Bill Disparity and Feeding Strategies Among Fossil and Modern Penguins
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JVP 26(3) September 2006—ABSTRACTS
Neoceti Symposium, Saturday 8:45 acid-prepared osteolepiforms Medoevia and Gogonasus has offered strong support for BODY SIZE AND CRYPTIC TROPHIC SEPARATION OF GENERALIZED Jarvik’s interpretation, but Eusthenopteron itself has not been reexamined in detail. PIERCE-FEEDING CETACEANS: THE ROLE OF FEEDING DIVERSITY DUR- Uncertainty has persisted about the relationship between the large endoskeletal “fenestra ING THE RISE OF THE NEOCETI endochoanalis” and the apparently much smaller choana, and about the occlusion of upper ADAM, Peter, Univ. of California, Los Angeles, Los Angeles, CA; JETT, Kristin, Univ. of and lower jaw fangs relative to the choana. California, Davis, Davis, CA; OLSON, Joshua, Univ. of California, Los Angeles, Los A CT scan investigation of a large skull of Eusthenopteron, carried out in collaboration Angeles, CA with University of Texas and Parc de Miguasha, offers an opportunity to image and digital- Marine mammals with homodont dentition and relatively little specialization of the feeding ly “dissect” a complete three-dimensional snout region. We find that a choana is indeed apparatus are often categorized as generalist eaters of squid and fish. However, analyses of present, somewhat narrower but otherwise similar to that described by Jarvik. It does not many modern ecosystems reveal the importance of body size in determining trophic parti- receive the anterior coronoid fang, which bites mesial to the edge of the dermopalatine and tioning and diversity among predators. We established relationships between body sizes of is received by a pit in that bone. The fenestra endochoanalis is partly floored by the vomer extant cetaceans and their prey in order to infer prey size and potential trophic separation of and the dermopalatine, restricting the choana to the lateral part of the fenestra. -
PTAČÍ FYLOGENEZE Dovnitř Patří Šplhavci
E Afroaves – řada linií této skupiny je velmi rozrůzněna v Africe a pro Afriku stepokur kropenatý typická (vlhy, zoborožci, mandelíci), ale fosilie africký původ nepodporují. Mnoho (Pterocles burchelli) linií bylo řazeno do řádu srostloprstých, ale ten není monofyletický, poněvadž PTAČÍ FYLOGENEZE dovnitř patří šplhavci. Patří sem i myšáci (Coliiformes) a trogoni. F Australaves – obdobně jako u Afroaves dnes sice řada linií žije jen FYLOGENEZE NA HRUBÉ ÚROVNI A PTAČÍ ŘÁDY v australasijské oblasti, ale fosilní zástupci byli nalezeni všude možně, takže o původu skupiny těžko něco soudit. Zobrazený fylogenetický stromeček zahrnuje jen velké skupiny, přičemž co je „velká“ skupina, je docela arbitrární. Aby vůbec bylo možné dělat O Passerimorphae (nebo Psittacopasserae) – papoušci korektní fylogenetické analýzy, nesmí být těch skupin moc a zároveň je potřeba, aby to byly skupiny, o nichž se předpokládá, že jsou a pěvci sdílejí řadu znaků, například schopnost učit se zpěvu, která byla monofyletické. Reálně se totiž sekvenují samozřejmě jednotliví zástupci (tedy jeden druh, respektive přímo jeden jedinec z dané skupiny), považovaná za klíčovou pro jejich sloučení (pokládala se za znak vzniklý jen a potřebujeme tedy, aby skutečně skupinu reprezentovali. Proto by nestačilo brát zástupce tradičních ptačích řádů, které známe z učebnic, jednou u jejich společného předka, tedy synapomorfii). Ovšem vzhledem k tomu, jelikož se ukázalo, že řada z nich monofyletických není. Tradiční „dravci“ byli například polyfyletičtí, poněvadž zahrnovali monofyletickou že u pěvců je s jistotou známa jen ve skupině Oscines (zpěvní) a navíc se vyskytuje skupinu složenou z orlů, jestřábů, káňat apod. (včetně kondorů), ale taky sokolovité (poštolky, sokoly apod.), kteří jsou ve skutečnosti blízce také u zcela nezávislých kolibříků, musela vzniknout víckrát konvergentně, a to příbuzní pěvcům a papouškům. -
The Tarsometatarsus of the Middle Eocene Loon Colymbiculus Udovichenkoi
– 17 – Paleornithological Research 2013 Proceed. 8th Inter nat. Meeting Society of Avian Paleontology and Evolution Ursula B. Göhlich & Andreas Kroh (Eds) The tarsometatarsus of the Middle Eocene loon Colymbiculus udovichenkoi GERALD MAYR1, LEONID GOROBETS2 & EVGENIJ ZVONOK3 1 Senckenberg Research Institute and Natural History Museum Frankfurt, Frankfurt am Main, Germany; E-mail: [email protected] 2 Taras Shevchenko National University of Kiev, Dept. of Ecology and Environmental Protection, Kiev, Ukraine 3 Institute of Geological Sciences of NAS of Ukraine, Branch of Paleontology and Stratigraphy, Kiev, Ukraine Abstract — We describe the previously unknown tarsometatarsus of the earliest unambiguously identified loon, Colymbiculus udovichenkoi, from the Middle Eocene of the Ukraine. Except for being more elongate and apart from details of the hypotarsus morphology, the bone resembles the tarsometatarsus of the Early Miocene Colym- boides minutus. We consider the hypotarsus morphology of Colymbiculus to be plesiomorphic for Gaviiformes. Colymboides and crown group Gaviiformes are each characterized by an autapomorphic hypotarsus morphology, which precludes the former from being directly ancestral to the latter. The similarities shared by C. udovichenkoi and C. minutus, including their small size, are likely to be plesiomorphic for Gaviiformes. Although the disap- pearance of small stem group Gaviiformes may be related to the retreat of loons to cold Northern latitudes, more data are needed to firmly establish this hypothesis. We finally note that early Paleogene stem group Gaviiformes markedly differ from putative Late Cretaceous loons, whose identification needs to be verified by further fossil specimens. Key words: Colymbiculus, Colymboides, fossil birds, Gaviiformes, Eocene, Lutetian, Ukraine Introduction 1982 from the Early Miocene of the Czech Republic (ŠVEC 1982), the earliest stem line- Loons (Gaviiformes) have a fairly comprehen- age representative, being distinctly smaller than sive Neogene fossil record (OLSON 1985; MAYR its modern congeners. -
Bayesian Total-Evidence Dating Reveals the Recent Crown Radiation of Penguins Alexandra Gavryushkina University of Auckland
Ecology, Evolution and Organismal Biology Ecology, Evolution and Organismal Biology Publications 2017 Bayesian Total-Evidence Dating Reveals the Recent Crown Radiation of Penguins Alexandra Gavryushkina University of Auckland Tracy A. Heath Iowa State University, [email protected] Daniel T. Ksepka Bruce Museum David Welch University of Auckland Alexei J. Drummond University of Auckland Follow this and additional works at: http://lib.dr.iastate.edu/eeob_ag_pubs Part of the Ecology and Evolutionary Biology Commons The ompc lete bibliographic information for this item can be found at http://lib.dr.iastate.edu/ eeob_ag_pubs/207. For information on how to cite this item, please visit http://lib.dr.iastate.edu/ howtocite.html. This Article is brought to you for free and open access by the Ecology, Evolution and Organismal Biology at Iowa State University Digital Repository. It has been accepted for inclusion in Ecology, Evolution and Organismal Biology Publications by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Syst. Biol. 66(1):57–73, 2017 © The Author(s) 2016. Published by Oxford University Press, on behalf of the Society of Systematic Biologists. This is an Open Access article distributed under the terms of the Creative Commons Attribution Non-Commercial License (http://creativecommons.org/licenses/by-nc/4.0/), which permits non-commercial re-use, distribution, and reproduction in any medium, provided the original work is properly cited. For commercial re-use, please contact [email protected] DOI:10.1093/sysbio/syw060 Advance Access publication August 24, 2016 Bayesian Total-Evidence Dating Reveals the Recent Crown Radiation of Penguins , ,∗ , ALEXANDRA GAVRYUSHKINA1 2 ,TRACY A. -
1931-15701-1-LE Maquetación 1
AMEGHINIANA 50 (6) Suplemento 2013–RESÚMENES REUNIÓN DE COMUNICACIONES DE LA ASOCIACIÓN PALEONTOLÓGICA ARGENTINA 20 a 22 de Noviembre de 2013 Ciudad de Córdoba, Argentina INSTITUCIÓN ORGANIZADORA AUSPICIAN AMEGHINIANA 50 (6) Suplemento 2013–RESÚMENES COMISIÓN ORGANIZADORA Claudia Tambussi Emilio Vaccari Andrea Sterren Blanca Toro Diego Balseiro Diego Muñoz Emilia Sferco Ezequiel Montoya Facundo Meroi Federico Degrange Juan José Rustán Karen Halpern María José Salas Sandra Gordillo Santiago Druetta Sol Bayer COMITÉ CIENTÍFICO Dr. Guillermo Albanesi (CICTERRA) Dra. Viviana Barreda (MACN) Dr. Juan Luis Benedetto (CICTERRA) Dra. Noelia Carmona (UNRN) Dra. Gabriela Cisterna (UNLaR) Dr. Germán M. Gasparini (MLP) Dra. Sandra Gordillo (CICTERRA) Dr. Pedro Gutierrez (MACN) Dr. Darío Lazo (UBA) Dr. Ricardo Martinez (UNSJ) Dra. María José Salas (CICTERRA) Dr. Leonardo Salgado (UNRN) Dra. Emilia Sferco (CICTERRA) Dra. Andrea Sterren (CICTERRA) Dra. Claudia P. Tambussi (CICTERRA) Dr. Alfredo Zurita (CECOAL) AMEGHINIANA 50 (6) Suplemento 2013–RESÚMENES RESÚMENES CONFERENCIAS EL ANTROPOCENO Y LA HIPÓTESIS DE GAIA ¿NUEVOS DESAFÍOS PARA LA PALEONTOLOGÍA? S. CASADÍO1 1Universidad Nacional de Río Negro, Lobo 516, R8332AKN Roca, Río Negro, Argentina. [email protected] La hipótesis de Gaia propone que a partir de unas condiciones iniciales que hicieron posible el inicio de la vida en el planeta, fue la propia vida la que las modificó. Sin embargo, desde el inicio del Antropoceno la humanidad tiene un papel protagónico en dichas modificaciones, e.g. el aumento del CO2 en la atmósfera. Se estima que para fines de este siglo, se alcanzarían concentraciones de CO2 que el planeta no registró en los últimos 30 Ma. La información para comprender como funcionarían los sistemas terrestres con estos niveles de CO2 está contenida en los registros de períodos cálidos y en las grandes transiciones climáticas del pasado geológico. -
Skull Shape Analysis and Diet of South American Fossil Penguins (Sphenisciformes) Claudia Patricia Tambussi & Carolina Acosta Hospitaleche
Skull shape analysis and diet of South American fossil penguins (Sphenisciformes) Claudia Patricia Tambussi & Carolina Acosta Hospitaleche CONICET & Museo de La Plata, Paseo del Bosque s/n, 1900 La Plata, Argentina [email protected] [email protected] ABSTRACT – Form and function of the skull of Recent and fossil genera of available Spheniscidae are analysed in order to infer possible dietary behaviors for extinct penguins. Skull shapes were compared using the Resistant-Fit Theta-Rho- Analysis (RFTRA) Procrustean method. Due to the availability and quality of the material, this study was based on six living species belonging to five genera (Spheniscus, Eudyptula, Eudyptes, Pygoscelis, and Aptenodytes) and two Miocene species: Paraptenodytes antarticus (Moreno and Mercerat, 1891) and Madrynornis mirandus Acosta Hospitaleche, Tambussi, Donato & Cozzuol. Seventeen landmark from the skull were chosen, including homologous and geometrical points. Morphologi- cal similarities among RFTRA distances are depicted using the resulting dendrograms for UPGMA (unweighted pair-group method using arithmetic average) cluster analysis. This shape analysis allows the assessment of similarities and differences in the skulls and jaws of penguins within a more comprehensive ecomorphological and phylogenetic framework. Even though penguin diet is not well known, enough data supports the conclusion that Spheniscus + Eudyptes penguins specialize on fish and all other taxa are plankton-feeders or fish and crustacean-feeders. We compared representative species of both ecomor- phological groups with the available fossil material to evaluate their feeding strategies. Penguins are the most abundant birds, indeed the most abundant aquatic tetrapods, in Cenozoic marine sediments of South America. The results arising from this study will be of singular importance in the reconstruction of those marine ecosystems. -
Bayesian Total Evidence Dating Reveals the Recent Crown Radiation of Penguins
Supporting Information Bayesian total evidence dating reveals the recent crown radiation of penguins Contents 1 Methods 1 1.1 Prior distributions for parameters..........................1 1.2 Data..........................................2 1.3 Fossil ages.......................................2 1.4 Validation.......................................8 2 Results 8 2.1 Divergence dates...................................8 2.2 Sensitivity to the choice of prior distributions for the net diversification rate.. 11 1 Methods 1.1 Prior distributions for parameters We used the following prior distributions for the parameters in all analyses except for the sensitivity analyses. For the net diversification rate, d, we place a log-normal prior distribution with parameters1 M = −3:5 and S2 = 1:5. The 2.5% and 97.5% quantiles of this distribution are 1:6 · 10−3 and 0:57 implying that it well covers the interval (with the lowest 5% quantile of 0.02 and the largest 95% quantile of 0.15) estimated in [1] Figure 1. We place a Uniform(0,1) prior distribution for the turnover, ν, and Uniform(0,1) prior distri- bution for the fossil sampling proportion, s. For birth, death and sampling rates (λ, µ, and ) we used log-normal distributions with pa- rameters1 M = −2:5 and S2 = 1:5 and 2.5% and 97.5% quantiles of 4:34 · 10−3 and 1:55. For the time of origin, tor, we use the Uniform prior distribution between the oldest sample and 160 Ma because Jetz et al. estimated the upper bound for the MRCA of all birds to be 1Parameters M and S for log-normal distributions here are the mean and standard deviation of the associated normal distributions 1 150 Ma ([1] Figure 1) and Lee et al. -
A Rhinopristiform Sawfish (Genus Pristis) from the Middle Eocene (Lutetian) of Southern Peru and Its Regional Implications
Carnets Geol. 20 (5) E-ISSN 1634-0744 DOI 10.4267/2042/70759 A rhinopristiform sawfish (genus Pristis) from the middle Eocene (Lutetian) of southern Peru and its regional implications Alberto COLLARETA 1, 2 Luz TEJADA-MEDINA 3, 4 César CHACALTANA-BUDIEL 3, 5 Walter LANDINI 1, 6 Alí ALTAMIRANO-SIERRA 7, 8 Mario URBINA-SCHMITT 7, 9 Giovanni BIANUCCI 1, 10 Abstract: Modern sawfishes (Rhinopristiformes: Pristidae) are circumglobally distributed in warm wa- ters and are common in proximal marine and even freshwater habitats. The fossil record of modern pristid genera (i.e., Pristis and Anoxypristis) dates back to the early Eocene and is mostly represented by isolated rostral spines and oral teeth, with phosphatised rostra representing exceptional occurren- ces. Here, we report on a partial pristid rostrum, exhibiting several articulated rostral spines, from middle Eocene strata of the Paracas Formation (Yumaque Member) exposed in the southern Peruvian East Pisco Basin. This finely preserved specimen shows anatomical structures that are unlikely to leave a fossil record, e.g., the paracentral grooves that extend along the ventral surface of the rostrum. Ba- sed on the morphology of the rostral spines, this fossil sawfish is here identified as belonging to Pristis. To our knowledge, this discovery represents the geologically oldest known occurrence of Pristidae from the Pacific Coast of South America. Although the fossil record of pristids from the East Pisco Basin spans from the middle Eocene to the late Miocene, sawfishes are no longer present in the modern cool, upwelling-influenced coastal waters of southern Peru. Given the ecological preferences of the extant members of Pristis, the occurrence of this genus in the Paracas deposits suggests that middle Eocene nearshore waters in southern Peru were warmer than today. -
Bulletin~ of the American Museum of Natural History Volume 87: Article 1 New York: 1946 - X X |! |
GEORGE GAYLORD SIMPSON BULLETIN~ OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 87: ARTICLE 1 NEW YORK: 1946 - X X |! | - -s s- - - - - -- -- --| c - - - - - - - - - - - - - - - - - -- FOSSIL PENGUINS FOSSIL PENGUINS GEORGE GAYLORD SIMPSON Curator of Fossil Mammals and Birds PUBLICATIONS OF THE SCARRITT EXPEDITIONS, NUMBER 33 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY VOLUME 87: ARTICLE 1 NEW YORK: 1946 BULLETIN OF THE AMERICAN MUSEUM OF NATURAL HISTORY Volume 87, article 1, pages 1-100, text figures 1-33, tables 1-9 Issued August 8, 1946 CONTENTS INTRODUCTION . 7 A SKELETON OF Paraptenodytes antarcticus. 9 CONSPECTUS OF TERTIARY PENGUINS . 23 Patagonia. 24 Deseado Formation. 24 Patagonian Formation . 25 Seymour Island . 35 New Zealand. 39 Australia. 42 COMPARATIVE OSTEOLOGY OF MIOCENE PENGUINS . 43 Skull . 43 Vertebrae 44 Scapula. 45 Coracoid. 46 Sternum. 49 Humerus. 49 Radius and Ulna. 53 Metacarpus. 55 Phalanges . 56 The Wing as a Whole. 56 Femur. 59 Tibiotarsus. 60 Tarsometatarsus 61 NOTES ON VARIATION. 65 TAXONOMY AND PHYLOGENY OF THE SPHENISCIDAE . 68 DISTRIBUTION OF MIOCENE PENGUINS. 71 SIZE OF THE FOSSIL PENGUINS 74 THE ORIGIN OF PENGUINS. 77 Status of the Problem. 77 The Fossil Evidence . 78 Conclusions from the Fossil Evidence. 83 A General Theory of Penguin Evolution. 84 A Note on Archaeopteryx and Archaeornis 92 ADDENDUM . 96 BIBLIOGRAPHY 97 5 INTRODUCTION FEW ANIMALS have excited greater popular basis for comparison, synthesis, and gener- and scientific interest than penguins. Their alization, in spite of the fact -
71St Annual Meeting Society of Vertebrate Paleontology Paris Las Vegas Las Vegas, Nevada, USA November 2 – 5, 2011 SESSION CONCURRENT SESSION CONCURRENT
ISSN 1937-2809 online Journal of Supplement to the November 2011 Vertebrate Paleontology Vertebrate Society of Vertebrate Paleontology Society of Vertebrate 71st Annual Meeting Paleontology Society of Vertebrate Las Vegas Paris Nevada, USA Las Vegas, November 2 – 5, 2011 Program and Abstracts Society of Vertebrate Paleontology 71st Annual Meeting Program and Abstracts COMMITTEE MEETING ROOM POSTER SESSION/ CONCURRENT CONCURRENT SESSION EXHIBITS SESSION COMMITTEE MEETING ROOMS AUCTION EVENT REGISTRATION, CONCURRENT MERCHANDISE SESSION LOUNGE, EDUCATION & OUTREACH SPEAKER READY COMMITTEE MEETING POSTER SESSION ROOM ROOM SOCIETY OF VERTEBRATE PALEONTOLOGY ABSTRACTS OF PAPERS SEVENTY-FIRST ANNUAL MEETING PARIS LAS VEGAS HOTEL LAS VEGAS, NV, USA NOVEMBER 2–5, 2011 HOST COMMITTEE Stephen Rowland, Co-Chair; Aubrey Bonde, Co-Chair; Joshua Bonde; David Elliott; Lee Hall; Jerry Harris; Andrew Milner; Eric Roberts EXECUTIVE COMMITTEE Philip Currie, President; Blaire Van Valkenburgh, Past President; Catherine Forster, Vice President; Christopher Bell, Secretary; Ted Vlamis, Treasurer; Julia Clarke, Member at Large; Kristina Curry Rogers, Member at Large; Lars Werdelin, Member at Large SYMPOSIUM CONVENORS Roger B.J. Benson, Richard J. Butler, Nadia B. Fröbisch, Hans C.E. Larsson, Mark A. Loewen, Philip D. Mannion, Jim I. Mead, Eric M. Roberts, Scott D. Sampson, Eric D. Scott, Kathleen Springer PROGRAM COMMITTEE Jonathan Bloch, Co-Chair; Anjali Goswami, Co-Chair; Jason Anderson; Paul Barrett; Brian Beatty; Kerin Claeson; Kristina Curry Rogers; Ted Daeschler; David Evans; David Fox; Nadia B. Fröbisch; Christian Kammerer; Johannes Müller; Emily Rayfield; William Sanders; Bruce Shockey; Mary Silcox; Michelle Stocker; Rebecca Terry November 2011—PROGRAM AND ABSTRACTS 1 Members and Friends of the Society of Vertebrate Paleontology, The Host Committee cordially welcomes you to the 71st Annual Meeting of the Society of Vertebrate Paleontology in Las Vegas. -
Onetouch 4.0 Scanned Documents
/ Chapter 2 THE FOSSIL RECORD OF BIRDS Storrs L. Olson Department of Vertebrate Zoology National Museum of Natural History Smithsonian Institution Washington, DC. I. Introduction 80 II. Archaeopteryx 85 III. Early Cretaceous Birds 87 IV. Hesperornithiformes 89 V. Ichthyornithiformes 91 VI. Other Mesozojc Birds 92 VII. Paleognathous Birds 96 A. The Problem of the Origins of Paleognathous Birds 96 B. The Fossil Record of Paleognathous Birds 104 VIII. The "Basal" Land Bird Assemblage 107 A. Opisthocomidae 109 B. Musophagidae 109 C. Cuculidae HO D. Falconidae HI E. Sagittariidae 112 F. Accipitridae 112 G. Pandionidae 114 H. Galliformes 114 1. Family Incertae Sedis Turnicidae 119 J. Columbiformes 119 K. Psittaciforines 120 L. Family Incertae Sedis Zygodactylidae 121 IX. The "Higher" Land Bird Assemblage 122 A. Coliiformes 124 B. Coraciiformes (Including Trogonidae and Galbulae) 124 C. Strigiformes 129 D. Caprimulgiformes 132 E. Apodiformes 134 F. Family Incertae Sedis Trochilidae 135 G. Order Incertae Sedis Bucerotiformes (Including Upupae) 136 H. Piciformes 138 I. Passeriformes 139 X. The Water Bird Assemblage 141 A. Gruiformes 142 B. Family Incertae Sedis Ardeidae 165 79 Avian Biology, Vol. Vlll ISBN 0-12-249408-3 80 STORES L. OLSON C. Family Incertae Sedis Podicipedidae 168 D. Charadriiformes 169 E. Anseriformes 186 F. Ciconiiformes 188 G. Pelecaniformes 192 H. Procellariiformes 208 I. Gaviiformes 212 J. Sphenisciformes 217 XI. Conclusion 217 References 218 I. Introduction Avian paleontology has long been a poor stepsister to its mammalian counterpart, a fact that may be attributed in some measure to an insufRcien- cy of qualified workers and to the absence in birds of heterodont teeth, on which the greater proportion of the fossil record of mammals is founded. -
REPORT 2015 Vol. 6
REPORT 2015 vol. 6 Forward-thinking articles from our global network of innovation ecosystem experts KFR Staff KF Board of Directors Publisher Brian Dovey, Chairman Kauffman Fellows Press Domain Associates Executive Editor Tom Baruch Phil Wickham Formation 8 Managing Editor Jason Green Anna F.Doherty Emergence Capital Partners Associate Editor Karen Kerr Leslie F. Peters Agile Equities, LLC Production and Design Audrey MacLean Anna F. Doherty Stanford University Leslie F. Peters Susan Mason Printer Aligned Partners Almaden Press www.almadenpress.com Jenny Rooke Copyright Fidelity Biosciences © 2015 Kauffman Fellows. All rights reserved. Christian Weddell Under no circumstances shall any of the information provided herein be construed as investment advice of any kind. Copan About the Editor Phil Wickham Anna F. Doherty is an accomplished editor and writing Kauffman Fellows coach with a unique collaborative focus in her work. She has 20 years of editing experience on three continents in a variety of business industries. Through her firm, Together Editing & Design, she has offered a full suite of writing, design, and publishing services to Kauffman Fellows since 2009. Leslie F. Peters is the Lead Designer on the TE&D team. www.togetherediting.com www.kauffmanfellows.org Town & Country Village • 855 El Camino Real, Suite 12 • Palo Alto, CA 94301 Phone: 650-561-7450 • Fax: 650-561-7451 iii Kauffman Fellows on the Science of Capital Formation Phil Wickham To describe the unique contribution of Kauffman Fellows to the venture capital ecosystem, the author introduces a Startup Capital Hierarchy of Needs. While financial capital and intellectual capital are most often discussed, three other “shadow” capital types are needed for success.