Midterm - Lecture Flashcards

(145 cards)

1
Q

What epochs are mammals present in?

A

Paleocene = 66 – 56 Ma
Eocene = 56 – 34 Ma
Oligocene = 34 – 23 Ma
Miocene = 23 – 5.3 Ma
Pliocene = 5.3 – 2.6 Ma
Pleistocene = < 2.6 Ma

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2
Q

What do boundaries in eons, eras, periods and epochs represent?

A

Major geological and paleontological events
- example: mass extinctions

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3
Q

What ERA is really important in the primate evolution?

A

The Cenozoic! (This is the last 65 million years when mammals were initially created)

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4
Q

What EPOCH is very important for primate evolution?

A

The Miocene Epoch
- 23 - 5.3 Ma
- Early Miocene (23 -16 Ma)
- Middle Miocene (16 - 11.6 Ma)
- Late Miocene (11.6 - 5.3 Ma)

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5
Q

What era is the suggested OLDEST fossil primate found in?

A

Up to 65 Ma (Paleocene)
- They are often controversial due to not being considered “true primates”
- Major lineages of primates split fairly early on too
- By the end of the Eocene, there are major primate groups that exist

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6
Q

When is it suggested that major primate lineages began to split?

A

Fairly Early
- Paleocene to Early Eocene
Potentially up to 65ma

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7
Q

How long ago do they believe primates became an order? How do they know this?

A

-Believe as long ago as 90 Ma
- This is through the help of molecular dating (use of DNA to estimate species divergences), but there is a possibility that it is TOO old
- Likely that they will find older fossils eventually to help them with this
- currently no evidence to support this

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8
Q

What is Stratigraphy?

A

The study of rock layers
(This is how we know how old fossils are with the help of the principle of superposition)

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9
Q

What is the principle of superposition?

A

Older layers deposited first, covered in younger layers over top
- Gives a suggestion on the relative age of certain things

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10
Q

What is Provenience?

A

The original source of a fossil. Where it came from

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11
Q

What are some dating techniques used to help identify remains?

A
  • Absolute dating techniques (gives an actual age of something)
  • Relative Dating techniques (age estimate relative to other sites and species

Regardless, need to know where the fossil came from

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12
Q

What radiometric dating techniques do they use to find primate fossils?

A

Argon-Argon dating
- can only find these when there is sub fossil sites and rocks are formed suddenly (Volcanism!) this is why you see more fossils in Asia and Africa

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13
Q

What is Paleomagnetism?

A

Magnetic field that aligns rocks to form the N/S poles of that time
- changes over times, great way to age a fossil

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14
Q

What is Faunal Correlation?

A
  • This is Biostratigraphy
  • A correlation you can use for other components of undated species to identify the certain time period.
  • Ex. SHELLS
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15
Q

What are the best dating techniques for primate fossils?

A

Lithostratigraphy (Stratigraphic positions)
Chronostratigraphy (Radiometic and Magentostratigaphy)
Biostratigraphy (Faunal Correlation)

When they are all combined together you get the best suggestion of an age of a fossil

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16
Q

What was the distribution of fossil primates?

A

From far north to far south
- Early primates were all the way up into Alberta and Saskatchewan
North America and Europe is where the oldest primate fossils are located

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17
Q

How did primates get to North America?

A

From the help of the continental drift, North America was still connected to Eurasia (Europe)

Climate has had slow cooling, only in the Oligocene did we see it to begin to drop. There would have been many tropical climates with thick forests which are perfect climates for primates

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18
Q

What epoch would primates have still been in the Northern parts of the world

A

The Palaeocene, still very high temperatures

Plate Tectonics and Climate Change is particular reasons as to why they are no longer alive in the Northern hemispheres

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19
Q

What is 2 main things in the primate fossil record?

A

Biased
- Body parts are often easier than others (specifically teeth)
- Body sizes, much easier for larger species to preserve than older species
- locations!
Incomplete
- many species and ghost lineages that we have not found that link lineages together
- many of the reasons for incompleteness is due to environment, needs to be in perfect condition in order to preserve

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20
Q

Who is in Strepsirrhines?

A

Lemurs, Lorises and Galagos (Lemuriformes and Lorisiformes)

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21
Q

Who is a part of Haplorhinies?

A

Tarsiers, New World Monkeys, Old World Monkeys and Apes
(Tarsiiformes, Platyrrhini (NWM), Catarrhini (OWM) )

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22
Q

What is the Evolutionary Taxonomy? What is the Phylogenetic Systematics?

A

Older: Prosimians and Antrhopoidea (Tarseris were in the Prosmii)

Newer: Strepsirhini and Haplorhini (Tarsiers and now in Haplorhini with monkeys, apes, and humans)

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23
Q

What is a Cladogram

A
  • A classification that groups animals together based on shared traits (common descent)
  • BRANCHING PATTERN
  • FOSSIL CLADOGRAMS ARE TENIOUS HYPOTHESIS ONLY
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24
Q

What is a phylogenetic tree?

A
  • genetically different
  • time differences
  • morphological and/or molecular data
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25
What is a primitive trait?
A Plesiomorphy (ancestral state) ex. HAIR
26
What is a derived trait
Apomorphy! - a new trait that is not present in the ancestral taxon ex. Bipedalism in Hominins
27
Apomorphy Traits of Primates
- Forward facing eyes - opposable thumbs - nails - reduced reliance on olfaction -post-orbital closure - Big brains relative to body size - Dental Formulas with a reduction in incisors (2.1.2.3 or 2.1.3.3)
28
Plesiomorphys of primates
An ancesteral character traiT 5 fingers generalized skeleton hair retained clavicle
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Symplesiomorphy
A shared primitive trait "-mplesio-" - not helpful in creating cladograms
30
What is a Synapomorphy?
Shared derived trait "napomor" - helpful in creating cladograms Rare (Large brains, arboreality, social behaviours etc. but characterize MOST primates)
31
Homoplasy
Convergence evolutions in traits not shared ancestry, but it was evolved
32
The 4 Complexes of Primate Characters:
1) Hands and Feet (nails, opposable) 2) Eyes and Orbits (Vision, Coloured) 3) The Ear (Auditory Bulla) 4) Dental and Postcranial Features Also: dental formula, molar morphology (low profile, expanded talonid), large brains, forward facing eyes, arboreality
33
Differences in Vision between all primates and higher primates (Anthropoids)
They all have postorbital bars Higher Primates have postorbital closure which includes a bony cup Non-primate mammals only have postorbital process This is a derived trait!
34
What is special in primate ears?
Auditory Bulla: the structure that houses middle and inner ears BULLA IS AN OUTGROWTH OF THE PETROSAL (temporal) BONE in primates
35
What is found in primate dentition
Ancestral Mammals: 3.1.4.3 Streps and Plats: 2.1.3.3. Catarrhines: 2.1.2.3 Molars have lower crests, there is more crushing rather than cutting
36
Primate Synapomorphies:
1) Post-orbital bar or closure 2) Middle and inner ear housed in an outgrowth of petrosal bone 3) Opposable 1st digits (on hands and feet) 4) Nails on at least some digits
37
Strepsirrhine Features:
- Split Nostiles - Long Snout - Post-orbital Bar - Unfused metopic suture (separate L/R frontal bones) - Unfused mandibular symphysis - Ectotympanic ring Tooth Comb - Grooming Claw on second toe bicornate uterus minimal parental care
38
Haplorhine features:
Unsplit nostrils Dry nose Short Snout Post-orbital closure fused metopic suture (single frontal bone) fused mandibular symphysis (except tarsiers) Ectotympanic tube No tooth-comb No grooming claw Unicornate uterus (except tarsier) extended parental care
39
Strepsirrhine vs Haprlorhine Reproduction differences
Steps - more layers between maternal and fetal blood which decreases transfer rates - placentas are around most of uterus - Bivornuate "two-horned" uterus
40
Tarsier features
Strep features - unfused mandibular symphais and frontal bone - Bicornate uterus - grooming class on 2nd toe Hap features * Dry nose, unfused to upper lip * NEAR postorbital closure * No tooth comb * Hemochorial placenta * No tapetum lucidum Unique tarsier features: * Huge eyes * Specialized ankles – leaping * Another grooming claw on 3rd toe! * Dental formula
41
What are the humps in an upper molar?
Trigon - Paracone - Metacone - Protocone Then the: - Paraconule - Metaconule
42
What are the ridges in a lower molar?
Trigonid - Protoconid - Paraconid - Metaconid Talonid Basin - Hypoconid - Hypoconulid - Entoconid
43
What does Buccal mean? Mesial? Lingual? Distal?
Mesial – toward anterior or mid-sagittal Distal – away from anterior or mid-sagittal Buccal/Labial – toward the cheeks or lips Lingual – toward the tongu
44
What is the tip of the trigon / trigonid always
The Protocone / Protoconid
45
What can help determine a fossil mammal?
Jaw Joint is a single bone. The jaw bones that were in non-mammals shrunk and moved to the middle ear in mammals Differentiated Teeth - Heterodont (non-mammal, teeth come back) - Diphyodont (2 sets of teeth deciduous and adult) Molar Occlusion (chewing ability) Secondary Bony Palate (separates wind pipe from food pipe) Two Occipital Condyles
46
When do the earliest representatives of living mammals become introduced?
The paleocene
47
What is considered a Mammalian Molar?
- Moves from singular cusped (only protocoled) to TRIBOSPHENIC ('tri' cusped, protocoled, paranoid, metaconid) - Upper molars have basic triangle cusp - Lower molars are triangles and extra teeth on back (how to grind the food efficiently)
48
Who is believed to be the mammalian cousin of Primates?
The Euarchontoglires - Scandentia (tree shrews) - Dermoptera (colugos) - Rodentia (rats, squirrels, beavers) Lagomorpha (rabbits and ochontids)
49
What are the Living Euarchontas?
Dermoptera ("Flying Lemurs") Scandentia (Tree Shrews) Primates Extincts: Plesiadapiformes
50
What are Dermoptera
Colugos or Flying Lemurs SE Asia Not diverse. Only two extant genera – Cynocephalus - Philippines – Galeopterus – mainland SE Asia, Borneo, Sumatra Gliders – Best mammalian gliders – Large flaps of skin between legs and tail, but also between fingers – aka ‘mitten glidders’ - Forward facing eyes
51
Physical Characteristics of Dermoptera:
* Tooth comb (but not like strepsirhines! Independent) each incisor has a tooth comb * No nails and no opposable hallux (poor climbers) * Incomplete post-orbital bar – Variable between species * Middle ear different from primates (not in Peterus bone)
52
What are Scandentia's?
* Tree Shrews * Southeast Asia * Used to be considered primitive primates * Insectivores and frugivores * Terrestrial and arboreal * Diurnal and nocturnal * About 20 living species Physical Characters: * Post-orbital bar * Middle ear morphology superficially similar to primates (but not a petrosal bulla) * Relatively large brain * Some have binocular vision * Some are arboreal * No nails and no opposable hallux
53
Physical Adaptations that help primates arboreality
Think this due to primate ancestors likely being this - Grasping hands and Opposable Hallux - Mobile Shoulders and Hips - Stereoscopic vision - Prehistoric tail Fruit eating?
54
What do we think goes along with arboreality?
- More primitive primates live in extreme forested areas - earliest primates likely arboreal Closest relatives are generally arboreal (e.g. dermopterans, some tree shrews) * Many early mammals were arboreal as well SMALL BODY SIZES CONNECTED WITH MOST PRIMITIVE PRIMATE ANCESTORS TOO
55
What are main traits that are looked for when trying to find a primate ancestor?
* Grasping extremities * Nails instead of claws * Optical convergence * Enhanced vision * Enlarged brains * Arboreal what could have led to these?
56
What are the 4 origin theories of primates?
* Arboreal Theory * Visual PredationTheory * Angiosperm Radiation Theory * Terminal Branch Theory They are not static so it is hard to test We are talking about Basal primates that look like modern primates Fossils are essential to determine these models!
57
Arboreal Hypothesis:
1920s - argument that primates evolved to live arboreal habitats (most early mammals were this, but primates stayed) - 3D environments with high risks - Grasping hands, stereoscopic vision are assets Problem: other mammals live like this without all primate adaptations and are just as well in trees (ex. squirrels)
58
Visual Predation Hypothesis:
1960-70s LEAST ACCEPTED HYPOTHESIS * Primate pattern represents an adaptation to forage for insects in the terminal branches of trees * Based on comparative method – Found similarities to other visual predators (felines, raptors) * Problems: few living primates are visual predators, why primate dental changes if eating insects?
59
Angiosperm Radiation Hypothesis:
1991: Sussman, COEVOLUTION THEORY * Fruit, flowers (nectar) offered new niches leading to primate specializations * Primates’ roles as pollinators and seed dispersers may have helped spread of flowering trees * Vision for distinguishing fruit in low light, grasping hands for better handling * Problems: flowering plants evolved before primates, early primates not nectar specialists, don’t need acute vision for fruit eating
60
Terminal Branch Hypothesis:
* Primate specializations evolve to move primates into the fine branches of trees using ‘acrobatic’ movements * Active locomotion in fine branches accounts for: – Grasping hand and feet – Hindlimb dominance – emphasis on vision – forward rotation of the orbits – stereoscopic vision – enlargement of the brain (especially visual cortex) * Applies to both predation and herbivorous diets – Many evolved in sequence
61
Where are most Plesiadapifrom primates found?
66-56 until about <40 my In North America and Europe (all above the equator
62
What are Plesiadapiforms?
Primate-like Mammals - HIGHLY SIMILAR to primates, but relationship is disputed - Originally considered primates, then placed into own order as sister taxons of Primates - Now many think that some of these Plesiadapiforms are EARLY PRIMATES Most common mammalian group in North America Paleocene
63
Adaptive Radiation of Plesiadapiforms
* Very diverse “order” * More than 140 species * 11 families * Large range of body size (from smaller than any living primate to ~ 3 kg) * Diverse dietary adaptations (dental diversity, size diversity) * Locomotor diversity (arboreal, terrestrial, gliding?) * Probably went extinct due to competition from rodents
64
Dental Characterisics of Plesiadapiforms
First things found of them, lots of them Molars are similar to primates - expanded talon basins - unreduced lower m3 - bunodont cusps are dull - low trigonids
65
Cranial Features of Plesiadapiforms? Post-Cranial Features?
* No post-orbital bar * Small brain * Wide zygomatic arches with divergent orbits * Long snouts * Low flat skulls * Petrosal auditory bulla – Long debate due to fragmentary material and fusing of sutures No nails, most retain claws No grasping hallux in most Arboreal majority
66
Similarities and Differences of Plesiadapiforms to Primates
Primate similarities: * Bunodont molar cusps * low trigonids and basin- shaped talonids * Petrosal bulla Primate dissimilarities: * no postorbital bar * non-convergent orbits * large rodent-like incisors * non-opposable hallux * no hindlimb dominance * claws instead of nails * 3.1.3.3 primitive tooth count (early primates 2.1.4.3)
67
What are the Fossil Plesiadapidiformes discussed in class?
Carpolestidae Paromomyidae (Phenacolemur) Purgatoriidae (Purgatorius)
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Carpolestidae
Plesioadapiforme North America and Asia Specialized Dentition - Enlarged premolars - Lower p4 is largest tooth - Upper P3 and P4 Dentition may have something to do with their diets, harder fruits tend to challenge accessibility
69
Paromomyidae
North America and Europe Early Palaeocene to Eocene (very long living group) Enlarged incisors - uppers have multiple cusps - lowers are huge and procumbent some have enlarged p4 Live at fairly northern latitudes - Ellesmere Island - Calgary, and Southern AB NOT OMOMYOIDS
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Genus Phenacolemur
From the Family Paromomyidae Well known post cranial remains Beard suggests gliding adaptations Close relationship to Demopterans? May be too large and robust due to new skeletal finds
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Purgatoriidae
IMPORTANT FAMILY Genus is Purgatories Early Palaeocene, North America 5 species Earliest group Potential ancestor to both Plesiadapiformes and Primates supported by some post-canine dentition - 3.1.4.3 in P. janisae - low molar trigonids - broad second lower molars - elongated third molars - specialized incisors which makes unlikely to be ancestral primates * I1, I2, and I3 are different sizes * I1 and I2 likely larger than the canine (a derived plesiadapiform character not found in primates) * Based on analysis of alveoli only
72
Why is there a common view that Plesiadapiformes are not Primates?
* Lacking primate synapomorphies * Too derived to be ancestral to primates * Have many of their own specialized dental features * Generally recognized as their own order – Plesiadapiformes –within Euarchontaglires
73
What did they find in NOVA in the Clark Fork Basin
Late Paleocene 3 partial skeletons (Paromomyidae, Micromomyidae, Carpolestidae)
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What are Euprimtes
'True Primates' Appear suddenly at the beginning of the Eocene 56MA (warm, humid) Appear simultaneously in North America and Europe (land bridge) Primate SYNAMPORPHIES: PO Bar, Nails, Larger brains, Stereoscopic Vision, less olfaction 50 genera from Eocene alone
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Eocene Primates
Diverse of THE primates - Adapiformes - Omomyiformes Sparse, but growing rapidly - Lorisoids - Tarsiers - Earliest Anthropoids
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Adapoids vs Omomyoids
Adapiformes (Streps) - small eyes (diurnal) - long snout - small incisors, large canines - ring in ear - general arboreal quadrupeds - premolars and molars have shearing crests Omomyiformes (Haps) - large eyes (nocturnal) - short face / snout - elongated tarus (ankle complex that are in modern tarsiers and lorises) - ear tube - large incisors, small canines - small species have sharp pointed molar cusps, Large later species with flat molar teeth
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What are Adapiformes?
Suborder: Strepsirhini Infraorder Adapiformes Superfamily: Adapoidea (Adapoids or adapiforms 6 families Similar to modern lemurs Larger bodied than other Eocene primates (range of sizes similar to living lemurs; 100g – ~ 7 kg) Reconstructed as largely diurnal Derived adapiforms can sometimes look a bit like anthropoids (convergence, but NOT ANCESTRAL)
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What are the 6 families of Adapiformes?
1) Notharctidae – mostly Nth America - Frugivorous or folivorous, large (1.5 kg – 7 kg) - Arboreal quadrupeds (including leapers), most like living lemurs 2) Cercomoniidae – mostly Europe – Dietarily diverse – insects through leaves, size diverse (100 g – 4 kg) 3) Caenopithecidae – Nth Am, Asia, Eur, Africa (widespread!) - Highly varied diets (500g – 3.5 kg) 4) Asiadapidae – India, late Eocene - Oligocene – Poorly known – 2 genera (200-600g) 5) Adapidae – mostly Europe (France) – Folivorous, medium to large (500 g – 4 kg), this was the first family name 6) Sivaladapidae – Eocene to MIOCENE in Asia – Folivorous, large (2.5 kg – 4.5 kg) - first in India, now widespread
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Dental Characteristics in Adapiforms
Unfused mandible (Fused in some later forms) Large unspecialized canines 2.1.4.3 dental formula in primitive forms (Later have 2.1.3.3 or even 2.1.2.3, like Catarrhines) Small lower incisors (No tooth comb) Hypocone is variable – None – True hypocone (OUTGROWTH from lingual cingulum) in CERCOMONIIDAE AND CAENOPITHECIDAE - Pseudohypocone (offshoot from protocone, separated from cingulum)
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What is a True Hypocone
An outgrowth of the lingulum cingulum
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Pseudohypocone
A fake Hypocone / Nonnopithex fold offshoot from the protocone Separated from the cingulum
82
What is a Cingulum?
A ridge of the tooth
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Cranial Characteristics of Adapiformes?
- All have post orbital bars - Slightly convergent orbits - Large brains - Petrosal bulla
84
Cranial Characteristics of Adapiforms? Postcranial Characteristics?
Post-orbital bar (not closure) Slightly convergent orbits Large brain Petrosal bulla Divergent hallux and pollex Nails Long trunk Long tail Long hindlimbs (Hindlimb dominance)
85
Notharctus (Notharctidae)
North American genus Multiple species, 3 – 7 kg Folivorous (Fused mandibular symphysis to allow more chewing power, shearing crests on molars) Sexually dimorphic (clear example of one of the first sexually dimorphic) Overall appearance somewhat Lemur-like Multiple almost complete skeletons Like Lemur or Varecia in limb proportions and architecture (Long hindlimbs and Long thorax) Mostly leaping? Foot morphology suggests vertical clinging and leaping (arboreal quads) Grooming claws in some Robust 1st metatarsal and stronger hallux grooming claws present, unsure of which toe
86
Donrussellia (Cercamoniidae)
Earliest (E. Eocene) and most primitive cercamonid 2.1.4.3; No hypocones; Simple trigonids Small body size with high-crowned cusps on premolars and molars suggests Insectivorous? Most primitive adapiform? Some phylogenies place it as a stem to all streps Primitive low molars with a simple trigonid and broad talon basin
87
Sivaladapidae
Adapoid Earliest are Eocene, but some lineages survive until the late Miocene, persists for 30my, not overwhelming amount Named after hide God Sheena Many genera – Sivaladapis = India, Pakistan, – Indraloris = India, Pakistan – Sinoadapis and others = China Survived global cooling by moving to tropics? Larger than early adapoids Fused mandibles Likely leaves and fruits
88
Darwinius maxillae (Ida)
Most complete fossil ever found, 1983 found from a private collection Middle Eocene 47 Ma in Messel, Germany Great preservation due to swamp environment
89
What did the press say about "Ida"? What did the Scientific Paper say?
It was everywhere, conferences, books, tv specials Created it to be more than a science event Stated that it was an Anthropoid Peer-Reviewed Journal A ceracomine adapoid (now seen as caenopithecid) Allows new information to show what early Eocene primates were like Conclude that: ALL adapiforms are likely more closely related to haplorhines than strepsirrhines (says not Anthropoid, but it is if it is a hap)
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Why is it suggested that they are calling "Ida" a Haplorrhine?
It is juvenile Lots of 'traits' that we see in Haplorrhines are argued to be seen in the fossil, but cannot say that it is like this in adulthood due to it not being an adult or the traits are not actually preserved
91
Afradapis
Another caenopithecid (Caenopithecidae) Fayum of Egypt, 37 Ma (later Eocene) Relatively large (2.5 – 3 kg) Living with early anthropoid primates (looks primitive) Highly specialized dentition: – Honing p3 and enlarged canines – Loss of P2 and p2 (2.1.2.3) – Folivory features on molars Convergent dental features with later catarrhines – Adapiforms go extinct due to competition? Demonstrates the highly varied nature of adapoid primates. – Complex biogeography – Multiple convergences with anthropoids, likely due to diet
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Adapids and Strepsirhinis
Share: * Petrosal bulla * Postorbital bar * Nails instead of claws * Divergent hallux and pollex * Ectotympanic ring * Similar molars with hypocone * Relatively long snout, many turbinates (well- developed olfaction) Unique Features of Streps - Tooth Combs - Grooming Claws
93
What are Omomyiformes?
Superfamily: Omoyoidea Suborder: Haplorhini 2 families (Omomyidae, Microchoeridae) Eocene radiation (Eocene: 54-34 Ma) North America, Europe, and some in Asia Diverse – 2 families, ~ 45 genera Generally small-bodied (<500g) – Smaller than most adapoids Insectivore/frugivores, ‘Tarsier-like’ (or ‘galago-like’) primates NOCTURNAL
94
Omomyidae vs Microchoeridae
The two familys in Omomyiformes Omomyidae: mostly N. America – Subfamily Anaptomorphinae (Nth Am, except Teilhardina and Baataromomys) Nocturnal, frugivorous or insectivorous, small (30 g – 600 g) Examples: Teilhardina (Europe, Nth Am, and Asia) – Subfamily Omomyinae (Early Eocene-Late Oligocene – Nth Am, Asia) Diurnal?, diverse diet, larger (100 g – 2 kg), leaping Microchcoeridae: mostly Europe - Generally smaller, nocturnal, diverse diet, diverse size (40 g – 1.8 kg), leaping
95
Dental Characters of Omomyoids
Large procumbent lower central incisors (no tooth comb), gap between the 2 front incisors (lip connected to nose?) Small canines (often “premolarized”) No canine sexual dimorphism True hypocone in many, but not all – Nannopithex-fold in some Lower molars have low trigonids and broad talonid basins V-shaped (Bell) tooth row Primitive forms have 2.1.4.3 Later forms 2.1.3.3 (acquired independently) and some are 2.1.2.3 Some reduce anterior dentition Large molars Unfused mandible Paraconids disappear (shorter front-back trigonid)
96
Cranial and Post-cranial Characteristics of Omomyiforms
Cranial: Postorbital bar, forward orbits, large brain, petrosal bulla Enlarged orbits Central protruding incisors Large brain Short snout with broad palate Mandibular symphysis unfused Postcranial: Limited remains! Nails and opposable hallux Elongated calcaneus (ankle region) Navicular often also elongated Some have fused distal tibia and fibula (or strong fibrous connection) These two things are connected to Modern clinging and leaping primates Generally looks like leaping adaptations More similar to cheirogalids (mouse lemurs) than to Tarsius
97
Ear morphology of Omomyoids
Ectotympanic ring fused to auditory bulla (See Fig 2.20) – with small bony tube in some species (external auditory meatus) Blood supply to the brain made up of both the stapedial and promentory arteries (see Fig 2.13 in textbook). Ties to Haps, but may be convergent
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What are the differences in structure of the Tympanics across Primates?
Lemuriforms: ring hanging within bulla cavity Lorisiforms: ring attached to inside wall of bulla NWM: ring attached to external wall Tarsiers and catarrhines: attached to external wall and extended into a tube – external auditory meatus
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Cranial Blood Supply in Primates
Stapedial blood supply (in ear) is in lemurs Us, Monkeys, and Apes have a protomontory branch of internal carotid - Stapedial artery absent in adults Lorisifomes and cheirogaleids are WEIRD with an External carotid supply with ascending pharengeal artery Eocene Euprimates typically have both a stapedial and promontory artery
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Telihardina
Primitive Anaptomorphine (Omomyidae) * Earliest and most primitive * Early Eocene of North American AND Europe and Asia * 2.1.4.3. dental formula * Orbits small: Diurnal? * Some consider probable ancestor of later omomyoids Waste basket taxon: chucks anything into it because they do not know what to do Teilhardina (omomyoid) looks like Donrussellia (early adapoid) But: jaws & snout are receding in omomyoids – trigonids mesiodistally compressed – small relative to talonids – p4 mesiodistally compressed – but buccolingually wider
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Anatopmorphines
Mostly Nth Am; Early Eocene common, rare by middle Eocene * Possible that early anaptomorphines are ancestral to both microchoerids and omomyines * Detailed record of speciation in Wyoming * Anterior teeth change in several lineages – note small canines * Some premolars lost, some enlarged * Hard to differentiate incisors, canines, and premolars (especially when some premolars are lost)
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Omomyinae
* Middle-late Eocene Nth Am (later than anaptomorphines) * Likely descendants of anaptomorphines * More diverse (body size, dental adaptations) * Steinius – 2.1.4.3. – Probably ancestral omomyine * Omomys – 2.1.3.3. – Slight p4 enlargement * Macrotarsius – Nth Am and China – Large plant eater (~2 kg)
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Microchoeridae
Family of omomyoids * Western Europe only * Early Eocene – early Oligocene * Tiny to medium-sized * Traditionally, researchers thought strong links to Tarsius * Could derive from European populations of Teilhardina * Relatively abundant in the fossil record * All have hypocones, unlike Tarsius * Nannopithex has just a Nannopithex fold – Nannopithex is most primitive * V-shaped palate * Very large eyes (not as big as Tarsius) * Long tarsals * Tibia-fibula fused * Tympanic ring within the bulla, but with a bony tube
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Similarities between Omomyoidea and Haplorhini
* Reduced facial and nasal region * Relatively reduced olfactory bulbs – Olfactory bulb lies above interorbital septum * Shape of the talus * Typmanic ring fused to bulla wall in some; within the bulla in others, and microchoerids have a tube * Blood supply - both stapedial and promontory arteries
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Omoyoidea and Tarsius
Share: * Rel. large orbits – BUT not structurally similar – Tapetum lucidum retained? * Tubular ectotympanic in some – BUT combined with an unfused ring – possibly convergent evolution * Elongated tarsals – BUT not long enough * Tendency toward tibiofibular fusion – BUT possibly convergent evolution * Similar dentition – BUT probably not homologous – hypocones on many omomyoids Likely are convergences, unlikely that any known omomyoid is directly ancestral to Tarsius, but may not have been found yet
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Adapiformes vs Omomyiformes
Adapiformes * Avg. body size: >500g * Mostly diurnal * Dental formula: 2.1.4-3-2.3 * Spatulate incisors * Dimorphic canine * Ring-like tympanic suspended within bulla * Rel. short tarsals * Rel. elongated snout * Rel. large infraorbital foramen * ‘Higher primate’ niches (frugivory, ?folivory) Omomyiformes * Avg. body size: <500g * Mostly nocturnal * Dental formula : 2.1.4-3-2.3 * Pointed incisors * Non-dimorphic canine * Tympanic ring fused to bulla in some; tubular ectotympanic in some * Elongated tarsals * Rel. reduced snout * Small infraorbital foramen * Mostly Galago-like niches (small, mostly insectivorous)
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What is the simpliest (Parsimony) suggestion to the origin of Crown strepsirrhines?
African Origin (They passed Adapidforms) Some in Africa split from all African and Asian crown streps (this is galagoids and African Lorids)
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What does genetic data suggest about mammal dispersals to Madagascar?
There were multiple dispersal events! Likely started in the Paelocene and Early Mioene Parsomonies believed it to be a single event very early on with lemurs (compatible with data, SPLIT OCCURS ON MADAGASCAR) But there are also huge error rates Also the fossil genus Propotto suggests a second and separate migration for chiromyiformes that must have occurred later on in the fossil record Fossil record if weird on Madagascar, with little preservation of Cenzoic record (lots of assumptions and guessing)
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What is seen with the early Strepsirrhine Fossil Record/
* Leaving adapiforms aside (the most basal strepsirhines) * Have a lineage of North African fossil species that seem to represent early strepsirrhines * Generally poorly known (fieldwork ongoing) Fayum is the most important fieldwork site!
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4 Stem Strepsirrhines:
1) Azibiidae 2) Djebelemuridae 3) Chiromyiformes 4) Propotto leakeyi
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Azibiidae: Stem Strep
Early-Middle Eocene of Algeria (52- 46Ma) Two genera: Azibius – weird premolars (Not sure it was even primate) Algeripithecus minutus (Originally described as anthropoid, New material: alveoli show canines might have been procumbent: i.e. an incipient tooth-comb?) Azibids in general: – Talus is strepsirrhine-like * Laterally flared; with sloping fibular facet - Limited material
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Djebelemuridae: Stem strep
Example: Djebelemur martinezi Early-Middle Eocene Originally described as a cercamoniine adapiform Then suggested to be anthropoid More detailed analysis says stem strepsirrhine – Stem: because no tooth comb present
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Chiromyiformes: Stem Strep
Previously called Plesiopithecidae One genus and species: – Plesiopithecus teras Latest Eocene-Early Oligocene (34 Ma) Quarry L-41, Fayum Egypt Very odd primate – Dental formula ?.1.3.3/0.1.4.3 – Large upper and lower canines – No hypocones – Skull with enlarged orbits and only post-orbital bar A possible aye-aye?
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Propotto leakeyi
Early Miocene of Kenya (20-17 Ma) First described as a lorisoid, Then, as a fruitbat. Now, tied to Plesiopithecus as an aye-aye relative – Enlarged anterior dentition – Similar molar morphology Suggested lineage on the mainland Africa
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What is the now suggestion about the Strepsirrhine split in Madagascar
Split of crown streps from Lemurs and Lorises, then a split again of Lemurs and Chiromyiformes Still early Miocene (split is now EARLIER) Crown Streps has tooth comb, which derived Loris and Lemurs OR non-tooth comb ancestor and they developed them convergently Maybe 40 Ma migration rather than 50-60 Ma? Also suggestion because would not be able to get to Madagascar today from Africa due to ocean currents, maybe it was reversed at time to go quicker Had to go through adaptive radiation to survive (activity, diet, vertical space, body size)
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Crown Strepsirrhines:
1) Karanisia clarki 2) Saharagalago 3) Wadilemur 4) Miocene Fossil Lorisiforms
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Karanisia clarki
Crown Strep Latest Eocene (37 Ma) Birket Qarun Formation, Fayum Depression, Egypt Lower canine appears to have been modified into a tooth comb Originally described as crown lorisid, but now considered a basal crown strepsirrhine or stem lemuriform
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Saharagalago: Crown Strep
37 Ma (Late Eocene) Fayum Depression, Egypt Lower molars unremarkable Upper molar: * Concave distal crown; Protruding lobe for hypocone * Feature only seen in galagids Late Eocene galago, Not surprising because knew lorisids and galagids must have existed by this time
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Wadilemur: Crown Strep
34 Ma (Eocene-Oligocene Boundary) Fayum, Egypt (L-41 Quarry) Described as cercamoniine adapiform (1997) from first mandible More material shows tooth-comb (protruding p2 and alveoli for canines and incisors) Upper molars galago-like Femur shows distinct leaping adaptations similar to those in living galagids Also interpreted as a galagid
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Miocene Fossil Lorisiforms
Early Miocene (20 – 17 Ma) Uganda and Kenya, East Africa Several fossil lorisiforms appear suddenly in high-quality fossil record Three genera appear at the same time: – Mioeuoticus – lorisid – Progalago – galago – Komba - galago Similar to modern African forms (size, dentition) – Large orbits and tooth combs present Postcranial adaptations: – for leaping (like galagos) – limited evidence for slow moving like lorises (one humerus) ASIA TOO! Middle and Late Miocene, 7-14 Ma Pakistan Nycticeboides simpsoni Placed in Lorisidae Similar to living genus Nycticebus (Slow Loris) – Including post-cranial anatomy – Adapted for slow vertical climbing in trees
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Fossil Tarsiers
ASIA: Miocene – Thailand – Tarsius thailandica (1 tooth) Eocene – – Afrasia - Myanmar/Burma (isolated teeth) – Xanthorhysis – China (Lower jaw, Eocene – almost identical to modern tarsiids) – Tarsius eocaenus – China (Isolated teeth and New facial fragment AFRICA Afrotarsius chatrathi * Early Oligocene of Fayum, Egypt (Quarry M) * Some favour placing Afrotarsius within anthropoids, or as sister-taxon to anthropoids (more closely related than extant tarsiers) * Tibiofibula almost identical to Tarsius – But it’s the proximal end! Not as diagnostic. * New material from Libya now – Primitive dentition that looks tarsier-like, not anthropoid like * Affinities unclear – may be an early tarsier or anthropoid
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What happened to Eocene Primates?
Adapoids and Omomyoids in northern continents die out at end of Eocene (mostly!) – Likely due to climate change (global cooling) at end of Paleocene- Eocene Thermal Maximum Majority of primates in Northern latitudes at the end of the Eocene go extinct 35 ma Stem strepsirrhines (e.g., Djebelemur) give rise to crown strepsirrhines – Via Saharagalago, Wadilemur One of the fossil tarsiers (probably unknown) ancestral to tarsier-anthropoid clade
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Where do all nested primate groups originate?
* Euarchontoglires: LAURASIA * “Primates” ( including Plesiadapiformes): LAURASIA * Euprimates: LAURASIA * Strepsirrhines: Basal streps in Laurasiatheria, Crown Streps Africa, Lemuriform Madagascar Haplorhines: LAURASIA * Anthropoids: SUGGESTED ASIA
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What are Subfossil Lemurs?
Living lemurs are part of a radiation “ravaged by extirpation and extinction” since the arrival of humans Radiation today is not representative of the diversity that existed even a few thousand years ago Not really fossil primates, only died very recently
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What has Madagascars Climate been like?
Mostly tropical with, dry and humid forests throughout A Central Plateau LONG HISTORY OF CLIMATE INSTABILITY Lots of biodiversity due to long isolation from other species (endemism) as well as the various forms of environments
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What are Lemuirform Traits and Behaviours?
tooth combs Grooming claws on 2nd digit long snouts post-orbital bar only (no closure) unfused mandibles reproduction differences (more invasive placenta and uterus shape) * Small group sizes – Many pair bonded species * Low basal metabolic rates (BMR) * Highly seasonal breeders – 1-2 weeks/year * Varied activity patterns * Female dominance – Sexually monomorphic – Priority of access to food – Lead group movements * Adaptations to unpredictable climate – Energy conservation
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Extant Lemur Families
Lemuridae (diverse, medium size) Indriidae (largest, arboreal and clinging) Lepilemuridae (small, nocturnal) Cheriogaleidae (smallest body size) Daubentoniidae (One species, specialized diet)
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How many extinct lemurs are there?
* In the last 1000 years, 8 genera and (at least) 17 species disappeared * All lemur fossils are 15,000 and YOUNGER (Holocene) * All are subfossils (found in caves and shelters, contain some organic material, easy to radiocarbon date and get likely age, WELL CALIBRATED
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Subfossil Lemurs:
1) Palaeopropithecidae 2) Archaeolemuridae 3) Megaladapidae 4) Lemuridae (Pachylemur) 5) Daubentoniidae *refer to details of them on handout
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Sub-Fossil Lemurs compared to Living Lemurs
Similarities * No sexual dimorphism in body size or canines – Behavior similar too? * Mostly arboreal * Small brains * Strepsirrhine synapomorphies – Toothcomb – Grooming claw * Focus on olfaction * Rapid growth and development Differences: * Larger body size (all extinct forms bigger than all living) * Some appear very terrestrial * More diurnal * More folivory and seed eating * Locomotion – Slow climbing – Suspensory locomotion like sloths
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When did lemurs start going extinct after human contact
Many of these extinct species cohabitated with humans for at least 1500 years – e.g. Paleopropithecus survives until at least 1300 CE Some semi-historical records from earliest Europeans (1500 CE) * Also oral traditions – kidoky perhaps seen as recently as 1952. Some claim there are still giant lemurs
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Anthropoid Dental Features:
* 2.1.3.3 or 2.1.2.3 dental formula * Vertical incisors (not procumbent like those of streps) * Canines larger and taller than the incisors * Most have canine sexual dimorphism (size) * p3 acts to sharpen/hone the upper canine * p4 is semi-molariform with (partial) trigonid and talonid (metaconid is present) * Hypocones on upper molars (so large that cannot tell if it is a true / pseudo hypocone, so just called Hypcones) * Reduced or absent conules on upper molars * Strong lingual cingulum on upper molars * Expanded talonid basins * Molar trigonids lacking paraconid (only 2 cusps and cusps in talon basin so you cannot see the triangle easily)
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Just ANTHROPOID FEATURES
Lack of ALL grooming claws Fused frontal and mandible bones Tympanic Ring fused to lateral wall of auditory bulla Larger Brains Reduced olfaction Majority of differences are found in only dental features!
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Eosimiidaes and their time periods in the Eocene? Amphipithecidae and their time periods in the Eocene?
EOSIMIIDAE Anthrasimias gujaratensis: Early Eosimias sinensis: Middle Eosimias centennicus: Middle Phenacopithecus krishtalkai: Middle Bahinia pondaungensis: Middle AMPHIPITHECIDAE Amphipithecus mogaungensis: Late Pondaungia cotteri: Late Siamopithecus eocaenus: Late Most later periods are from Myanmar
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Anthrasimias:
55 Ma (Earliest Eocene) – Oldest possible anthropoid material Vastan Coal Mine, India Known only from isolated teeth – Small hypocone (true) – Reduced conules Teeth are highly controversial – Most now treat these as adapiform and reassign this material to Marcgodinotius (Asiadapidae) Postcrania from the site are unassociated, but some resemble eosimiids / any primitive haplorhine – So there seems to be some kind of primitive haplorrhine at the site
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Eosimias:
Middle Eocene AKA "dawn monkey from China" 40 - 45 million year old deposits in China Very small: 65-130 grams One isolated petrosal that is omomyoid-like (Not tarsier-like!) 2.1.3.3 dental formula More skull material needed to evaluate these traits * 4 species known – Eosimias sinensis – Eosimias centennicus Dental Features: Vertical and planted incisors Large, projecting canines (DISTINCT) Broad premolars with obliquely oriented roots (almost as wide as molars) (more than one cusp, talon basin on p4 and angled roots: getting bigger?) Anthropoid-like mandible – Vertical symphysis – Deep corpus – BUT unfused! Wide talonids, Cranium is unknown Triangular upper molars No Nannopithex-fold No hypocone Para- and metaconules reduced Postprotocrista reaches metacone Complete lingual cingulum Only isolated post-cranial remains Indicates Eosimias was like some platyrrhines in its postcrania Mixture of arboreal quadrupedalism and leaping
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Bahinia
Late Middle Eocene, Myanmar (SE Asia) More complete remains Cranium with orbits indicating diurnality More convincing anthropoid: – 2.1.3.3, with P2 reduced – Vertical upper incisors – Enlarged round canines – Strong lingual cingulum – Reduced conules – Small hypocone? But: – Single cusp on p4 (anthropoids have 2) – Unfused mandibular symphysis
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Amphipithecidae
ARE ANTHROPOIDS LATE EOCENE in SE Asia (Burma and Thailand) and early Oligocene Much larger than Eosimiidae (500g-10kg) 6 genera, but 3 most important Dental features * First, fragmentary remains only * Deep mandible with vertical symphysis that is almost fused * Strong P4 metaconid (two cusped premolar) (meta = two) * Bunodont (broad, low-crowned circles) molars, wear easily * Clear large hypocones on upper molars THIS CREATES A SQUARED LOOK RATHER THAN TRIANGLE * Large canines Postcrania * All isolated * Not clearly attributed to amphipithecids (or to specific species within amphipithecidae) – But likely are amphipithecid! * Some resemble anthropoids, but others are similar to adapiformes Will only learn if they look 'advanced' if there is more fossils found
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Altiatlaisus
Late Paleocene (>56 Ma), Morocco Oldest fossil euprimate known Oldest Euarchontan in Africa Little material – isolated teeth and one small mandible fragment Relationships controversial – Plesiadapid – Haplorhine – Stem euprimate – Earliest stem anthropoid Most consider this some kind of stem euprimate or an early strepsirrhine Dental features seem advanced for its early age (e.g. hypoconulid) So old cannot compare to Adapoids, most leave unassigned (FLOATER)
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What site helps determine the best fossil records in all of Primate fossils
Fayum Most important Late Eocene - Early Oligocene fossil mammal locality in Africa – First documented in 1877 – Includes the Eocene-Oligocene boundary at 34 Ma – Located in the Sahara seventy miles southwest of Cairo – Deposits uncovered by wind erosion that began in the Pliocene or Pleistocene – 5 decades of research by Elwyn Simons of Duke University MANY SITES, but Quarry BQ-2 (37 Ma) is the most important, can see the main changes occurring throughout time
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What was the Fayum like?
- Very tropical like (like SE Asian forests) - At Eocene/Oligocene barrier Streps go extinct in North Africa and Anthropids become dominant * Small-medium sized * Fruit and seed eaters, no committed folivores * Arboreal quadrupeds and leapers * Adaptively most similar to platyrrhines, and a radiation of similar diversity to platyrrhines
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Platyrrhini vs Catarrhini in Anthropoid fossils
Plats: Zygomatic-Parietal Contact (Frontal and Spinal bones aren't touching 3 premolars No ear tube Cats: Frontal-Sphenoid bones Contact 2 premolars Ear tube
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What IS an Anthropoid
* Diurnality * Postorbital closure * Advanced visual systems * Enlarged brains * Sexual dimorphism in canines * Promentory branch supplies the brain * Fused frontals and mandibles * That list of dental features - Hypocones: believed they converged separately many times, Maybed Anthropids made once and everyone gets one afterwords (Synamorphy)
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Stem vs Crown Groups
Crown: smallest group/clade that includes ALL LIVING MEMBERS of the group (can include extinct taxa) ex. All living fossils of anthropoids descend from LCA of extant plats and cats Stem: set of EXTINCT taxes that are not in the crown, but are more closely related to that crown group than to any other group or clade ex. Ancestral node of tarsiers! More closely related to crown anthropoids than they are to tarsiers BUT outside crown
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Stem vs Crown Anthropoids
Stem Anthropoids * Eosimiidae * Amphipithecidae * Parapithecoidea – Parapithecidae – Biretia * Proteopithecidae Crown Anthropoids * Propliopithecidae * Oligopithecidae