GEOL431
Vertebrate Paleobiology
Spring 2025
M 1:00 - 1:50 (SPH 0302), TTH 12:30 - 1:45 (ATL1106).

Instructor: Dr. Thomas R. Holtz, Jr.
Office: Chemistry 1225B
Phone: x5-6965
E-mail: tholtz@umd.edu
Office Hours: Th 2:30-3:30

Instructor: Dr. John W. Merck, Jr.
Office: Geology 1119, Sometimes Centreville 1218
Phone: x5-4379
E-mail: jmerck@umd.edu
Office Hours: Thu 2:00 - 4:00 or by appointment. Recommend you schedule appointments in advance.

Description:

A survey of the evolution of the vertebrates, encompassing information from the diversity of living members, but concentrating on the contribution of the fossil record. Emphasis is on the phylogenetic systematics, comparative anatomy, developmental biology, and geochronology of major extinct and extant groups.

Prerequisites:

At least one of the following: BSCI207, BSCI392, GEOL104, GEOL204, BSCI333/GEOL331, HONR219D, or permission of the Geology Department.

Texts:

• Benton, Michael J. 2024. Vertebrate Palaeontology. 5th Edition. Wiley Press. 660 pp. ISBN 978-1-394-19508-4
• Additional readings from the technical literature: some listed below, others will be provided via ELMS

Emphasis:

An advanced survey of vertebrate evolution as revealed by the fossil record and of the methods by which this pattern is illuminated. This course is intended for students with serious professional or avocational interests in vertebrate evolution and some prior knowledge of:
• The history of life
• The methods of phylogenetic systematics (cladistics)
• The rock record
Our aim is to provide the necessary background knowledge in vertebrate paleobiology properly to equip a student to enter graduate studies in vertebrate paleontology and/or systematics, or related disciplines.

Course Organization:

With the assumption that the entire semester will be in-person:
• Two 75 min. lectures weekly, TTH 12:30 - 1:45, PLS 1172
• One 50 min. discussion session weekly - MON 1:00 - 1:50, PLS 1184.

Credit:

Four credit hours.

Course Website:

http://www.geol.umd.edu/~jmerck/geol431/

ELMS:

The ELMS Canvas site will include announcements concerning the class; copies of the handouts; and so forth. If you have not already done so, make sure that you get access to ELMS.

Grades:

Final grades will be based on the following items:
• TBD

Discussion: The weekly discussion will be devoted either to explorations of the technical literature for selected specific topics or to exercises designed to improve your knowledge of background information not suitable for lecture. All students are expected to attend every meeting and be an active participant when appropriate. In some classes, there may be directed interactive activities or discussions. Some of the discussions will involve creating (individually or collaboratively) anatomical drawings based on vertebrate specimens.

Homework: Four graded homework assignments throughout the course help make sure that you are familiar with some of the content and concepts.

Quizzes: There will be five quizzes. We drop the lowest score to calculate your quiz grade. These will be administered via ELMS

Mid-term exams: There will be two midterms covering material since the beginning of the semester or the previous midterm.

Final exam: The final will be administered according to the standard schedule during finals week. Although cumulative in nature, It will emphasize material since the last mid-term .

Grade calculation: With diligent work it is possible for each student to attain an A in this class. Grading will be based on points gained from the examinations listed above, as follows:

100-97% = A+	96-94% = A	93-90% = A-
89-87% = B+	86-84% = B	83-80% = B-
79-77% = C+	76-74% = C	73-70% = C-
69-67% = D+	66-64% = D	63-60% = D-
<60% = F

Class description and attendance policy: Attendance won't be taken, however attendance is required. Exams will be based on lecture and discussion material and reading assignments. A participation score will make up 15% of your discussion grade. Posted web notes are intended as a synopsis of lecture material only and are made available as current and future reference material. We recommend printing these prior to class and making marginal notations. Actual lectures may include additional information. If you miss a lecture you must get full notes from a colleague.

Learning Outcomes:

By the end of the semester, every student should be able to:
• Identify, locate, and interpret technical literature in vertebrate paleontology
• Correctly interpret major features and many details of the osteology of extinct and extant vertebrates
• Display proficient knowledge of the pattern of vertebrate evolution
• Understand major connections between vertebrate development and evolution
• Critically evaluate paleontological analyses in the technical literature.

POLICIES

It is our shared responsibility to know and abide by the University of Maryland's policies that relate to all courses, which include topics like:

• Academic integrity
• Student and instructor conduct
• Accessibility and accommodations
• Attendance and excused absences
• Grades and appeals
• Copyright and intellectual property

Please visit https://www.ugst.umd.edu/courserelatedpolicies.html for the Office of Undergraduate Studies' full list of campus-wide policies and follow up with me if you have questions.

NOTE: As part of the nature of the course, there will be a lot of memorization (less than a foreign language class, but more than that found in more mathematically-oriented introductory science classes). This will include lots of anatomical, geological, and paleontological terms, as well as evolutionary and temporal relationships. If you have difficulty memorizing, this may not be the class for you. Also, if there are words or concepts with which you are not familiar, feel free to ask Drs. Holtz or Merck (in discussion, at office hours, over email, etc.) for an explanation or clarification.

Copyright: © 2025 John W. Merck, Jr. and Thomas R. Holtz, Jr. as to this syllabus and all lectures. Students are prohibited from copying and selling course materials, from selling lecture notes, and from being paid to take lecture notes without the express written permission of the faculty teaching this course.

Part I: Introduction and Context

Date	Topic
Week one (Jan. 27-31):	1/27 Discussion: REVIEW: Geologic Time & Phylogenetic Systematics and Reading Cladograms (TH); introducing Anatomical Directions (JM). 1/28: Introduction to Vertebrata (TH) & The Fossil Record and Vertebrate Taphonomy (JM). Reading: Chap. 2 1/30 Vertebrates within Metazoa (JM). Reading: Chap. 1

Week two (Feb. 3 - Feb. 7):

2/3 Discussion: The Vertebrate Cranium I: Regions and Structures (JM) 2/4 Body-Building - Vertebrate Development (JM) 2/6 Fossil and jawless vertebrates (JM). Reading: Chap. 3, pp. 63-79 Quiz I due 2/* Homework I (Geologic Time & Phylogenetics) assigned, due 2/* 2/7 - Schedule adjustment ends.

Week three (Feb. 10-14):

2/10 Discussion:The Vertebrate Cranium II: Just Look at the Bones! (JM) 2/11 The rise of Gnathostomata and "Placodermi" (JM). Reading: Chap. 3, pp. 80-89 2/13 Eugnathostomata, Chondrichthyes, and "spiny sharks" (JM). Reading: Chap. 3, pp. 90-93 & Chap. 7, pp. 243-255

Week four (Feb. 17-21):

2/17 Discussion: Vertebrate skeletal anatomy III - The axial skeleton (JM) 2/18 The rise of Osteichthyes and basal Actinopterygii (JM). Reading: Chap. 3, pp. 94-97 & Chap. 7, pp. 255-260 2/20 Neopterygii (JM). Reading: Chap. 7, pp. 261-279 Homework I due

Week five (Feb. 24-28):

2/24 Discussion: The Appendicular Skeleton - Anatomy (JM) 2/25 Sarcopterygii toward Tetrapoda (JM). Reading: Chap. 3, pp. 98-107 & Chap. 4, pp. 115-131 2/27 Crown Tetrapoda and Amphibia (JM). Reading: Chap. 4, pp. 132-150 Quiz II due 2/*

Week six (Mar. 3 - Mar. 7):

3/3 Discussion: The Appendicular Skeleton - Development (JM) 3/4 Reptiliomorpha and Amniote origins (JM). Reading: Chap. 5, pp. 157-167 3/6 Introduction to Synapsida (TH). Reading: Chap. 5, pp. 176-188 Homework II ("Look at the Bones!") assigned. Be sure to use osteology reference and cheat-sheet! Due 3/*

Week seven (Mar. 10-14):

3/10 Discussion: The Amniote Skull (TH) 3/11 Non-mammalian Therapsida and the Rise of Mammals (TH). Reading: Chap. 11, pp. 459-474 3/13 Mesozoic Mammal Diversity; Diversification of Crown-Mammals; Marsupialia (TH). Reading: Chap. 11, pp. 475-500. Midterm Exam I due 3/*

Week eight (Mar. 24-28):

3/24 Discussion: Mammalian Odontography (TH) 3/25 Placentalia I: Origins and Atlantogenata. Reading: Chap. 11, pp. 500-511 3/27 Placentalia II: Euarchontoglires & Laurasiatheria (TH). Reading: Chap. 12, pp. 525-540, 556-574 & Chap. 13 Homework II due Homework III (Odontography) assigned, due 3/*

Week nine (Mar. 31 - Apr. 3):

3/31 Discussion:Biomechanics of terrestrial locomotion (TH) 4/1 Placentalia III: Ungulatamorpha (TH). Reading: Chap. 12, pp. 540-555 4/3 Introduction to Sauropsida (JM). Reading: Chap. 5, pp. 169-175 Quiz III due 3/* Homework III (Odontography) due

Week ten (Apr. 7-11):

4/7 Discussion: Biomechanics of aquatic locomotion (JM) 4/8 Lepidosauromorpha (JM). Reading: Chap. 9, pp. 374-382 4/10 Euryapsida (JM). Reading: Chap. 6, pp. 202-208 & Chap. 9, pp. 367-373

Week eleven (Apr. 14-18):

4/14 Discussion: The Biomechanics of Flight (JM) 4/15 Introduction to Archosauromorpha (JM). Reading: Chap. 6, pp. 209-211 4/17 Archosauriformes through Pseudosuchia (TH). Reading: Chap. 6, pp. 212-219 Midterm Exam II due 4/* Apr. 11, Last day to withdraw with a W

Week twelve (Apr. 21-25):

4/17 Discussion: Bone Histology (TH) 4/20 Introduction to Crocodylomorpha (TH). Reading: Chap. 8, pp. 353-358 4/25 Introduction to Ornithodira; Pterosauria (TH). Reading: Chap. 6, pp. 220-223 & Chap. 9, pp. 359-366

Week thirteen (Apr. 28- May 2):

4/28 Discussion: Thermal Metabolism of Fossil Vertebrates (TH) 4/27 Dinosauria Origins; Ornithischia (TH). Reading: Chap. 6, pp. 224-235 & Chap. 8, pp. 309-331 5/2 Saurischia (TH). Reading: Chap. 8, pp. 287-308 Quiz IV due 4/* Homework IV (Locomotion) assigned

Week fourteen (May 5-9):

5/5 Discussion: TBD 5/6 Pennaraptora (including Aves) (TH). Reading: Chap. 10 5/8 "God's noblest creation"; Let's Go Terps! (JM). Reading: Chap. 9, pp. 343-352 Homework IV due

Week fifteen (May 12-13):

5/12 Discussion: Vertebrates through time (TH) 5/13 Outstanding Problems in Vertebrate Paleontology (JM) Quiz V due 5/13

Finals week:

May *** Final Exam

There is no perfect textbook for GEOL431, although Benton 2024 provides a good background to the information that vertebrate paleontologists work with. Serious students of vertebrate paleontology, however, should be aware of the landmark textbooks in the field, past and present. Before you start grad school, definitely acquaint yourself with the following landmarks:

• The inspiration of three generations of paleontologists from an age where there could be an undisputed master:
  Alfred S. Romer, 1966. Vertebrate Paleontology. University of Chicago Press. 468 pp.

• The ultimate resource for illustrations and morphology of fossil vertebrates:
  Robert L. Carroll. 1990. Vertebrate Paleontology and Evolution. W. H. Freeman and Company. 698 pp.

• A function-and-developmental biology oriented vertebrate paleontology source:
  Matthew F. Bonnan. 2016. The Bare Bones: An Unconventional Evolutionary History of the Skeleton. Indiana University Press. 508 pp.

• The grand resource of the future - eagerly awaited:
  Darren Naish. 20??. Working Title: The Vertebrate Fossil Record.

Additional reading on specific topics is cited in many lecture notes.

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PDF version of the Syllabus and Lecture schedule available on ELMS Latest revision: 10 January 2025