Mesozoic Mammaliaforms Early evolution of mammals

1. Mesozoic Mammaliaforms Early evolution of mammals

2. http://scienceblogs.com/pharyngula/2008/05/10/monotreme_cladogram.jpghttp://scienceblogs.com/pharyngula/2008/05/10/monotreme_cladogram.jpg

3. Monotremes Placentals Marsupials Synapsid Transition Basal Synapsida Mammals Dimetrodon Edaphosaurus

4. What is a mammal?? For Linnaeus animals with hair covering the body, mammary glands to feed their offspring, and development of endothermy.

5. What is a mammal?? For Simpson and Kermack animals with a dentary-squamosal joint; the presence of one bone in the lower jaw, three auditory ossicles in the middle ear Osteological Diagnosis

6. Seven to eight bones are present in the lower jaw of synapsids Most of then decrease in size, and only the dentary increase in size Three of the bones of the lower jaw present in basal Therapsids continues in mammals dentary, which is the only bone of the lower jaw articular, which is incorporated to the middle ear as thehammer angular, also in the middle ear as thetympanicand that support the tympanic membrane quadrate, from the skull is also incorporated to the middle ear as theanvil

7. Early Mammal squamosal jaw joint dentary Changes summary Therapsid dentary Pelycosaur quadrate jaw joint dentary articular Postdentary bones

8. Mesozoic mammals at the end of the ’70s: monophyletic group, that diverged early into Prototheria/Theria, with representatives of the group already in the Triassic Living PROTOTHERIA: monotremes Living THERIA: marsupials and placentals CromptonandJenkins, 1979

9. 2 1 Mesozoic mammals at the end of the ’80s and onward: monophyletic group, without the early divergence. Pr are groups included in prototheria in the previous phylogeny, indicating that prototheria is not monophyletic Conflict in definition of Mammalia Arrow 1, Mammalia based on osteological diagnosis Arrow 2, Mammalia based on common ancestry of extant groups Rowe, 1988 Osteological diagnosis

11. EXTANT MAMMALS MONOTREMATA High degree of specialization • Only three species restricted to Australia and New Guinea • Oviparous • Aquatic and terrestrial ant-eaters platypus Long beaked echidna short beaked echidna

12. Lack of teeth, or only a couple of teeth in the juvenile platypus • sprawling posture • limb and girdles bones of robust construction

13. EXTANT MAMMALS MONOTREMATA Fossil History in the Mesozoic The earliest Monotreme is know from the Early Cretaceous of Australia. Lower jaw fragment with three distinctive teeth remarkably similar to those of the juvenile platypus Steropodon Other two species represented by jaw fragment with tooth were discovered in the Lower Cretaceous, demonstrating an early radiation of monotrems Monotremes and closely related forms only known in Australia, Madagascar and South America form a quite recently proposed group: AUSTRALOSPHENIDA (more about this latter)

15. EXTANT MAMMALS MARSUPIALIA 275 living species within two basic groups: AUSTRALIDELPHIA, mainly in the Australian region, and AMERIDELPHIA in the New World Viviparous, with short gestation period and neonates with minimal anatomical development. Most of them with abdominal pouches Variable habits: Ameridelphians are mainly arboreal and omnivorous. Australidelphians are mostly terrestrial, but some species are arboreal or burrower. Carnivorous, herbivorous and omnivorous Opposum The most primitive living marsupial is the American DIDELPHIDAE Among diagnostic features are the inflected angle of the dentary and replacement of only one tooth (the last premolar) Presence of incisors, canines, premolar and molar, with the primitive forms showing 5/4 incisors, the highest among mammals Kangaroo Koala

16. Alphadon EXTANT MAMMALS MARSUPIAL Fossil History in the Mesozoic The earliest metatheria are recorded in the Early/Late Cretaceous of North America and Asia They are represented by mandibular and skull fragments and many isolated teeth Metatherian were diverse in the Late Cretaceous of North America, but their diversity drops after the Cretaceous/Tertiary extinction. They became extinct in North America in the tertiary Probably at the end of the Late Cretaceous they reach South America Didelphodon

17. EXTANT MAMMALS MARSUPIAL Fossil History in the Mesozoic The earliest and most primitive METATHERIA (monophyletic group that includes extant marsupials and fossil forms most closely related with marsupials) is know from the Early Cretaceous in the Liaoning Province of China (125 ma) Sinodelphis is represented by nearly complete skeleton, surrounded by well-preserved impressions of fur It was a small animal (15 cm in body length) and weighed approximately 30 grams It was insectivorous and arboreal

18. EXTANT MAMMALS PLACENTALS 4162 living species in 17 groups. Cosmopolitan Viviparous with a long gestation period. Neonate may be born with minimal development (many rodents), or in a more developed stage All kind of habits and life styles The most primitive living placental in phylogenies based in morphological characters are the American Edentates (e.g. armadillos, sloths and anteaters) The most primitive living placental in phylogeny based in molecular characters are Afrotheria Placental oldest member is from the Early Paleocene

19. EXTANT MAMMALS Two recent phylogenies give different perspectives of interrelationships and origin of placental mammals Bininda Emmons et al. 2007 (Supertree) Wible et al. 2007 (Morphological)

20. EXTANT MAMMALS

21. EXTANT MAMMALS PLACENTAL Fossil History in the Mesozoic The earliest and most primitive EUTHERIA (monophyletic group that includes extant placentals and fossil forms most closely related with placentals) is know from the Early Cretaceous in the Liaoning Province of China (125 ma) Eomaia is represented by a nearly complete skeleton, surrounded by well-preserved impressions of fur It was insectivorous and a climber. It could grasp branches and move very fast in trees and on the ground. It was nearly 15 cm long and weighed about 25 grams In the same Early Cretaceous are at least three other younger representatives of the eutherians, indicating an early radiation of the group. At this time they are arboreal as well as terrestrial

22. Triconodont Holotherians Tribosphenida TOOTH HISTORY IN MESOZOIC TIMES Something like 10 to 20 years ago the changes in the molar pattern was an easy sequence to follow: First came the triconodont pattern: three main cusps in a straight line, with or without outer cingulum and accessory cusps Then in holotherians the central cusp moved outward or inward to create a triangular pattern, known as trigon/trigonid The apex of the triangle of the upper tooth points inwards, while that of the lower points outwards. This relationships is known as reversed triangle Finally the true tribosphenic molar developed: A new cusp (protocone) is added to the trigon, which occludes against a basin added in the lower teeth (talonid) The earliest record of a Tribosphenida is from the Middle Jurassic of Madagascar

23. TOOTH HISTORY IN MESOZOIC TIMES Coming back to the ’70s picture: The Triconodont pattern of teeth was typical of Prototherian, a group that included extant monotremes While reversed triangles and tribosphenic molars were found in the Theria lineage, including marsupials and placentals But in the ’80s, a close relationship between monotremes, marsupials and placentals was favoured This hypothesis gained support with the discovery in Australia of the Cretaceous monotreme Steropodon This fossil showed that monotremes had ancestors with triangular pattern of the molars,

24. TOOTH HISTORY IN MESOZOIC TIMES Until recently the earliest records of tribosphenic molars were restricted to northern continents (e.g. Asia, North America) The development of the tribosphenic molar is interpreted as key in the evolutionary success of mammals, and as having evolved once in the northern continents But in the last five years fossils with tribosphenic molars begin to appears in faunas from Australia, Madagascar and quite recently Argentina This began to change ideas about the place and time of origin of the tribosphenic molar A recent hypothesis suggests that tribosphenic molars evolved independently in two ancient groups during the Jurassic /Early Cretaceous One in Laurasia: Boreosphenida, that gave rise to marsupials and placentals, and the other in Gondwanan: Australosphenida, including monotremes

25. Mesozoic diversity of Mammaliaformes Many different groups of Mammaliaforms were represented during the Mesozoic: Stem Mammaliaformes Triconodonta Gobiconodonta Docodonta Symmetrodonta Dyolestidae Multituberculata Eupantotheria Monotremata Metatheria Eutheria Two major periods of Mammaliaform diversification: 1. From Early Jurassic to Early Cretaceous: radiation of forms that in the main did not survive past the Mesozoic 2. From the Early Cretaceous: first records of forms closely related to the three extant mammals, and the beginning of a great diversification of multituberculates Mesozoic Mammaliaformes are widely distributed around the world, Europe, Asia, North and South America, Australia and Africa In Africa are known in South Africa, Tanzania, northwestern Africa and Madagascar

26. Stem Mammaliaformes These is a paraphyletic group of basal mammaliaformes, not included in a family and with conflicting phylogenetic positions. Many of them were previously included in Triconodonta Adelobasileus, the earliest and most basal Mammaliaform, is known from the Upper Triassic of Texas (230 million years ago) Megazostrodon is the stem Mammaliaform known from the Early Jurassic of South Africa and Lesotho Morganucodon from the Early Jurassic of U.K. and China is the best known stem Mammaliaform Main cusp, one accessory anterior two accessory posterior and cingulum Dentary squamosal craniomandibular joint Presence of the promontorium Retention of postdentary bones

27. Docodonta Small group of mouse-to-rat sized animals, restricted to Early Jurassic/Early Cretaceous faunas of northern continents The best known docodontid is Haldanodon a small insectivorous from the Upper Jurassic of Portugal Lower jaw with many bones Very complex bucco-lingually expanded molars, associated with a sophisticated occlusion

28. Docodonta Castorocauda, Middle Jurassic lake beds of China; more than 6 cm in skull length and at least 42.5 cm in body length earliest-known swimming mammal with teeth for eating fish “the most primitive-known mammal to be preserved with hairs” beaver-like tail, strong arms for digging, and sharp teeth specialized for aquatic feeding, similar to the modern river otter

29. Triconodonta Considered by most authors as a paraphyletic group. This group originally included representatives of stem Mammaliaformes (Sinoconodon, Morganucodon) But they include the monophyletic GOBICONODONTIDS, group known from the Early Cretaceous of northern continents This is the only group of mammals in which replacement of molar teeth is known The angular process of the mandible is absent Gobiconodon Extremely derived morphology in the antemolar dentition (incisors looks like canine, canine is very reduced), but the molars present a generalized triconodontid pattern

30. Fruitafossor or “Popeye” 15 cm long and weighed about 30 grams. Late Jurassic (150 Ma), Colorado U. S. Single- and open-rooted tubular molars, suggesting continuous growth in life, and similar to the teeth of armadillos and aardvarks Extremely robust humerus and claws adapted for digging; vertebrae with extra joints The first known mammal adapted for feeding on ants or termites and for digging and burrowing

31. Multituberculata The most successful group of mammals to have ever lived: from late Jurassic to the late Eocene. They lived over 100 million years and survived the K/T extinction. Mostly restricted to northern continents but with some conflicting record in South America Over 200 species are known, some as small as a tiny mice, the largest the size of beavers. They were terrestrial, arboreal and burrowing They had large incisors, looking somewhat like modern rodents, and had long multi-cusped molars. Remarkable feature in the skull is a tiny jugal bone in the zygoma Arboreal multituberculates with tarsal features similar to mammals that descend tree headfirst and have prehensile tails Ptilodus

32. Multituberculata Conflict in the phylogenetic placement of Multituberculata: Are Multituberculata more closely related to theria (placentals and marsupials) than monotremes (and thus included in mammals)? Are Monotremata more closely related to theria than multituberculates (and thus the last group is outside of mammals)?

33. “Symmetrodonta”/Spalacotheroidea Many authors consider this group as paraphyletic. They are represented largely by isolated teeth and mandibular fragments. Recently was found a new species from the Early Cretaceous of China with a nearly complete skeleton and a partial skull and lower jaw Symmetrodonta are characterized by postcanine teeth with an imperfectly symmetrical, triangular arrangement of the principal cusps The front legs are interpreted as splayed out, similar to the condition of reptiles

34. “Pretribosphenid Therians” Vincelestes, from the Early Cretaceous of Argentina, is one of the best represented “pretribosphenid therians”, a paraphyletic group of mammals Vincelestes skulls is about 7cm long and the body length 30 cm, without considering the long tail Upper molars in Vincelestes show a small protocone and lower molars a short talonid with only one cusp

37. Mesozoic Mammaliaforms Epilogue Even when not commonly dominant in Mesozoic faunas, Mammaliaforms were notably diverse Most of their record is represented by isolated teeth and fragments of mandibles with teeth. In the last ten years many complete specimens of different groups have been found in China Mesozoic Mammaliaforms were extraordinarily small. The largest size (in gobiconodontids and metatherians) were similar to the Opossum (half a metre in body length) They are distributed all around the world, except in Antarctica. Outcrops with their remains are notably more common in northern continents Of the living mammals, Monotremes have representatives in the Early Cretaceous of Australia, Marsupials in the Late Cretaceous of North America and Placentals in the Early Paleocene But the earliest records of both Metatheria (group that includes marsupials) and Eutheria (group that includes placentals) occurs in the Early Cretaceous of Liaoning Province, China approximately 125 My ago The most successful group of mammals (both extant or extinct) are Multituberculates, that lived for a period of 100 My, surviving the K/T extinction Much disagreement among researchers remain regarding phylogenetic placement of Mammaliaform groups. This is particularly evident in the relationships of many Stem Mammaliaformes and in the placement of Multituberculates. There is also discussion about the monophyly of Triconodontids