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 Atlas of Advanced
Shoulder Arthroscopy
 Atlas of Advanced
Shoulder Arthroscopy

edited by
Andreas B. Imhoff
Department of Orthopedic Sports Medicine
Technical University of Munich
Munich, Germany

Jonathan B. Ticker
College of Physicians and Surgeons of Columbia University
New York, New York
Orlin & Cohen Orthopedic Group
Merrick, New York

Augustus D. Mazzocca
Department of Orthopedic Surgery
University of Connecticut Health Center
Farmington, Connecticut

Andreas Voss
Department of Orthopedic Sports Medicine
Technical University of Munich
Munich, Germany
CRC Press
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Library of Congress Cataloging‑in‑Publication Data

Names: Imhoff, Andreas B., editor. | Ticker, Jonathan B., editor. | Mazzocca,
Augustus D., editor. | Voss, Andreas, M.D., editor.
Title: Atlas of advanced shoulder arthroscopy / Andreas B. Imhoff, Jonathan
B. Ticker, Augustus Mazzocca, Andreas Voss.
Description: Boca Raton : CRC Press, [2018] | Includes bibliographical
references and index.
Identifiers: LCCN 2017027654| ISBN 9781498787482 (hardback : alk. paper) |
ISBN 9781315148687 (eBook - General) | ISBN 9781351372763 (ebook - pdf) |
ISBN 9781351372756 (ebook - epub) | ISBN 9781351372749 (ebook - mobipocket)
Subjects: | MESH: Shoulder Joint--surgery | Arthroscopy | Atlases
Classification: LCC RD686 | NLM WE 17 | DDC 617.4/720597--dc23
LC record available at https://lccn.loc.gov/2017027654

Visit the Taylor & Francis Web site at

http://www.taylorandfrancis.com
and the CRC Press Web site at
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To my lovely and beautiful wife, Susann, for her love and continuous support, to our son Florian and his
wife Mareen, with our grandson Louis, and to my sons Pascal and Dominik with his wife Dominique for
encouragement and accepting the lifestyle of an orthopaedic surgeon. To my mother, Marianne, and in
memory of my father, Urs, for their love. Finally, to all my fellows in the last 30 years for their support
and questions to help me in treating patients better.

Andreas B. Imhoff

To Alyse.

Jonathan B. Ticker

To my wonderful and beautiful wife, Jennifer, who holds our whole family together and is dedicated to
making the world a better place. My wonderful children, Gus, Nico, and Jillian, who have given me my
greatest joy to see them work hard and become successful adults. Finally, my parents, Gus and D’Ann,
who taught me that hard work and honesty will always succeed.

Augustus D. Mazzocca

To my wonderful wife, Maria, and my parents, Marita and Jörg, for their continuous support.

Andreas Voss
Contents

Preface xi
Editors xiii
Contributors xvii

Section I  BASICS OF SHOULDER ARTHROSCOPY

1 Anatomy and biomechanics of the shoulder 3

Lucca Lacheta and Bastian Scheiderer
2 Physical examination of the shoulder 15
Florian B. Imhoff and Andreas Voss
3 Imaging of the shoulder 23
Pia M. Jungmann

Section II  GLENOHUMERAL INSTABILITY AND BONY DEFECTS

4 Arthroscopic anterior instability repair 37

Hardeep Singh, Andreas Voss, and Robert A. Arciero
5 Anterior instability repair (5:30 portal) 45
Knut Beitzel and Frauke Wilken
6 Anterior glenoid bone graft 51
Marvin Minkus, Elisabeth Böhm, and Markus Scheibel
7 Posterior shoulder instability repair: Indications and arthroscopic technique 59
Fotios P. Tjoumakaris, Weilong J. Shi, and James P. Bradley
8 Posterior glenoid bone graft 67
Matthew T. Provencher, Anthony Sanchez, E. Stefan Vutescu, and George Sanchez
9 McLaughlin/Neer procedure of subscapularis transfer into a reverse Hill–Sachs lesion 75
Felipe Eggers, Sepp Braun, Andreas Voss, and Andreas B. Imhoff
10 Remplissage technique for Hill–Sachs lesion 79
Jiwu Chen
11 Anterior glenoid bone loss: Japan 83
Eiji Itoi, Nobuyuki Yamamoto, Taku Hatta, and Jun Kawakami
12 The arthroscopic Latarjet 91
Johannes E. Plath and Laurent Lafosse
13 Latarjet (congruent arc technique) 99
Andrew J. Sheean, Robert U. Hartzler, and Stephen S. Burkhart
14 Arthroscopic HAGL and RHAGL repair 109
Stephen Thon, Felix H. Savoie III, and Michael J. O’Brien
15 Arthroscopic repair of Type IIa and IIb SLAP lesions 117
Kevin P. Shea
16 Pearls and pitfalls in stabilizing surgery 123
S. Tal Hendrix and John M. Tokish

Section III  ROTATOR CUFF PATHOLOGY

17 Single-row rotator cuff repair 133

Carl K. Schillhammer and Dan Guttmann

vii
viii Contents

18 Knotless, double row, transosseous equivalent rotator cuff repair (USA) 141
Brandon J. Erickson and Anthony A. Romeo
19 Double row cuff repair (Europe) 149
Emilio Calvo and Gonzalo Samitier
20 PASTA repair 157
Bastian Scheiderer and Andreas B. Imhoff
21 Arthroscopic repair of the subscapularis 163
Brett A. Lenart and Jonathan B. Ticker
22 Arthroscopic transosseous rotator cuff repair 177
Ofer Levy and Ehud Atoun
23 Arthroscopic superior capsular reconstruction 183
Matthew P. Noyes, Patrick J. Denard, and Stephen S. Burkhart
24 Biological advancements in rotator cuff repair 189
Felix Dyrna, Mary-Beth McCarthy, Megan Wolf, Jennifer Brewer, Andreas Voss, Andreas B. Imhoff, and Augustus D. Mazzocca
25 Arthroscopic revision of failed rotator cuff reconstruction 197
Eduard Buess and Michael Hackl
26 Capsular release 207
John G. Horneff III and Mark D. Lazarus
27 Value of the arthroscopic rotator cuff repair 215
Catherine J. Fedorka and Laurence D. Higgins

Section IV PROXIMAL BICEPS PATHOLOGY

28 Arthroscopic biceps tenodesis (intra-articular—high in the groove) 223

Paul Brady
29 Suprapectoral biceps tenodesis 229
Paul Sethi and WM Grant Franco
30 Arthroscopic suprapectoral biceps tenodesis with dual expansion PEEK anchor and bone socket fixation 237
Joseph C. Tauro and Jonathan B. Ticker
31 Open subpectoral biceps tenodesis: Interference screw fixation 245
Jeremiah D. Johnson, Felix Dyrna, and Augustus D. Mazzocca
32 Tenodesis of the long head of the biceps tendon (subpectoral) 251
Felipe Eggers, Andreas Voss, and Sepp Braun

Section V FRACTURES AND CHONDRAL DEFECTS

33 Treatment of glenohumeral chondral defects 257

Timothy S. Leroux, Michael L. Redondo, and Brian J. Cole
34 Proximal humeral fracture repair (greater tuberosity) 265
J. Christoph Katthagen, Alexander Ellwein, and Helmut Lill
35 Glenoid fracture repair using the 5:30-o’clock portal 273
Andreas Voss, Knut Beitzel, and Andreas B. Imhoff
36 Anatomical humeral head resurfacing 279
Anthony Miniaci, Ashraf M. Elbanna, and Ronak M. Patel
37 Shoulder arthroplasty 285
Werner Anderl, Brenda Laky, and Philipp R. Heuberer

Section VI  AC AND SC JOINT

38 Operative management of acromioclavicular joint injuries 295

Felix Dyrna, Brendan Comer, and Augustus D. Mazzocca
39 Acromioclavicular stabilization 307
Frauke Wilken, Knut Beitzel, Andreas B. Imhoff, and Sepp Braun
40 Arthroscopic excision of the sternoclavicular joint 313
Frank Martetschläger and Graham Tytherleigh-Strong
 Contents ix

Section VII  SPECIALIZED ARTHROSCOPIC TECHNIQUES

41 Arthroscopic inferior transverse scapular ligament release at the spinoglenoid notch and ganglion cyst
decompression using the extra-articular Plancher portal 321
Stephanie C. Petterson, Joseph M. Ajdinovich, and Kevin D. Plancher
42 Arthroscopic superior transverse scapular ligament release for suprascapular nerve compression at the
suprascapular notch 333
Michael J. Messina, Marie Walcott, Michelle J. Chang, and Jon J.P. Warner
43 Arthroscopic resection of the superomedial scapula and scapulothoracic bursectomy 339
Robert Z. Tashjian
44 Arthroscopic treatment of symptomatic pre- and meso-acromion 349
Brandon J. Erickson and Scott W. Trenhaile
45 The comprehensive arthroscopic management (CAM) procedure for the treatment of glenohumeral osteoarthritis
in young patients 355
Justin J. Mitchell, J. Christoph Katthagen, Salvatore J. Frangiamore, Sandeep Mannava, and Peter J. Millett

Index 363
Preface
After the success of our 1st and 2nd editions, focusing on
the basics of shoulder surgery and shoulder arthroscopy,
we are pleased to introduce the 3rd edition. The aim of
this book is to create a bridge between basic and advanced
operative procedures in shoulder arthroscopy. Therefore,
we are delighted that numerous specialized shoulder sur-
geons from all over the world have contributed to this
edition. Their insight in special procedures will help the
reader to understand and to treat difficult pathologies of
the shoulder. We have also aimed to make allowance for
the diversity of operative procedures for treating the same
pathology. Therefore, we have provided alternative opera-
tive techniques for identical or similar pathologies. We
hope that through this variety the reader will find the one
option which fits best for him or her, as well as the patient
and surgical setup.
Each chapter is structured to show: indication; opera-
tion principles; preoperative assessment, with clinical
examination, patient information, and consent; relevant
imaging; positioning and preparation; operative tech-
nique; and postoperative management, as well as follow-
up treatment.
A big thank you to our families, partners, and children,
who have supported our activities and duties much more
than normal.
We hope that those who laid their hands on the 1st edi-
tion 20 years ago, as well as those who show their curiosity
for the first time, will read the chapters and study the illus-
trations with great interest, so expanding their knowledge
and surgical horizons.
Andreas B. Imhoff
Jonathan B. Ticker
Augustus D. Mazzocca
Andreas Voss
xi
Editors
Professor Andreas B. Imhoff, MD,
graduated from Basel University in
1980. He started his residency in
General Surgery and Traumatology
in Davos, Switzerland, under
the guidance of Professor Peter
Matter, and in General Surgery,
Traumatology, and Knee Surgery
at the University Hospital
Bruderholz-Basel under the guid-
ance of Professor Werner Müller
and Professor Peter Waibel in 1982. He completed his
Orthopedic Residency at the Balgrist University Hospital,
University of Zurich, under the direction of Professor Adam
Schreiber, Professor Hans Zollinger, and Professor Christian
Gerber between 1983 and 1996, after 1987 as head of differ-
ent departments for knee, shoulder, and sports medicine. A
one-year exchange fellowship brought him to the depart-
ment of orthopedic surgery in the Center of Sports Medicine
at the University of Pittsburgh, PA, USA, under the direc-
tion of Professor Freddie Fu, Professor Jon J.P. Warner, and
Professor Chris Harner, but also at the Musculoskeletal
Research Center, University of Pittsburgh under the direc-
tion of Professor Savio L.-Y. Woo between 1994 and 1995.
He was elected as a Professor of Orthopedic Surgery
and Arthroscopy at the faculty Medical School and faculty
School of Sport Science and Health, Technical University
Munich, to serve as director and chairman of the
Department of Orthopedic Sports Medicine from 1996.
Professor Imhoff is an active member of the AGA
(German Society of Arthroscopy and Joint Surgery) which
currently has 4500 members. He was a founder in 1983,
congress chairman 1999, president 2000–2003, general
secretary, and is now still a member of the executive board.
He is again congress chairman in 2017.
He has been on the board of directors of the DGOOC
(German Society of Orthopaedics), DGOU (German Society
of Orthopaedics and Traumatology) since 1996. He is also
member of ISAKOS (International Society of Arthroscopy,
Knee Surgery, and Orthopedic Sports Medicine), started as
a founding member of the IAA/ISK in October 1994, and
has been a member of the board and different commit-
tees since 1996. He has also been chairman of the ISAKOS
shoulder committee since 2015. He was awarded as
Corresponding member by the ASES (American Shoulder
and Elbow Society) in 2004, where he is actually working
in the program committee and the research committee. He
is a member of several societies, including the SGO (Swiss
Society for Orthopaedics), ESSKA (European Society of
Sports Traumatology, Knee Surgery, and Arthroscopy),
SECEC (European Society for Surgery of the Shoulder and
the Elbow), AOSSM (American Orthopedic Society for
Sports Medicine), DVSE (German Society for Shoulder and
Elbow), DGU (German Society for Traumatology), SICOT
(Société Internationale de Chirurgie Orthopédique et de
Traumatologie), and has been a member of the board of
AFOR (Association of Orthopedic Research) since 2010,
and of DKG (German Knee Society) since 2012.
Professor Imhoff has received many awards from dif-
ferent countries all over the world since 1999, and he has
edited 34 books and published over 365 journal articles,
cited in Pubmed. He was awarded and listed by the FOCUS
company every year since 2010. As one of the Top Surgeons
in Shoulder, Knee and Sportsorthopaedics in Germany.
He has been an honorary member of the AANA
(Arthroscopy Association of North America) since
2004, Miembro Honorario of the Sociedad Argentina de
Cirurgia de Hombro y Codo since 2006, he received the
Malaysian Federal Honorary Award in 2010, has been an
honorary member of AGA since 2013, and an honorary
member of IAS (Indian Arthroscopic Society) since 2014.
Professor Imhoff serves as an editor of journals
including Arthroscopy, Operative Orthopaedics and
Traumatology, Journal of Shoulder and Elbow Surgery,
Knee Surgery, Sports Traumatology, Arthroscopy (KSSTA),
American Journal of Sports Medicine (AJSM) and the Open
Access Journal of Sports Medicine as Editor-in-Chief.
Jonathan B. Ticker, MD, is an
Assistant Clinical Professor
of Orthopaedic Surgery at
the College of Physicians and
Surgeons of Columbia University,
New York. Dr. Ticker is Chair
of the Board of Trustees of the
American Shoulder & Elbow
Surgeons Foundation, and is a
past member of the Executive
Committee of ASES. He has been
Co-Chair of the Industry and
xiii
xiv Editors
Program Committees for ASES, as well as Chair of the
Research Committee for the Arthroscopy Association of
North America. His academic endeavors have included
research on the shoulder in basic science and clinical
subjects. These interests have resulted in numerous peer-
reviewed journal publications, as well as book chapters.
In addition, Dr. Ticker has co-edited two shoulder texts:
An Atlas of Shoulder Surgery and An Atlas of Shoulder
Arthroscopy, along with Freddie Fu and Andreas Imhoff.
He edited American Shoulder & Elbow Surgeons 25th
Anniversary, a book devoted to the history of this Society.
His emphasis on education includes instructing at interna-
tional, national, regional, and local meetings and courses,
with didactic and technique-oriented efforts, as well as
orthopedic residency training at Northwell Health’s Long
Island Jewish Medical Center. Dr. Ticker is in private
practice on Long Island, New York, with Orlin & Cohen
Orthopaedic Group. His clinical and operative practice
is exclusive to the shoulder (www.LIshoulder.com). He
has been married for over 30 years, with two wonderful
children.
Professor Augustus D. Mazzocca,
MS, MD, is the Director of the
UConn Musculoskeletal Institute
and Chairman, Department
of Orthopedic Surgery, at the
University of Connecticut Health
Center. He is the Director of the
University of Connecticut Human
Soft Tissue Research Laboratory,
which consists of integrated transla-
tional laboratories incorporating cell and molecular biology,
histology, biomechanics, and clinical outcomes research.
He is also the Director of the University of Connecticut
Bioskills Laboratory and holds the Harry and Helen Gray,
Harry R. Gossling, MD Chair in Orthopaedic Surgery.
Professor Mazzocca also holds a joint faculty appointment
at the University of Hartford in the Department of Civil,
Environmental, and Biomedical Engineering, College of
Engineering Technology and Architecture.
International collaboration in both education and
research is a top priority for Professor Mazzocca bridg-
ing six countries, including Brazil, Japan, Austria,
Germany, Italy, and France and five of the seven conti-
nents. He has had several international sports medicine
research fellows in his laboratories, and he is internation-
ally renowned for his work in the following areas: biceps
tenodesis, distal biceps for the elbow, anatomic coraco-
clavicular reconstruction for the treatment of chronic
acromioclavicular separation, and biologic augmentation
of failed rotator cuff repair using concentrated bone mar-
row and platelet rich plasma. The extent of this research
has led to 70 book chapters, 138 abstracts, posters, 162
peer reviewed journal articles, and 252 invited national
and international talks.
Professor Mazzocca has also held many important posi-
tions. He was the Program Director for the American
Orthopaedic Society for Sports Medicine (AOSSM) for the
2015 International Meeting, and a member at large for the
AOSSM Nominating Committee from 2014–2015. In 2014
he served on the Upper Extremity Program Committee for
Specialty Day. Professor Mazzocca has been a part of the
American Shoulder and Elbow Society (ASES) Continuing
Education Committee since 2009, and has been a member
of the Closed Meeting Committee for 2015 and 2016. He
has also been a member of the Arthroscopy Association of
North America (AANA) Research Committee since 2010.
In 2003, Professor Mazzocca was a founder of the New
England Shoulder and Elbow Society (NESES) and contin-
ues to be part of its executive governing board. He remains
an active member of AOSSM, ASES, AANA, and NESES,
as well as the following professional societies: American
Academy of Orthopaedic Surgeons (AAOS), International
Society of Arthroscopy, Knee Surgery and Orthopaedic
Sports Medicine (ISAKOS), The American Orthopaedic
Association (AOA), Orthopaedic Research Society (ORS),
European Society for Surgery of the Shoulder and Elbow
(ESSE), American College of Sports Medicine Member
(ACSM), and the Connecticut Academy of Science and
Engineering (CASE).
Professor Mazzocca has served on the editorial board
of several orthopaedic publications including: Orthopedics
Today, Basic Science & Technology Section Editor 2014,
Orthopedics Today Editorial Board from 2013 to present,
Techniques in Shoulder and Elbow Surgery, Editorial Board
from 2010 to present, Associate Editor of the Journal of
Bone and Joint Surgery–Shoulder and Elbow Newsletter
from 2011 to present, Section Editor–Arthroscopy Section
for the AAOS Orthopaedic Knowledge Update 4th Edition
in 2011, co-editor of the AAOS Monograph Disorders of
the Proximal Biceps Tendon in 2011. He has also received
more than 50 awards, honors, and grants.
Andreas Voss, MD, is a resident
in training to become an ortho-
paedic and trauma surgeon, and
also works at the Department
of Orthopedic Sports Medicine
at the Hospital Rechts der Isar
of the Technical University of
Munich. During his training at
the Department of Orthopedic
Sports Medicine he spent one
year as a Sports Medicine
Research Fellow at the University of Connecticut
(Farmington, CT, USA), focusing on shoulder research
and surgery. In addition to receiving his medical degree
from the University of Regensburg (Germany), he also
graduated from the SRH University of Riedlingen as a
health economist, specializing in hospital management.
He is a member of the following professional asso-
ciations: Association for Arthroscopy and Joint
Surgery (AGA), German Association for Shoulder and
Elbow Surgery (DVSE), European Society of Sports
Traumatology, Knee Surgery, and Arthroscopy (ESSKA),

and the International Society of Arthroscopy, Knee
Surgery, and Orthopaedic Sports Medicine (ISAKOS).
Within the ISAKOS he works as a guest member in the
shoulder committee.
Besides his enthusiasm for and interest in research, he
also greatly supports medical education. Therefore, he
was one of the founding members of a special student’s
board within the AGA. Using a unique step-by-step
Editors xv
curriculum, the medical students are taught in the
basic principles of joint surgery to spark their inter-
est in orthopedic surgery. He also holds the certificate
of higher ­education of Bavaria, and has received sev-
eral grants and awards (e.g., co.don Research Award,
German Research Foundation grant, German Society
of Orthopedics and Traumatology grant, Bavarian
Research Alliance grant).
Contributors

Joseph M. Ajdinovich and

Plancher Orthopaedics and Sports Medicine
Department of Orthopaedic Sports Medicine
New York, New York
Technical University of Munich
and Munich, Germany

Orthopaedic Foundation Jennifer Brewer

Stamford, Connecticut Department of Orthopaedic Surgery
University of Connecticut
Werner Anderl Farmington, Connecticut
St. Vincent Shoulder and Sports Clinic
and Eduard Buess
Austrian Research Group for Regenerative and Orthopedic Orthopaedic Practice “Shouldercare”
Medicine Bern, Switzerland
Vienna, Austria Stephen S. Burkhart
Robert A. Arciero The San Antonio Orthopaedic Group
Department of Orthopaedic Surgery Burkhart Research Institute for Orthopaedics (BRIO)
University of Connecticut Health Center San Antonio, Texas
Farmington, Connecticut Emilio Calvo
Department of Orthopaedic Surgery and
Ehud Atoun
Traumatology
Ben-Gurion University
Universidad Autónoma
Barzilai Medical Center
Madrid, Spain
Ashkelon, Israel
Michelle J. Chang
Elisabeth Böhm
Department of Orthopedics
Department of Shoulder and Elbow Surgery
Boston Shoulder Institute
Charité-Universitaetsmedizin Berlin
Massachusetts General Hospital
Berlin, Germany
Boston, Massachusetts
Knut Beitzel
Department of Orthopaedic Sports Medicine Jiwu Chen
Technical University of Munich Huashan Hospital, Fudan University
Munich, Germany Shanghai, China
Brian J. Cole
James P. Bradley
Department of Orthopedics
Clinical Professor of Orthopaedic Surgery
and
University of Pittsburgh Medical Center
Department of Surgery
Burke and Bradley Orthopaedics
Rush OPHShoulder, Elbow and Knee Surgery
Pittsburgh, Pennsylvania
and
Paul Brady Cartilage Restoration Center at Rush
Tennessee Orthopaedic Clinics Rush University Medical Center
Knoxville, Tennessee Chicago, Illinois
Sepp Braun Brendan Comer
Gelenkpunkt Department of Orthopaedic Surgery
Sports and Joint Surgery Innsbruck University of Connecticut Health Center
Innsbruck, Austria Farmington, Connecticut

xvii
xviii Contributors

Patrick J. Denard S. Tal Hendrix

Southern Oregon Orthopedics Steadman Hawkins Clinic of the Carolinas
Medford, Oregon Greenville, South Carolina
Felix Dyrna Philipp R. Heuberer
Department of Orthopaedic Sports Medicine Austrian Research Group for Regenerative and Orthopedic
Technical University of Munich Medicine
Munich, Germany and
Felipe Eggers St. Vincent Shoulder and Sports Clinic
Department of Orthopaedic Sports Medicine Vienna, Austria
Technical University of Munich Laurence D. Higgins
Munich, Germany Boston Shoulder Institute, Brigham and Women’s Hospital,
Harvard Medical School
Ashraf M. Elbanna
Boston, Massachusetts
Orthopaedic Surgeon
Memorial Healthcare John G. Horneff III
Owosso, Michigan Department of Orthopaedic Surgery
Thomas Jefferson University
Alexander Ellwein
Philadelphia, Pennsylvania
Department of Trauma and Orthopaedic Surgery
Diakovere Hospital Friederikenstift Andreas B. Imhoff
Hannover, Germany Department of Orthopedic Sports Medicine
Technical University of Munich
Brandon J. Erickson
Munich, Germany
Department of Shoulder and Sports Medicine
Hospital for Special Surgery Florian B. Imhoff
New York City, New York Department of Orthopaedic Sports Medicine
Technical University of Munich
Catherine J. Fedorka
Munich, Germany
Cooper Bone and Joint Institute
Cooper University Hospital Eiji Itoi
Camden, New Jersey Department of Orthopaedic Surgery
Tohoku University School of Medicine
Wm Grant Franco
Sendai, Japan
ONS Foundation for Clinical Research and
Education Jeremiah D. Johnson
Greenwich, Connecticut Department of Orthopaedic Surgery
University of Connecticut Health Center
Salvatore J. Frangiamore
Farmington, Connecticut
The Steadman Clinic and Steadman Philippon Research
Institute Pia M. Jungmann
Vail, Colorado Department of Radiology
Technical University of Munich
Dan Guttmann Munich, Germany
Director of Shoulder and Elbow Surgery
Taos Orthopaedic Institute Sports Medicine Fellowship and and
Research Foundation Department of Radiology
Taos, New Mexico Orthopedic University Hospital Balgrist
Michael Hackl Zurich, Switzerland
Klinik und Poliklinik für Orthopädie und Unfallchirurgie, J. Christoph Katthagen
Universitätsklinikum Department of Trauma, Hand and Reconstructive Surgery
Köln, Germany University Hospital Münster
Robert U. Hartzler Münster, Germany
The San Antonio Orthopaedic Group Jun Kawakami
Burkhart Research Institute for Orthopaedics Department of Orthopaedic Surgery
(BRIO) Tohoku University School of Medicine
San Antonio, Texas Sendai, Japan
Taku Hatta Lucca Lacheta
Department of Orthopaedic Surgery Department of Orthopaedic Sports Medicine
Tohoku University School of Medicine Technical University of Munich
Sendai, Japan Munich, Germany
 Contributors xix

Laurent Lafosse Anthony Miniaci

Alps Surgery Institute Cleveland Clinic Sports Health Center
Clinique Générale Garfield Heights, Ohio
Annecy, France
Marvin Minkus
Brenda Laky Department of Shoulder and Elbow Surgery
Austrian Research Group for Regenerative and Orthopedic Charité-Universitaetsmedizin Berlin
Medicine Berlin, Germany
and
Justin J. Mitchell
St. Vincent Shoulder and Sports Clinic
Gundersen Health System Department of Sports Medicine
Vienna, Austria
La Crosse, Wisconsin
Mark D. Lazarus
Matthew P. Noyes
Department of Orthopaedic Surgery
Southern Oregon Orthopedics
Thomas Jefferson University
Medford, Oregon
Philadelphia, Pennsylvania
Michael J. O’Brien
Brett A. Lenart
Department of Orthopedics
Orlin & Cohen Orthopaedic Group
Tulane University
Merrick, New York
New Orleans, Louisiana
Timothy S. Leroux
Ronak M. Patel
Department of Orthopaedic Surgery
Hinsdale Orthopaedic Associates
University of Toronto
Hinsdale, Illinois
and
Toronto Western Hospital Stephanie C. Petterson
University Health Network Orthopaedic Foundation
Toronto, Ontario, Canada Stamford, Connecticut

Ofer Levy Kevin D. Plancher

Reading Shoulder Unit Orthopaedic Foundation
Royal Berkshire Hospital and Berkshire Independent Hospital Stamford, Connecticut
Reading, UK and
Helmut Lill Plancher Orthopaedics and Sports Medicine
Department of Trauma and Orthopaedic Surgery New York, New York
Diakovere Hospital Friederikenstift
Hannover, Germany Johannes E. Plath
Alps Surgery Institute
Sandeep Mannava Clinique Générale
The Steadman Clinic and Steadman Philippon Research Institute Annecy, France
Vail, Colorado
Matthew T. Provencher
Frank Martetschläger The Steadman Clinic and Steadman Philippon Research Institute
Department of Orthopaedic Sports Medicine Vail, Colorado
Technical University of Munich
Michael L. Redondo
Munich, Germany
Department of Orthopaedic Surgery
Augustus D. Mazzocca Rush University Medical Center
Department of Orthopedic Surgery Chicago, Illinois
University of Connecticut Health Center
Anthony A. Romeo
Farmington, Connecticut
Department of Shoulder and Elbow Surgery
Mary-Beth McCarthy Midwest Orthopaedics at Rush
Department of Orthopaedic Surgery Rush University Medical Center
University of Connecticut Chicago, Illinois
Farmington, Connecticut
Gonzalo Samitier
Michael J. Messina Department of Orthopaedic Surgery and Traumatology
Valley Forge Orthopaedics Hospital General de Villalba
Paoli, Pennsylvania Madrid, Spain
Peter J. Millett Anthony Sanchez
The Steadman Clinic and Steadman Philippon Research Institute Steadman Philippon Research Institute
Vail, Colorado Vail, Colorado
xx Contributors

George Sanchez Jonathan B. Ticker

Geisel School of Medicine at Dartmouth College of Physicians and Surgeons of Columbia University
Hanover, New Hampshire New York, New York
Felix H. Savoie III and
Department of Orthopedics
Orlin & Cohen Orthopaedic Group
Tulane University
Merrick, New York
New Orleans, Louisiana
Fotios P. Tjoumakaris
Markus Scheibel
Sidney Kimmel College of Medicine, Jefferson Medical College
Department of Shoulder and Elbow Surgery
Rothman Institute Orthopaedics
Charité-Universitaetsmedizin Berlin
Egg Harbor Township, New Jersey
Berlin, Germany
Bastian Scheiderer John M. Tokish
Department of Orthopaedic Sports Medicine Steadman Hawkins Clinic of the Carolinas
Technical University of Munich Greenville, South Carolina
Munich, Germany and
Carl K. Schillhammer Mayo Clinic Arizona
Pintler Surgical Specialists Phoenix, Arizona
Community Hospital of Anaconda
Anaconda, Montana Scott W. Trenhaile
Orthoillinois
Paul Sethi Rockford, Illinois
ONS Foundation for Clinical Research and Education
Greenwich, Connecticut Graham Tytherleigh-Strong
Department of Orthopaedic Sports Medicine
and Technical University of Munich
Clinical Instructor in Orthopedic Surgery Munich, Germany
Yale University School of Medicine Andreas Voss
New Haven, Connecticut Department of Orthopedic Sports Medicine
Kevin P. Shea Technical University of Munich
Division of Sports Medicine and Shoulder Surgery Munich, Germany
University of Connecticut Health Center E. Stefan Vutescu
Farmington, Connecticut Rhode Island Hospital
Andrew J. Sheean Department of Orthopaedic Surgery
Department of Orthopaedic Surgery Brown University
San Antonio Military Medical Center Providence, Rhode Island
San Antonio, Texas
Marie Walcott
Weilong J. Shi Agility Orthopedics
Sidney Kimmel College of Medicine, Jefferson Medical College Stoneham, Massachusetts
Rothman Institute Orthopaedics Jon J.P. Warner
Philadelphia, Pennsylvania Department of Orthopedics
Hardeep Singh Boston Shoulder Institute
Department of Orthopaedic Surgery Massachusetts General Hospital
University of Connecticut Health Center Boston, Massachusetts
Farmington, Connecticut Frauke Wilken
Robert Z. Tashjian Department of Orthopaedic Sports Medicine
University of Utah Orthopaedic Center Technical University of Munich
Salt Lake City, Utah Munich, Germany
Joseph C. Tauro Megan Wolf
Rutgers Medical School Department of Orthopaedic Surgery
Ocean County Sports Medicine University of Connecticut
Toms River, New Jersey Farmington, Connecticut
Stephen Thon Nobuyuki Yamamoto
Department of Orthopedic Surgery Department of Orthopaedic Surgery
Tulane University Tohoku University School of Medicine
New Orleans, Louisiana Sendai, Japan
 Section    I
Basics of Shoulder Arthroscopy
 1
Anatomy and biomechanics of the shoulder

LUCCA LACHETA and BASTIAN SCHEIDERER

CONTENTS
Introduction 3
Landmarks 4
Acromioclavicular joint 4
Acromion 4
Coracoid process 4
Glenohumeral joint 4
Humerus and glenoid 4
Glenoid labrum 5
Capsule 6
Muscles 6
Rotator cuff 6
Subscapularis 6
Infraspinatus 6
Teres minor 6
Supraspinatus 7
Biceps brachii 7
Deltoid muscle 8
Ligaments 8
Coracohumeral ligament 8
Superior glenohumeral ligament and middle glenohumeral ligament 8
Inferior glenohumeral ligament complex 8
Biomechanics 9
Passive constraints 9
Scapula 9
Labrum 9
Ligaments and capsule 9
Cohesion and adhesion 10
Negative intra-articular pressure 10
Dynamic stabilizers 10
Rotator cuff and deltoid muscle 10
Biceps brachii 10
References 11

INTRODUCTION The mobility of the upper extremity is based on the

Pathologies of the glenohumeral joint frequently require
surgery. Therefore, knowledge of the anatomy and its bio-
mechanical impact is essential for orthopedic surgeons in
order to treat patients most effectively. This chapter pro-
vides an overview of structures surgeons should take into
consideration pre-, intra-, and postoperatively.
anatomy and functional interaction of the shoulder com-
plex, which enables the widest range of motion of all joints
in the human body.
The bony shoulder complex consists of scapula, clavicle,
and humerus. These bones form the glenohumeral (GH)
joint, the acromioclavicular (AC) joint, and the sternocla-
vicular (SC) joint.1

3
4 Anatomy and biomechanics of the shoulder
The osteoarticular configuration of the glenohumeral
joint offers six degrees of freedom—three in rotation and
two in translation. The extent of translation varies in indi-
viduals, mainly depending on stiffness and laxity of the soft
tissue (capsule and ligaments) and less so on the muscu-
larly setting.
In the following the important anatomical structures
and their biomechanical relevance for motion and stabil-
ity are described.
LANDMARKS
Even experienced shoulder surgeons identify and point
out the important anatomical landmarks prior to surgery.
These relevant bony structures are the acromion, the spine
of scapula, the AC joint, the clavicle and the coracoid pro-
cess (Figure 1.1).
Acromioclavicular joint
The AC joint with the clavicle in prolongation, is the only
connection between scapula and bony thorax. It is a diar-
throdial joint with a disc in between the joint partners.2
Superiorly the AC joint is covered by the insertion of tra-
pezius and deltoid muscles. Inferior it is characterized by
S can be palpated in the deltopectoral groove. The coracoid
AC CL
A
C attached, medially the pectoralis minor muscle inserts.
Figure 1.1 Right shoulder. Anatomical landmarks: clavicula
(CL), acromioclavicular joint (AC), acromion (A), spina scapulae
(S), and coracoid process (C).
the coracoclavicular ligaments originating from the cora-
coid process, inserting to the lateral part of the clavicle.
The coracoclavicular ligaments are divided into conoid
band medially and trapezoid band laterally. The mean dis-
tances between the lateral edge of the clavicle to the inser-
tion site of the conoid ligament are 24.9 ± 3.8 mm (range
18.3–31.8 mm) and 46.3 ± 5.1 mm (range 21.9–43.9 mm)
to the conoid tuberosity.3
Beside the coracoclavicular ligaments, which stabi-
lize the clavicle in the vertical direction, the acromiocla-
vicular ligament strengthens the AC-capsule, preventing
enhanced horizontal translation.4 The AC ligament can be
separated into a superoposterior bundle and the anteroin-
ferior bundle. The superoposterior bundle runs posteriorly
toward the distal clavicle from the acromion at an average
angle of 30° to the joint surface.5
Acromion
The acromion forms the “roof of the shoulder.” The mean
distance between the apex of the humeral head and the
acromion is 9–10 mm on anteroposterior (a.p.) radio-
graphs.6 An acromiohumeral interval of less than 6 mm is
pathological, indicating a rotator cuff tear.6
Anatomical variations of the acromial arch were
described by Bigliani et al.7,8 They distinguish three mor-
phologic types on outlet radiographs: type I “flat,” type II
“curved,” and type III “hooked.” Type III acromion spurs
were found to be associated with superior rotator cuff dis-
ease in 70% of cases.7
Coracoid process
The coracoid process originates from the anterior superior
neck of the scapula. Its horizontal part curves lateral and
apex gives insertion to the conjoined tendons (short head
of the biceps brachii, coracobrachialis) (Figure 1.2). Lateral
at the horizontal portion the coracoacromial ligament is
Located close to the coracoid base, the coracoclavicular
ligaments originate at an average distance of 28.5 mm
from the anterior tip.9 The vertical part of the coracoid
process is supplied by the supra-scapular artery and the
horizontal part by branches of the axillary artery. It was
thought that preservation of the axillary artery branches
could be a possible solution to prevent non-union and lysis
of the bone transfer in the Latarjet procedure.10
GLENOHUMERAL JOINT
Humerus and glenoid
The humeral head has an articular surface of approxi-
mately 24 cm2 and is 2.5 cm in radius (a.p.-front).11
 Glenohumeral joint 5

PM C

CT

α
Figure 1.2 Left shoulder. Deltopectoral approach dur-
ing Latarjet procedure with exposed coracoid process (C),
conjoined tendons inferior (CT), and pectoralis minor tendon
medial (PM).

The humeral inclination angle varies between 130° and

150°, with 30° to 45° of retrotorsion11–13 (Figure 1.3).
In contrast, the glenoid fossa provides a relatively small
articular surface of 6 cm2.11,13 The face of the glenoid sur-
face appears pear-shaped, narrow superior and wider
inferior.14 In the normal population the glenoid fossa is 7° Figure 1.4 Axial CT-scan right shoulder. The glenoid fossa
retrotilted (range 3°–11°) to the scapula,15 with an average is on average 7° (α) retrotilted (between 3° and 11°) to the
superior glenoid tilt of 5°11,16 (Figure 1.4) scapula.

Due to the size mismatch of the joint components, only

part of the humeral head (25%–30%) is in contact with the
glenoid fossa, depending on the position of the arm.17,18

Glenoid labrum
The 4 mm wide and thick labrum consists of fibrocartilage
and surrounds the glenoid fossa.1,11 The superior pole gives
α origin to the long head of the biceps tendon (Figure 1.5).

LHB

HH L
G

Figure 1.5 Arthroscopy of a left shoulder via a poste-

Figure 1.3 Anteroposterior radiograph of a left shoul- rior standard portal. The long head of the biceps (LHB) origi-
der: Inclination (α) and retrotorsion of the humeral head are, nates from the superior labrum (L) and supraglenoid tubercle.
between 130° and 150° and 30° and 45°, respectively. (HH = humeral head, G = glenoid fossa).
6 Anatomy and biomechanics of the shoulder
Anterior inferior the labrum serves as attachment to the
inferior glenohumeral ligament, one of the most impor-
tant ligamentous stabilizers of the glenohumeral joint.11
Between the articular surface and the glenoid labrum
a thin gap is frequently observed. The outer labrum is
attached to the periost of the scapula neck, while the inner
labrum is covered with synovia.11
Cooper et al. distinguish two parts of the labrum.19
The inferior portion is more fibrous, elevated and directly
attached to the glenoid than the loosely attached and
meniscal-like superior one.19 Anatomical variations mainly
present in the superior labrum, like the sublabral foramen,
where the upper labrum is completely detached from the
glenoid. The frequency is reported to be 12%–18%.20,21
The congenital Buford complex is less common (fre-
quency 1.5%–6%). Here the anterosuperior labrum is
absent at the 1–3-o’clock position, instead it is replaced by
a thick middle glenohumeral ligament (MGHL), originat-
ing directly from the superior glenoid.20,22,23 Both, sub-
labral foramen and Buford complex are non-pathologic
variations and should not be misinterpreted as a SLAP- or
Bankart-lesion.19
The vascularization of the labrum by the surrounding
capsule is discussed, and should be considered with regards
to healing potential in case of surgical refixation.19,24
Capsule
The cylindric joint capsule originates from the glenoid
neck and inserts at the cartilage–bone interface of the
humeral head.25 However, the anterior attachment varies
between (1) insertion to the anterior labrum, (2) inser-
tion next to the scapular neck, and (3) insertion further
medial.25
The capsule is circumferentially tightened with the
exception of the axillary pouch. Ciccone et al. described
a global capsule thickness ranging from 1.32 to 4.47 mm.
The thickness increased from an average of 2.42 mm ante-
riorly to 2.80 mm in the inferior capsular pouch, with
close relation to the axillary nerve, and again thinned to
2.22 mm posteriorly.26 However, Itoi et al. reported a nar-
rower global thickness of 1.0–1.8 mm, whereas the poste-
rior capsule was also found to be thinnest.27
In a further cadaveric study Nimura et al. showed that
the superior capsule occupies a substantial part of the
greater tuberosity. The thinnest point was 11 mm poste-
rior to the anterior margin of the greater tuberosity and
close to the posterior edge of the supraspinatus tendon
insertion. The authors assumed this could contribute to
the etiology of degenerative rotator cuff tears in that area.28
The ligamentum semicirculare humeri or rotator cable
can be identified as a parallel-running bundle of connec-
tive tissue fibres forming a curved capsular–ligamentous
structure in the superolateral part of the glenohumeral
joint capsule. It arises from the superior part of the lesser
tubercle and inserts in the region between the middle and
inferior facets of the greater tubercle.29
MUSCLES
Rotator cuff
SUBSCAPULARIS
The subscapularis muscle is the largest muscle–tendon
unit of the rotator cuff, with an average maximum width
of 20 mm and maximum length of 40 mm.30 Whereas the
superior articular margin is tendinous, the inferior part
has a musculocapsular attachment30 (Figure 1.6). Its inser-
tion footprint at the lesser tuberosity shows a broad and
wide superior attachment that narrows distally to form a
trapezoidal shape. Cadaveric studies reported the mean
length of the footprint to be 2.5 cm. The mean width at
the most superior aspect of the insertion site was 1.8 cm
(range 1.5–2.6 cm), which was maintained for the upper
60% of the tendon insertion. The most inferior aspect of
the footprint was much narrower, with a mean width of
0.3 cm (range 0.1–0.7 cm).31
INFRASPINATUS
Second in size, with an average maximum width of 19 mm
and length of 29 mm, the infraspinatus muscle has a trap-
ezoidal footprint.30 The insertion site shows a medial
anteroposterior length of 22.6 ± 3.0 mm and an antero-
posterior length of 25.4 ± 3.3 mm. The maximal medio-
lateral width varies between 12.0 ± 2.7 mm.32
Superior the infraspinatus footprint passes into the
supraspinatus insertion site, inferiorly an intra-articular
cartilage-free area between the articular surface and the
infraspinatus insertion can be found—the “bare area”30
(Figure 1.7). The more inferior the infraspinatus gets
shortened and muscular.30
TERES MINOR
The teres minor muscle forms the smallest muscle–­
tendon unit, consisting predominantly of muscle tis-
sue and few tendinous fibers.30 It inserts below the
SSC
L
MGHL
HH
Figure 1.6 Arthroscopy of a left shoulder via a posterior
standard portal. Superior intra-articular tendinous unit of the
subscapularis muscle (SSC) behind slackened middle glenohu-
meral ligament (MGHL). (HH = humeral head, L = labrum).

ISP
BA
AS
Figure 1.7 Arthroscopy of a left shoulder via a posterior
standard portal. Gap between the articular surface (AS) and
the infraspinatus (ISP) insertion, so called “bare area” (BA).
infraspinatus muscle to the inferior facet on greater tuber-
osity of the humerus.
SUPRASPINATUS
The footprint of the supraspinatus is triangular. Based
on cadaveric studies width of the insertion site varies
between 6.7 and 16 mm. Medial and lateral length was
reported, respectively, as 20.9–32 mm and 1.3–6.4 mm
(Figure 1.8).28,30,32–35
The tendinous unit rises over the greater tuberosity with
an average width of 16 mm and length of 23 mm.30
BICEPS BRACHII
The biceps brachii tendon can be separated into a short
head (SHB) and long head (LHB). The SHB tendon inserts
to the tip of the coracoid process, lateral to the coraco-
brachialis muscle (Figure 1.2). The LHB tendon passes
the bicipital groove to intra-articular and turns, while
guided by the biceps pulley (see below), to the superior
SSP
BG
HH
Figure 1.8 Arthroscopy of a left shoulder via a posterior
standard portal. Medial intraarticular footprint of the anterior
edge of the supraspinatus tendon (SSP). Anterior to the SSP the
bicipital groove (BG) opens up. (HH = humeral head).
Muscles 7
SSP LHB
HH
Figure 1.9 Arthroscopy of a left shoulder via posterior
standard portal. “Pulley,” consisting of mainly ligamentous
structures. Posterior it is limited by the fibers of the supraspi-
natus (SSP) tendon. (HH = humeral head, LHB = long head of
the biceps tendon).
labrum (Figure 1.5). In a cadaver study including 100
shoulders, 50% of the LHBs originated directly from the
superior glenoid labrum. 36 Vangsness et al. distinguish
four types of attachment: (1) entire attachment to the
posterior labrum (22%), (2) major labral contribution is
posterior (33%), (3) equal contribution of anterior and
posterior labrum (37%), and (4) major attachment ante-
rior with minor contribution of the posterior labrum
(8%). 36
The intra-articular part of the LHB is routed by the
so-called “pulley” (Figures 1.9 and 1.10), consisting of the
coracohumeral ligament (CHL), the superior glenohu-
meral ligament (SGHL), and fibers of the subscapularis
and supraspinatus tendon.37,38
LHB
CHL
HL
SG
SSC
Figure 1.10 Arthroscopy of a left shoulder via a posterior
standard portal. The coracohumeral ligament (CHL) is passing
the rotator interval anterior to the supraspinatus tendon and
superior to the upper subscapularis margin (SSC). The superior
glenohumeral ligament (SGHL) is heading to the lesser tuber-
osity, inserting on the medial ridge of the bicipital groove dis-
tal to the CHL. (LHB = long head of the biceps tendon).
8 Anatomy and biomechanics of the shoulder
DELTOID MUSCLE
The deltoid muscle consists of three portions and seven
segments: (1) the anterior part originates from the lateral
end of the clavicle and anterior-third of the acromion, (2)
the middle part attaches to the mid-third of the lateral
aspect of the acromion, and (3) the posterior part to the
posterior third of the lateral acromion as well as the scapu-
lar spine.39,40
The insertion area of the deltoid muscle is the proximal-
lateral shaft of the humerus. Therefore the deltoid muscle
covers the glenohumeral joint from anterior, lateral and
posterior.39
LIGAMENTS
Coracohumeral ligament
The coracohumeral ligament (CHL) is divided into two
major bands, the anterior band and the posterior band.16
It originates from the lateral aspect of the base of the cora-
coid process, passing the rotator interval anterior to the
supraspinatus tendon and superior to the upper subscapu-
laris margin (Figure 1.10).41
The posterior part of the CHL irradiates into the fas-
cia of the supraspinatus tendon and capsule, anteriorly
to the insertion of the subscapularis tendon at the lesser
tuberosity. The two bands form a part of the biceps pul-
ley. Furthermore, the inferior part of the CHL unites with
the superior fibers of the superior glenohumeral ligament
(SGHL).
Superior glenohumeral ligament and middle
glenohumeral ligament
The superior glenohumeral ligament (SGHL) is the thin-
est of the three glenohumeral ligaments. It originates from
the supraglenoid tubercle, anterior to the insertion of the
long head of the biceps tendon. The SGHL passes the rota-
tor interval heading to the lesser tuberosity, inserting to
the medial ridge of the bicipital groove distal to the CHL
(Figure 1.10).42 In 3% of shoulders it is not verifiable.43 The
SGHL consists of direct and oblique fibers with different
origins, vectors, and insertions.
The direct fibers originate from the glenoid labrum
and the oblique fibers from the supraglenoid tubercle.44
However, in the absence of the MGHL they have a com-
mon origin from the glenoid labrum.
The direct fibers run parallel with the LHB to the medial
ridge of the bicipital groove. They partially bridge the
bicipital groove and form the superior part of the trans-
verse humeral ligament.44
The oblique fibers course over the intra-articular part
of the LHB and insert below the coracohumeral ligament
(CHL) into the humeral semicircular ligament.44
Compared to the SGHL, the MGHL is larger and more
variable in size. The width and thickness are 1–2 cm and up
to 4 mm, respectively.16 The origin is found on the supra-
glenoid tubercle and the anterosuperior labrum between
the 1- and 3-o’clock positions. Therefore, its anatomy is
similar to the SGHL. The fibers of the MGHL proceed
across the anterior capsule and blend with the posterior
portion of the subscapularis tendon approximately 2 cm
medial to its insertion on the lesser tuberosity.45 Cadaveric
studies show a verifiability of between 63% and 85%.23,42
The MGHL varies in length and thickness depending
on age. In the younger population it is well defined, but
in the elderly it gets thin and it sometimes can hardly be
identified.16 The MGHL has been shown to be a secondary
restraint for anterior and inferior translation. A variation
is a cord-like MGHL (Buford complex, see above). Thus,
decreased anterior translation is ensured when the MGHL
is morphologically more cord-like.46
Inferior glenohumeral ligament complex
Consisting of an anterior ligament, axillary pouch and
posterior ligament, the inferior glenohumeral ligament
complex (IGHLC) has the most important stabilizing
effect in the anteroinferior direction in extended exter-
nal and internal rotation while abducting the shoulder.38
The anterior and posterior inferior glenohumeral ligament
(IGHL) spans the inferior capsule and axillary pouch. The
anterior band is pulled taut in abduction and external
rotation (Figure 1.11). while the posterior band is pulled
taut in abduction and internal rotation.38
The anterior IGHL is attached to the glenoid in the
2- to 5-o’clock position. Two types of attachment can be
identified: major origin from the labrum with some fibers
extending to the glenoid neck (Type 1, 80%) and isolated
insertion to the glenoid neck (Type 2, 20%).19,47,48
Itoigawa et al. demonstrated in a cadaveric study that
the thickest part of the anterior IGHL–labrum complex is
alGHL
HH
Figure 1.11 Arthroscopy of a left shoulder via a posterior
standard portal. The anterior inferior glenohumeral ligament
(aIGHL) is pulled taut in abduction and external rotation.
(HH = humeral head).
Another random document with
no related content on Scribd:
received a dorsally-placed accessory gland, open to the exterior by a
median aperture placed ventrally a little way behind the mouth.
Life-history.—The egg undergoes a large portion of its
development within the body of the mother. In Linguatula
taenioides, which lives in the nasal cavities of the dog, the eggs pass
away with the nasal excretions. If these, scattered about in the grass,
etc., be eaten by a rabbit, the egg-shell is dissolved in the stomach of
the second host and a small larva is set free. In Porocephalus
proboscideus and others, which inhabit the lungs of snakes, the eggs
pass along the alimentary canal and leave the body with the faeces.
They also must be eaten by a second host if development is to
proceed.
The larva which emerges when
the egg-shell is dissolved has a
rounded body provided with two
pairs of hooked appendages, and
a tail which is more or less
prominent in different species
(Figs. 259, 260). Each appendage
bears a claw, and is strengthened
by a supporting rod or skeleton.
Anteriorly the head bears a
boring apparatus of several
chitinous stylets. The various
internal organs are in this stage
already formed, though in a Fig. 259.—A late larval stage of
somewhat rudimentary state, and Porocephalus proboscideus, seen from
the side. Highly magnified. (From
it is doubtful if the anus has yet Stiles.) 1, primordium of first pair of
appeared. chitinous processes; 2, primordium of
By means of its boring second pair of chitinous processes; 3,
apparatus, and aided by its mouth; 4, ventral ganglion; 5,
hooked limbs, the larva now receptaculum seminis; 6, oviduct; 7,
ovary; 8, anus; 9, vagina.
works its way through the
stomach-walls of its second host,
and comes to rest in the liver or in some other viscus. Its presence in
the tissues of its second host causes the formation of a cyst, and
within this the larva rests and develops. In man, at least, the cysts
often undergo a calcareous degeneration, and Virchow states “dass
 beim Menschen das
Pentastomum am häufigsten von
allen Entozoen zu
Verwechselungen mit echten
Tuberkeln Veranlassungen giebt.”
The larva moults several times,
and loses its limbs, which seem to
have no connexion with the
paired hooks in the adult (Fig.
256). The internal organs slowly
assume the form they possess in
the adult. The larva is at first
quite smooth, but as it grows the
annulations make their
appearance, arising in the middle
and spreading forward and
Fig. 260.—Larva of Porocephalus backward (Fig. 259). In this
proboscideus, seen from below. Highly encysted condition the larva
magnified. (From Stiles.) 1, Boring, remains coiled up for some
anterior end; 2, first pair of chitinous months, according to Leuckart;
processes seen between the forks of the
second pair; 3, ventral nerve-ganglion;
six in the case of L. taenioides,
4, alimentary canal; 5, mouth; 6 and 7, and a somewhat shorter
[385]
period,
gland-cells. according to Stiles, in the case
of P. proboscideus.
The frequency of what used to
be called Pentastoma denticulatum (= the larval form of L.
taenioides) in the body of man depends on the familiarity of man
with dogs. Klebs and Zaeslin found one larva in 900 and two in 1914
autopsies. Laenger[386] found the larva fifteen times in about 400
dissections, once in the mesentery, seven times in the liver, and
seven times in the wall of the intestine. After remaining encysted for
some time it may escape, and begins wandering through the tissues,
aided by its hooks and annulations, a proceeding not unaccompanied
by danger to its host. Should the latter be eaten by some carnivorous
animal, the larva makes its way into the nasal cavities or sinuses, or
into the lungs of the flesh-eating creature, and there after another
ecdysis it becomes adult. If, however, the second host escapes this
fate, the larvae re-encyst themselves, and then if swallowed they are
said to bore through the intestine
of the flesh-eater, and so make
their way to their adult abode.
Systematic.[387]—The
Pentastomida are a group much
modified by parasitism, which
has so deeply moulded their
structure as to obscure to a great
extent their origin and affinities.
The larva, with its clawed limbs,
recalls the Tardigrades and
certain Mites, e.g. Phytoptus,
where only two pairs of limbs
persist, and where the abdomen Fig. 261.—Encysted form of
is elongated and forms a large Porocephalus protelis, × 1, lying in the
proportion of the body. The mesentery of its host. (From Hoyle.)
resemblances to a single and
somewhat aberrant genus must
not, however, be pressed too far. The striated muscles, the ring-like
nature of the reproductive organs and their ducts, perhaps even the
disproportion both in size and number of the females to the males,
are also characters common to many Arachnids.
The Pentastomida include three genera, Linguatula, Fröhlich,
Porocephalus, Humboldt, and Reighardia, Ward.[388] The first two
were regarded by Leuckart as but sub-genera, but Railliet[389] and
Hoyle[390] have raised them to the rank of genera. They are
characterised as follows:—
Linguatula, body flattened, but dorsal surface arched; the edges of
the fluke-like body crenelated; the body-cavity extends as diverticula
into the edges of the body.
Porocephalus, body cylindrical, with no diverticula of the body-
cavity.
Reighardia, devoid of annulations, transparent, with poorly
developed hooks and a mouth-armature.
The following is a list of the species with their primary and
secondary or larval hosts:—
i. Linguatula pusilla, Diesing, found in the intestine of the fresh-
water fish Acara, a South American genus of the Cichlidae. This
is possibly the immature form of L. subtriquetra.
ii. L. recurvata, Diesing, found in the frontal sinuses and the
trachea of Felis onca.
iii. L. subtriquetra, Diesing, found in the throat of Caiman
latirostris and C. sclerops, perhaps the mature form of L. pusilla.
iv. L. taenioides, Lamarck, found in the frontal sinuses and nasal
chambers of the dog and ounce, and in the nasal cavities of the
wolf, fox, goat, horse, mule, sheep, and man, and in the trachea
of the ounce. The immature form has been found in or on the
liver of the cat, guinea-pig, and horse; in the lungs of the ox, cat,
guinea-pig, porcupine, hare, and rabbit; in the liver and
connective tissue of the small intestine of man; and in the
mesenteric glands of the ox, camel, goat, sheep, antelope, fallow-
deer, and mouse.
v. Porocephalus annulatus, Baird, found in the lungs of the
Egyptian cobra, Naja haje; the immature form is thought to live
encapsuled in a species of Porphyrio[391] and in the Numidian
Crane.
vi. P. aonycis, Macalister, from the lungs of an Indian otter taken
in the Indus.
vii. P. armillatus, Wyman, found in the adult state in the lungs of
certain African pythons, and in the lion; in the larval form it
occurs encysted in the abdomen of the Aard-wolf, the mandril,
and man—usually in negroes. Its migrations in the body of its
second host sometimes cause fatal results.
viii. P. bifurcatus, Diesing, found in the body-cavity of certain
snakes, and in the lungs of boa-constrictors and the legless
lizard, Amphisbaena alba. Possibly an immature form.
ix. P. clavatus, Lohrmann, found in the lungs of the Monitor lizard.
x. P. crocidura, Parona, found in the peritoneum of the “musk-rat”
Crocidura in Burmah. Probably a larval form.
xi. P. crotali, Humboldt, found in the lungs, body-cavity, kidneys,
spleen, and mesentery of many snakes and lizards, and of the
lion and leopard. The immature forms occur in the liver and
abdominal cavity of species of opossum, armadillo, mouse,
raccoon, bat, and marmoset.
xii. P. geckonis, Dujardin, found in the lungs of a Siamese gecko.
xiii. P. gracilis, Diesing, found free in the body-cavity or
encapsuled on the viscera and mesenteries of South American
fishes, snakes, and lizards,
xiv. P. heterodontis; Leuckart, found encapsuled in the abdominal
muscles and mesentery of a species of Heterodon.
xv. P. indicus,[392] v. Linst., found in the trachea and lungs of
Gavialis gangeticus.
xvi. P. lari, Mégnin, found in the air-sacs of the Burgomaster or
Glaucous gull, Larus glaucus of the Polar seas.
xvii. P. megacephalus, Baird, found embedded in the flesh of the
head of an Indian crocodile, C. palustris, the “Mugger.” Probably
a larval form.
xviii. P. megastomus, Diesing, found in the lungs of a fresh-water
tortoise, Hydraspis geoffroyana.
xix. P. moniliformis, Diesing, found in the lungs of pythons.
xx. P. najae sputatricis, Leuckart, found encapsuled in the
abdominal muscles and peritoneum of the cobra, Naja
tripudians. Probably a larval form.
xxi. P. oxycephalus, Diesing, found in the lungs of crocodiles and
alligators.
xxii. P. platycephalus, Lohrmann, habitat unknown.
xxiii. P. subuliferus, Leuckart, in the lungs of the cobra Naja haje.
xxiv. P. teretiusculus, Baird, found in the lungs and mouth of
certain Australian snakes.
xxv. P. tortus, Shipley, found in the body-cavity of a snake,
Dipsadomorphus irregularis, taken in New Britain.
xxvi. Reighardia, sp., Ward, found in the air-sacs of Bonaparte’s
gull and the common North American tern.
 PYCNOGONIDA

BY

D’ARCY W. THOMPSON, C.B., M.A. Trinity College

Professor of Natural History in University College, Dundee
 CHAPTER XXI
[393]
PYCNOGONIDA

Remote, so far as we at present see, from all other Arthropods, while yet
manifesting the most patent features of the Arthropod type, the Pycnogons
constitute a little group, easily recognised and characterised, abundant and
omnipresent in the sea. The student of the foreshore finds few species and seldom
many individuals, but the dredger in deep waters meets at times with prodigious
numbers, lending a character to the fauna over great areas.
The commonest of our native species,
or that at least which we find the oftenest,
is Pycnogonum littorale (Phalangium
littorale, Ström, 1762). We find it under
stones near low water, or often clinging
louse-like to a large Anemone. The squat
segmented trunk carries, on four pairs of
strong lateral processes, as many legs,
long, robust, eight-jointed, furnished each
with a sharp terminal claw. In front the
trunk bears a long, stout, tubular
proboscis, at the apex of which is the
mouth, suctorial, devoid of jaws; the body
terminates in a narrow, limbless,
unsegmented process, the so-called
“abdomen,” at the end of which is the
anal orifice. The body-ring to which is
attached the first pair of legs, bears a
tubercle carrying four eye-spots; and Fig. 262.—Pycnogonum littorale, Ström, × 2.
below, it carries, in the male sex, a pair of
small limbs, whose function is to grasp
and hold the eggs, of which the male animal assumes the burden, carrying them
beneath his body in a flattened coherent mass. In either sex a pair of sexual
apertures open on the second joints of the last pair of legs. The integument of body
and limbs is very strongly chitinised, brown in colour, and raised into strong bosses
or tubercles along the middle line of the back, over the lateral processes, and from
joint to joint of the limbs. The whole animal has a singular likeness to the Whale-
louse, Cyamus mysticeti (well described by Fr. Martins in 1675), that clings to the
skin of the Greenland Whale as does Pycnogonum to the Anemone, a resemblance
close enough to mislead some of the older naturalists, and so close that Linnaeus,
though in no way misled thereby, named it Phalangium balaenarum. The substance
of the above account, and the perplexity attending the classification of the animal,
are all included in Linnaeus’s short description:[394] “Simillimus Onisco Ceti, sed
pedes omnes pluribus articulis, omnes perfecti, nec plures quam octo. Dorsum
rubrum, pluribus segmentis; singulis tribus mucronibus. Cauda cylindrica,
brevissima, truncata. Rostrum membranaceum, subsubulatum, longitudine pedum.
Genus dubium, facie Onisci ceti; rostro a reliquis diversum. Cum solo rostro absque
maxillis sit forte aptius Acaris aut proprio generi subjiciendum.... Habitat in mari
norvegico sub lapidibus.”[395]
The common Pycnogonum is, by
reason of the suppression of certain
limbs, rather an outlying member than a
typical representative of the Order, whose
common characters are more strikingly
and more perfectly shown in species, for
instance, of Nymphon. Of this multiform
genus we have many British species, some
of the smaller being common below tide-
marks, creeping among weeds or clinging
like Caprellae with skeleton limbs to the
branches of Zoophytes, where their
slender forms are not easily seen. In
contrast to the stouter body and limbs of
Pycnogonum, the whole fabric of
Nymphon tends to elongation; the body is
drawn out so that the successive lateral
processes stand far apart, and a slender
neck intervenes between the oculiferous
tubercle and the proboscis; the legs are
produced to an amazing length and an
extreme degree of attenuation: “mirum
tam parvum corpus regere tam magnos
pedes,” says Linnaeus. Above the base of
the proboscis are a pair of three-jointed
appendages, the two terminal joints of
which compose a forcipate claw; below
Fig. 263.—Dorsal view of Nymphon brevirostre, and behind these come a pair of delicate,
Hodge, × 6. Britain. palp-like limbs of five joints; and lastly,
on the ventral side, some little way
behind these, we find the ovigerous legs
that we have already seen in the male Pycnogonum, but which are present in both
sexes in the case of Nymphon. At the base of the claw which terminates each of the
eight long ambulatory legs stands a pair of smaller accessory or “auxiliary” claws.
The generative orifices are on the second joint of the legs as in Pycnogonum, but as
a rule they are present on all the eight legs in the female sex, and on the two
hindmost pairs in the male. One of the Antarctic Nymphonidae (Pentanymphon)
and one other Antarctic genus less closely related (Decolopoda) have an extra pair of
legs. No other Pycnogon, save these, exhibits a greater number of appendages than
Nymphon nor a less number than Pycnogonum, nor are any other conspicuous
organs to be discovered in other genera that are not represented in these two: within
so narrow limits lie the varying characters of the group.
In framing a terminology for the parts and members of the body, we encounter an
initial difficulty due to the ease with which terms seem applicable, that are used of
more or less analogous parts in the Insect
or the Crustacean, without warrant of
homology. Thus the first two pairs of
appendages in Nymphon have been
commonly called, since Latreille’s time,
the mandibles and the palps (Linnaeus
had called them the palps and the
antennae), though the comparison that
Latreille intended to denote is long
abandoned; or, by those who leaned, with
Kröyer and Milne-Edwards, to the
Crustacean analogy, mandibles and
maxillae. Dohrn eludes the difficulty by
denominating the appendages by simple
numbers, I., II., III., ... VII., and this
method has its own advantages; but it is
better to frame, as Sars has done, a new
nomenclature. With him we shall speak of
the Pycnogon’s body as constituted of a
trunk, whose first (composite) segment is
the cephalic segment or head, better
perhaps the cephalothorax, and which Fig. 264.—Nymphon brevirostre, Hodge. Head,
terminates in a caudal segment or from below, showing chelophores, palps, and
abdomen; the “head” bears the proboscis, ovigerous leg.
the first appendages or “chelophores,” the
second or “palps,” the third, the false or
“ovigerous” legs, and the first of the four pairs of “ambulatory” legs. The chelophores
bear their chela, or “hand,” on a stalk or scape; the ambulatory legs are constituted
of three coxal joints, a femur, two tibial joints, a tarsus, and a propodus, with its
claws, and with or without auxiliary claws.
The Body.—The trunk with its lateral processes may be still more compact than
in Pycnogonum, still more attenuated than in Nymphon.
In a few forms (e.g. Pallene, Ammothea, Tanystylum, Colossendeis) the last two,
or even more, segments of the trunk are more or less coalescent. In Rhynchothorax
the cephalic segment is produced into a sharp-pointed rostrum that juts forward
over the base of the proboscis. The whole body and limbs may be smooth,
tuberculated, furnished with scattered hairs, or sometimes densely hispid.
 Fig. 265.—A, Colossendeis proboscidea, Sabine, Britain; B, Ammothea echinata,
Hodge, Britain; C, Phoxichilus spinosus, Mont., Arctic Ocean. (The legs omitted.)

The proboscis varies much in shape and size. It may be much longer or much
shorter than the body, cylindrical or tumid, blunt or pointed, straight or (e.g.
Decolopoda) decurved; usually firmly affixed to the head and pointing straight
forwards; sometimes (Eurycide, Ascorhynchus) articulated on a mobile stalk and
borne deflexed beneath the body.
Chelophores.—The first pair of appendages or chelophores are wanting in the
adult Pycnogonum, Phoxichilus, Rhynchothorax, and Colossendeis.[396]
In Ammothea and its allies they are extremely rudimentary in the adult, being
reduced to tiny knobs in Tanystylum and Trygaeus, and present as small two-
jointed appendages in Ammothea; in this last, if not in the others also, they are
present in complete chelate form in the later larval stages.
 Fig. 266.—A, B, Chelophores of Ascorhynchus abyssi, G.O.S. A, Young; B, adult.
(After Sars.) C, Anterior portion of Ammothea hispida, Hodge, Jersey: late larval
stage (= Achelia longipes, Hodge), showing complete chelae. D, Chela of
Eurycide hispida, Kr.

In Eurycide, Ascorhynchus, and Barana they are usually less atrophied, but yet
comparatively small and with imperfect chelae, while in some Ascorhynchi (A.
minutus, Hoek) they are reduced to stumps.

Fig. 267.—Chelae of species of Nymphonidae: A, Nymphon brevirostre, Hodge;

B, Boreonymphon robustum, Bell; C, Chaetonymphon macronyx, G.O.S.; D,
Nymphon elegans, Hansen.

In Pallenopsis the scape of the chelophore consists of two joints, as also in

Decolopoda and some Ascorhynchus: in Nymphon, Phoxichilidium, Pallene, and
Cordylochele of one only; in all these the terminal portion or “hand” forms a
forcipate “chela,” of which the ultimate joint forms the “movable finger.” In some
species of Nymphon the chela is greatly produced and attenuated, and armed with
formidable serrate teeth on its opposing edges; in others it is shortened, with blunter
teeth; in Boreonymphon robustum the
claws are greatly curved, with a wide gap
between. In this last, and in
Phoxichilidium, the opposing edges are
smooth and toothless. In Cordylochele
the hand is almost globular, the movable
finger being shortened down, and half
enclosed by the other.
Palpi.—The second pair of
appendages, or palps, are absent, or all
but absent, in the adult Pycnogonum,
Phoxichilus, Phoxichilidium, Pallene, and
their allies. In certain of these cases, e.g.
Phoxichilidium, a knob remains to mark
their place; in others, e.g. Pallenopsis, a
single joint remains; in a few Pallenidae a
sexual difference is manifested, reduction
of the appendage being carried further in
the female than in the male. The Fig. 268.—Proboscis and chelophores of
composition of the palps varies in the Cordylochele longicollis, G.O.S. (After Sars.)
genera that possess them. In Nymphon
there are five joints, and their relative
lengths (especially of the terminal ones) are much used by Sars in defining the many
species of the genus. The recently described Paranymphon, Caullery, has palps of
six or seven joints. In the Ammotheidae the number of joints ranges from five or six
in Tanystylum to nine (as a rule) in Ammothea and Oorhynchus, or ten, according
to Dohrn, in certain species of Ammothea. Colossendeis and the Eurycididae have a
ten-jointed palp, which in this last family is very long and bent in zigzag fashion, as
it is, by the way, also in Ammothea. The terminal joints of the palp are in all cases
more or less setose, and their function is conjecturally tactile.
Ovigerous Legs.—Custom sanctions for these organs an inappropriate name,
inasmuch as it is only in the males that they perform the function which the name
connotes.[397] They probably also take some part, as Hodgson suggests, in the act of
feeding.
Fig. 269.—Eurycide hispida, Kr., showing
stalked proboscis and zigzag palps.

Fig. 270.—Ovigerous legs of A, Phoxichilus spinosus, Mont.; B, Phoxichilidium

femoratum, Rathke; C, Anoplodactylus petiolatus, Kr.; D, Colossendeis
proboscideus, Sab.
 In Pycnogonum, Phoxichilus,
Phoxichilidium, and their immediate
allies they are absent in the female; in all
the rest they are alike present in both
sexes, though often somewhat smaller in
the female than in the male. They are
always turned towards the lower side of
the body, and in many cases even their
Fig. 271.—Terminal joints of ovigerous leg of point of origin is wholly ventral. The
Rhynchothorax mediterraneus, Costa. number of joints varies: in

Phoxichilidium five, Anoplodactylus six,

Phoxichilus seven; in Paranymphon
eight; in Pycnogonum nine, with, in
addition, a terminal claw; in the
Ammotheidae from seven (Trygaeus) to
ten, without a claw; in Pallenidae ten,
with or without a claw; in
Rhynchothorax, Colossendeis, Eurycide,
Ascorhynchus, Nymphon, ten and a claw.
The appendage, especially when long, is
apt to be wound towards its extremity
into a spiral, and its last four joints
usually possess a peculiar armature. In
Rhynchothorax this takes the form of a
stout toothed tubercle on each joint; in
Colossendeis of several rows of small Fig. 272.—Nymphon brevirostre, Hodge.
Terminal joints of ovigerous leg, with magnified
imbricated denticles; in Nymphon and “tooth.”
Pallene of a single row of curious serrate
and pointed spines, each set in a little
membranous socket.
 Fig. 273.—Nymphon strömii, Kr. Male carrying egg-masses on his ovigerous
legs.

Fig. 274.—Terminal joints (tarsus and propodus) of legs. 1, Chaetonymphon

hirtum, Fabr.; 2, N. strömii, Kr.; 3, Nymphon brevirostre, Hodge; 4, Ammothea
echinata, Hodge; 5, Ascorhynchus abyssi, G.O.S. (All after Sars.)

Legs.—The four pairs of ambulatory legs are composed, in all cases without
exception, of eight joints if we exclude, or nine if we include, the terminal claw. They
vary from a length about equal to that of the body (Pycnogonum, Rhynchothorax,
Ammothea) to six or seven times as much, perhaps more, in Nymphon and
Colossendeis, the fourth, fifth, and sixth joints being those that suffer the greatest
elongation. The seventh joint, or tarsus, is usually short, but in some Nymphonidae
is much elongated; the eighth, or propodus, is usually somewhat curved, and usually
possesses a special armature of simple or serrate spines. The auxiliary claws,
sometimes large, sometimes small, lie at the base of the terminal claw in
Ammotheidae, Phoxichilidae, in Phoxichilidium, in most Pallenidae, in nearly all
Nymphonidae. Their presence or absence is often used as a generic character,
helping to separate, e.g., Pallene from Pseudopallene and Pallenopsis, and
Phoxichilidium from Anoplodactylus; nevertheless they may often be detected in a
rudimentary state when apparently absent. The legs are smooth or hirsute as the
body may happen to be.
Fig. 275.—Legs of A, Pallene brevirostris, Johnston; B, Anoplodactylus
petiolatus, Kr.; C, Phoxichilus spinosus, Mont.; D, Colossendeis proboscidea,
Sabine; E, Ammothea echinata, Hodge, ♂.
 Fig. 276.—Boreonymphon robustum, Bell. Male with young, slightly enlarged.
Faeroe Channel.

Glands.—In some or all of the appendages of the Pycnogonida may be found

special glands with varying and sometimes obscure functions. The glands of the
chelophores (Fig. 280, p. 522) are present in the larval stages only. They consist of a
number of flask-shaped cells[398] lying within the basal joint of the appendage, and
generally opening at the extremity of a long, conspicuous, often mobile, spine (e.g.
Ammothea (Dohrn), Pallene, Tanystylum (Morgan), Nymphon brevicollum and N.
gracile (Hoek)). They secrete a sticky thread, by means of which the larvae attach
themselves to one another and to the ovigerous legs of the male parent. In Nymphon
hamatum, Hoek, the several filaments secreted by the separate sacculi of the gland
issue separately. In Pycnogonum the spine on which the gland opens is itself
prolonged into a long fine filament, and here, according to Hoek, the gland is in all
probability functionless and rudimentary. Hoek has failed to find the gland in
Ascorhynchus, and also in certain Nymphonidae (e.g. Boreonymphon robustum,
Bell), in which the young are more than usually advanced at the time of hatching.
The gland has also been described by Lendenfeld and others in Phoxichilidium,
whose larvae do not cling together but live a parasitic life; in this genus the long
spine or tubercle is absent on which the orifice is usually situated, and, according to
Lendenfeld, the secretion issues from many small orifices set along the opposing
edges of the chela. Of the two species described by Dohrn as Barana castelli and B.
arenicola, the former has the spine of inordinate length, more than twice as long as
the whole body, chelophore and all; while in the latter (which species rather
resembles Ascorhynchus) the spine is altogether absent.
In the palps and ovigerous legs of the adult are found glandular bodies of a hollow
vesicular form with a simple lining of cells, the vesicle being divided within by a
septum with a central orifice, the outer and smaller half opening to the exterior.
These glands are probably of general occurrence, but they have been but little
investigated. They lie usually in the fourth and fifth joints of the palp, and the third
and fourth joints of the ovigerous leg. Hoek describes them in Discoarachne
(Tanystylum) as lying within the elongated third joint of the palp, and opening by a
sieve-plate at the end of the second joint. In Ammothea (Dohrn) and Ascorhynchus
(Hoek) they open on a small tubercle situated on the fifth joint of the palp. In
Nymphon, Hoek describes them as opening by a small pore on the fourth joint of the
ovigerous leg. Dohrn failed to find them in Pycnogonum, but in Phoxichilus,
Phoxichilidium and Pallene he discovered the glands appertaining to the palps,
though the palps themselves have disappeared in those genera; he has found the
glands also in Ammothea, in larvae that have not yet attained their full complement
of legs.
The males in nearly all cases are known to possess glands in the fourth joints or
thighs of all the ambulatory legs, and these glands without doubt act as cement-
glands, emitting, like the chelophoral glands of the larvae, a sticky thread or threads
by which the eggs and young are anchored to the ovigerous legs. In some species of
Nymphon and of Colossendeis Hoek could not find these, and he conjectures them
to be conspicuous only in the breeding season. While in most cases these glands
open by a single orifice or by a few pores grouped closely together, in Barana,
according to Dohrn, and especially in B. arenicola, the pores are distributed over a
wide area of the femoral joint.[399] In Discoarachne (Loman) and Trygaeus they
open into a wide chitinised sac with tubular orifice. While the function of these last
glands and of the larval glands seems plain enough, that of those which occur in the
palps and ovigerous legs of both sexes remains doubtful.
In their morphological nature the two groups of glands are likewise in contrast,
the former being unicellular glands, such as occur in various parts of the integument
of the body and limbs of many Crustacea; while the latter are segmentally arranged
and doubtless mesoblastic in origin, like the many other segmental excretory organs
(or coelomoducts) of various Arthropods.
By adding colouring matters (acid-fuchsin, etc.) to the water in which the animals
were living, Kowalevsky demonstrated the presence of what he believed to be
excretory organs in Phoxichilus, Ammothea, and Pallene. These are small groups of
cells, lying symmetrically near the posterior borders of the first three body-
segments, and also near the bases of the first joints of the legs, dorsal to the
alimentary canal.[400]
Alimentary System.—The proboscis is a very complicated organ, and has been
elaborately described by Dohrn.[401] It is a prolongation of the oral cavity, containing
a highly developed stomodaeum, but showing no sign of being built up of limbs or
gnathites. The mouth, situated at its apex, is a three-sided orifice, formed by a
dorsal[402] and two lateral lobes; and hence the proboscis has been assumed by some,