (Mammalia). Geologica et Mohr, B.A.R. 1989. New palynological information on the age and Palaeontologica 33: 156. environment of Late Jurassic and Early Cretaceous vertebrate Kielan-Iaworowska, Z. and Ensom, P.e. 1994. Tiny plagiaulacoid localities of the Iberian Peninsula (eastern Spain and multituberculate mammals from the Purbeck Limestone Portugal). Berliner geowissenschaftliche Abhandlungen 106: 291- Formation of Dorset, England. Palaeontology 37: 17-31. 301. Krebs, B. 1985. (Mammalia) aus der Unterkreide von Galve Prothero, D.R. 1981. New Jurassic mammals from Como Bluff, (Provinz Teruel, Spanien). Berliner geowissenschaftliche Wyoming, and the interrelationships of non-tribosphenic Abhandlungen A 60: 29-48. Theria. Bulletin of the American Museum of Natural History 167: Krebs, B. 1991. Das Skelett von Henkelotherium guimarotae gen. et 281-325. sp. nov. aus dem Oberen Jura von Portugal. Berliner geowis- Sigogneau-Russell, D. 1999. Réévaluation des Peramura senschaftliche Abhandlungen A 133: 1-110. (Mammalia, Cladotheria) sur la base de nouveaux spécimens Krebs, B. 1998. Drescheratherium acutum gen. et sp. nov., ein neuer du Crétacé inférieur dAngleterre et du Maroe. Geodiversitas Eupantotherier (Mammalia) aus dem Oberen Jura von 21: 93-127. Portugal. Berliner geowissenschaftliche Abhandlungen E 28: 91- 111. Kretzoi, M. 1946. On Docodonta, a new order of [urassic Mammalia. Annales Historico-Naturales Musei Nationalis Hungarici 39: 108-111. Krusat, G. 1969. Ein Pantotheria-Molar mit dreispitzigem Talonid aus dem Kimmeridge von Portugal. Pataontologíeche Zeitschrift 43: 52-56. Accepted: December 20'h,2000.

A.P.A. Publicación Especial 7, 2001