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Sports
Cardiology
Care of the Athletic Heart
from the Clinic to the Sidelines
David J. Engel
Dermot M. Phelan
Editors

123
Sports Cardiology
David J. Engel • Dermot M. Phelan
Editors

Sports Cardiology
Care of the Athletic Heart from the Clinic
to the Sidelines
Editors
David J. Engel Dermot M. Phelan
Division of Cardiology Sports Cardiology Center
Columbia University Irving Medical Center Hypertrophic Cardiomyopathy Center
New York, NY Atrium Health Sanger Heart &
USA Vascular Institute
Charlotte, NC
USA

ISBN 978-3-030-69383-1    ISBN 978-3-030-69384-8 (eBook)

https://doi.org/10.1007/978-3-030-69384-8

© Springer Nature Switzerland AG 2021

This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part of
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The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
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Preface

Participation in organized sports across the globe has markedly increased over the
past decade, and, in parallel, the clinical practice and research activity centered on
the cardiac care for athletes within the field of sports cardiology has expanded expo-
nentially. Recognizing the unique diagnostic and management challenges in opti-
mizing the heart health of athletes and reflective of the increasing importance
assigned to protecting the hearts of athletes, the American College of Cardiology
(ACC) in 2011 launched the ACC Section of Sports and Exercise Cardiology.
A foundation in the growth of sports cardiology has been the development of a
refined and enhanced understanding of the physiological manifestations of exercise
on the heart. This improved characterization of exercise-induced cardiac remodel-
ing, recognizing the relative influence of such modifiers as sport type, duration and
intensity of training, age, gender, race, size, and genetics, has vastly improved our
ability to screen for subclinical cardiac disease and differentiate normal physiology
from pathology. It is essential for healthcare providers who screen and treat athletes
at all skill levels to have a firm grasp of the tenets of sports cardiology and readily
available reference data encompassing the key elements within this growing field.
The cumulative clinical experience gained from caring for athletes training and
competing with existing cardiac conditions has resulted in the rapid evolution of
recommendations guiding sporting participation and the recognition of the impor-
tance of shared decision-making. A contemporaneous challenge has been the devas-
tation wrought by the COVID-19 pandemic. Sport and health organizations now
confront significant challenges designing and implementing safe athlete return to
play (RTP) strategies. In this textbook, we will review the critical issues and data
surrounding concerns of potential cardiac sequelae of COVID-19, and their impact
on athlete screening and RTP plans, as the newest element in the field of sports
cardiology.
Finally, the field of sports cardiology has pushed the practicing cardiologist from
the clinical facilities to the sports training facilities where they must participate in
the acute evaluation and management of athletes in addition to provide guidance on
effective emergence action plans.

v
vi Preface

Reflecting these challenges, this textbook is divided into three parts:

1. Pre-participation Cardiac screening of Athletes
2. Management and Recommendations for Athletes with Existing Cardiac
Disorders
3. Sideline Management of Acute Cardiac Conditions in Athletes
The purpose of this textbook is to assist healthcare providers manage the cardiac
care of athletes across the spectrum of these essential components. We will review
best practices for using and interpreting diagnostic tests commonly employed in the
cardiac evaluation of athletes, including the 12-lead electrocardiogram, advanced
cardiac imaging, and genetic testing. Treatment of cardiac disorders, ranging from
acute symptoms that develop suddenly during competition to chronic conditions
that require longitudinal management and assessment, will be reviewed with incor-
poration of latest guideline recommendations. This textbook will provide a frame-
work to aid in the provision of optimal care for athletic patients of all ages both on
and off the playing field.

New York, NY, USA David J. Engel

Charlotte, NC, USA Dermot M. Phelan
Contents

1 The Cardiovascular History and Examination ������������������������������������    1

John DiFiori, Chad Asplund, and James C. Puffer
2 Using an Electrocardiogram as a Component of
Athlete Screening ������������������������������������������������������������������������������������   13
David J. Engel
3 Diagnostic Approach after Initial Abnormal Screening ����������������������   31
Matthew W. Martinez
4 Practical Use of Genetic Testing in Athletes������������������������������������������   53
Isha Kalia, Farhana Latif, Muredach P. Reilly, and Marc P. Waase
5 Management of Hypertension in Athletes����������������������������������������������   69
D. Edmund Anstey and Daichi Shimbo
6 Valvular Heart Disease����������������������������������������������������������������������������   85
Tamanna K. Singh
7 Hypertrophic Cardiomyopathy��������������������������������������������������������������   97
Dermot Phelan and John Symanski
8 Other Cardiomyopathies������������������������������������������������������������������������ 111
Bradley Lander and David J. Engel
9 Inflammatory Cardiac Disorders in the Athlete ���������������������������������� 129
Kenneth G. Zahka, Nishant P. Shah, and Kara Denby
10 Atrial Fibrillation������������������������������������������������������������������������������������ 143
Kyle Mandsager and Dermot M. Phelan
11 Sports Participation in Patients with Congenital
Long QT Syndrome �������������������������������������������������������������������������������� 157
Salima Bhimani, Jared Klein, and Peter F. Aziz

vii
viii Contents

12 Other Arrhythmic Disorders: WPW, CPVT,

Brugada and Idiopathic VF/VT�������������������������������������������������������������� 171
Jeffrey J. Hsu and Eugene H. Chung
13 Cardiovascular Implantable Electronic Devices in Athletes���������������� 195
Benjamin H. Hammond and Elizabeth V. Saarel
14 Diagnosis and Management of Coronary
Artery Disease in Athletes ���������������������������������������������������������������������� 211
Prashant Rao and David Shipon
15 Marfan Syndrome and Other Genetic Aortopathies���������������������������� 227
Jeffrey S. Hedley and Dermot M. Phelan
16 Congenital Heart Disease: Approach to Evaluation,
Management, and Physical Activity ������������������������������������������������������ 245
Silvana Molossi and Hitesh Agrawal
17 Sleep Disorders in Athletes���������������������������������������������������������������������� 275
Meeta Singh, Michael Workings, Christopher Drake,
and Thomas Roth
18 Chest Pain and Dyspnea�������������������������������������������������������������������������� 297
David C. Peritz and John J. Ryan
19 The Evaluation of Palpitations and Dizziness in the Athlete �������������� 321
Brad Witbrodt and Jonathan H. Kim
20 The Collapsed Athlete������������������������������������������������������������������������������ 343
Justine S. Ko and George Chiampas
21 Cardiac Arrest in Athletes���������������������������������������������������������������������� 361
Brian J. Cross, Shayna Weinshel, and Marc Estes
22 Commotio Cordis in Athletes������������������������������������������������������������������ 375
Mohita Singh and Mark S. Link
23 The Impact of COVID-19 on Sports Cardiology���������������������������������� 383
Bradley Lander, David J. Engel, and Dermot M. Phelan
Index������������������������������������������������������������������������������������������������������������������ 395
Contributors

Hitesh Agrawal, MD, MBA, FSCAI Department of Pediatric Cardiology,

University of Texas, Austin, TX, USA
D. Edmund Anstey, MD, MPH The Columbia Hypertension Center, Columbia
University Irving Medical Center, New York, NY, USA
Chad Asplund, MD, MPH Department of Orthopedics and Sports Medicine,
Mayo Clinic, Minneapolis, MN, USA
Peter F. Aziz, MD Cleveland Clinic Children’s, Cleveland, OH, USA
Salima Bhimani, MD Cleveland Clinic Children’s, Cleveland, OH, USA
George Chiampas, DO, CAQSM, FACEP Departments of Emergency Medicine
and Orthopedic Surgery, Feinberg School of Medicine, Northwestern University,
Chicago, IL, USA
Eugene H. Chung, MD Department of Medicine, University of Michigan, Ann
Arbor, MI, USA
Brian J. Cross, MD Division of Cardiology, VA Pittsburgh Health System,
Pittsburgh, PA, USA
Kara Denby, MD Department of Cardiovascular Medicine, Cleveland Clinic,
Cleveland, OH, USA
John DiFiori, MD Sports Medicine Institute, Hospital for Special Surgery, New
York, NY, USA
Christopher Drake, PhD Department of Sleep Medicine, Henry Ford Health
System, Detroit, MI, USA
David J. Engel, MD, FACC Division of Cardiology, Columbia University Irving
Medical Center, New York, NY, USA
Marc Estes, MD Heart and Vascular Institute, University of Pittsburgh,
Pittsburgh, PA, USA

ix
x Contributors

Benjamin H. Hammond, MD Division of Pediatric Cardiology, Cleveland Clinic

Children’s and Pediatric Institute, Cleveland, OH, USA
Jeffrey S. Hedley, MD Department of Cardiovascular Medicine, Section of
Cardiac Pacing and Electrophysiology, Cleveland Clinic Foundation,
Cleveland, OH, USA
Jeffrey J. Hsu, MD, PhD Department of Medicine (Cardiology), University of
California, Los Angeles, CA, USA
Isha Kalia, MS, MPH Division of Cardiology, Columbia University Irving
Medical Center, New York, NY, USA
Jonathan H. Kim, MD, MSc Emory Clinical Cardiovascular Research Institute,
Atlanta, GA, USA
Jared Klein, MD, MPH Cleveland Clinic Children’s, Cleveland, OH, USA
Justine S. Ko, MD Department of Emergency Medicine, Feinberg School of
Medicine, Northwestern University, Chicago, IL, USA
Bradley Lander, MD Division of Cardiology, Columbia University Irving Medical
Center, New York, NY, USA
Farhana Latif, MD Division of Cardiology, Columbia University Irving Medical
Center, New York, NY, USA
Mark S. Link, MD Department of Medicine, Cardiology Division, Cardiac
Electrophysiology, UT Southwestern Medical Center, Dallas, TX, USA
Kyle Mandsager, MD Centennial Heart, TriStar Centennial Heart and Vascular
Center, Nashville, TN, USA
Matthew W. Martinez, MD, FACC Director of Sports Cardiology, Morristown
Medical Center, Atlantic Health System, Morristown, NJ, USA
Silvana Molossi, MD, PhD Department of Pediatric Cardiology, Texas Children’s
Hospital, Baylor College of Medicine, Houston, TX, USA
David C. Peritz, MD Heart and Vascular Center, Dartmouth Hitchcock Medical
Center, Lebanon, NH, USA
Dermot M. Phelan, MD, PhD, FASE, FACC Sports Cardiology Center,
Hypertrophic Cardiomyopathy Center, Atrium Health Sanger Heart & Vascular
Institute, Charlotte, NC, USA
James C. Puffer, MD Division of Sports Medicine, Department of Family
Medicine, David Geffen School of Medicine at UCLA, Los Angeles, CA, USA
Prashant Rao, MBBS, MRCP(UK) Beth Israel Deaconess Medical Center,
Harvard Medical School, Boston, MA, USA
Muredach P. Reilly, MBBCh, MSCE Division of Cardiology, Columbia
University Irving Medical Center, New York, NY, USA
Contributors xi

Thomas Roth, PhD Department of Sleep Medicine, Henry Ford Health System,
Detroit, MI, USA
John J. Ryan, MD Division of Cardiovascular Medicine, Department of Internal
Medicine, University of Utah, Salt Lake City, UT, USA
Elizabeth V. Saarel, MD Division of Pediatric Cardiology, St. Luke’s Health
System, Boise, ID, USA
Nishant P. Shah, MD Department of Cardiovascular Medicine, Cleveland Clinic,
Cleveland, OH, USA
Daichi Shimbo, MD The Columbia Hypertension Center, Columbia University
Irving Medical Center, New York, NY, USA
David Shipon, MD, FACC Thomas Jefferson University Hospital,
Philadelphia, PA, USA
Meeta Singh, MD Department of Sleep Medicine, Thomas Roth Sleep Disorders
Center, Henry Ford Health System, Detroit, MI, USA
Mohita Singh, MD Department of Medicine, Cardiology Division, Cardiac
Electrophysiology, UT Southwestern Medical Center, Dallas, TX, USA
Tamanna K. Singh, MD, FAAC Sports Cardiology Center, Heart, Vascular and
Thoracic Institute, Cleveland Clinic Foundation, Cleveland, OH, USA
John Symanski, MD Sports Cardiology Center, Hypertrophic Cardiomyopathy
Center, Sanger Heart & Vascular Institute, Atrium Health, Charlotte, NC, USA
Marc P. Waase, MD, PhD Division of Cardiology, Columbia University Irving
Medical Center, New York, NY, USA
Shayna Weinshel, BS, MS Department of Medicine, University of Central Florida,
Orlando, FL, USA
Brad Witbrodt, MD Emory Clinical Cardiovascular Research Institute,
Atlanta, GA, USA
Michael Workings, MD Department of Family Medicine, Henry Ford Health
System, Detroit, MI, USA
Kenneth G. Zahka, MD Department of Pediatric Cardiology, Cardiovascular
Medicine, Cleveland Clinic, Cleveland, OH, USA
Chapter 1
The Cardiovascular History
and Examination

John DiFiori, Chad Asplund, and James C. Puffer

Introduction

A comprehensive preparticipation evaluation (PPE) is recommended prior to the

initiation of training and competition for organized sports at the high school level,
the NCAA, professional sports organizations, and most national and international
sport governing bodies [1–6]. While the PPE is felt to be an important first step to
ensuring athlete health and well-being, there is variation in how the PPE is per-
formed among state scholastic programs and even among higher levels of sport
competition [1, 7–10].

The Cardiovascular Component of the PPE

Given that the primary goal of the PPE is to promote the health and safety of the
athlete [1], the cardiovascular (CV) screening portion of the PPE is perhaps the
most essential piece of this assessment. The CV component aims to identify and
evaluate symptoms or exam findings that may lead to the diagnosis of underlying

J. DiFiori (*)
Sports Medicine Institute, Hospital for Special Surgery, New York, NY, USA
e-mail: [email protected]
C. Asplund
Department of Orthopedics and Sports Medicine, Mayo Clinic, Minneapolis, MN, USA
J. C. Puffer
Division of Sports Medicine, Department of Family Medicine, David Geffen School of
Medicine at UCLA, Los Angeles, CA, USA
e-mail: [email protected]

© Springer Nature Switzerland AG 2021 1

D. J. Engel, D. M. Phelan (eds.), Sports Cardiology,
https://doi.org/10.1007/978-3-030-69384-8_1
2 J. DiFiori et al.

cardiac conditions that could result in cardiac morbidity, sudden cardiac arrest, or
sudden cardiac death. The American College of Cardiology and the American Heart
Association state that “the principal objective of screening is to reduce the cardio-
vascular risks associated with organized sports and enhance the safety of athletic
participation; however, raising the suspicion of a cardiac abnormality on a standard
screening examination is only the first tier of recognition, after which subspecialty
referral for further diagnostic testing is generally necessary” [11].
Consensus statements and recommendations for the PPE include specific details
for the cardiac history and physical examination [1, 11–13]. Despite these published
standards, there remains a lack of consistency in their implementation [7–10].
Further, it is important to understand that there is debate about the ability of the CV
history and physical exam to detect significant CV conditions during PPEs.
However, it is well recognized that no screening algorithm is capable of detecting
all clinically relevant cardiac disorders [2, 12]. These important issues are beyond
the scope of this chapter and are discussed in detail in other sections of this
publication.
With these issues in mind, the goal of this chapter is to delineate the key features
of the CV history and physical examination of the PPE.

Organization and Planning

A successful CV screening process is dependent upon planning. Organization

should begin several months in advance. Planning meetings should include team
physicians, team athletic training staff, coaching staff, and administrative staff (e.g.,
staff from the school, athletic department, and/or sport operations). Setting the date
for the PPE with the key stakeholders is the first order of business. The date will
need to consider the timing of the onset of the training and the travel schedules and
availability of the athletes. For PPEs that are intended to be performed for a group
of athletes at a set time, the availability of medical facilities should be confirmed.
Key consultants in cardiology and radiology should be identified and informed of
the dates of the screening, so that their availability for athletes potentially requiring
further evaluation can be established, which will then help expedite the process for
follow-up testing.
The organization process should also include the development of policies regard-
ing issues such as liability coverage (for physicians, athletic trainers, and any other
clinical staff), medical record documentation, and the use of chaperones. If an
online medical history questionnaire is being used, information technology staff
should ensure that the site is secure. Testing of the online process should be per-
formed to identify any technical issues so that they can be resolved in advance.
The personnel needed to perform the CV screening should be identified. In many
cases, especially at the collegiate and professional levels, the CV history and exam
are performed by designated team physicians who are board certified in a primary
specialty and also have completed fellowship training and are certified in sports
1 The Cardiovascular History and Examination 3

medicine. In other situations, it may be ideal for an athlete who has an ongoing
relationship with a personal physician to have that physician perform the screening
examination [1]. This may be the best approach for children and adolescents who
are participating in programs that do not have an identified team physician.
Cardiologists may be used to perform the screening CV history and exam, but they
are more commonly relied upon to evaluate concerning findings. In some cases, a
nurse practitioner or a physician assistant may perform the screening [1]. Regardless
of the certification of the clinician, it is critical that the individual performing the
CV history and exam has had clinical training in this component of the PPE, an
intimate knowledge of the nuances of CV screening in athletes, and the necessary
clinical experience to identify a potential concern in this population.
Once the screening date is set, the athletes (and if minors, their families) should
be notified well in advance. This will allow sufficient time to complete the CV his-
tory (especially if performed online) and obtain any pertinent documents related to
prior screening and/or records involving CV diagnoses and treatment. In cases
where athletes will have the PPE performed by a personal physician or provider, this
provides ample time to arrange the examination.
Finally, the planning should occur with the understanding that the history and/or
physical examination may raise suspicion for the presence of a cardiac condition
that then requires additional evaluation. In such cases, screening events that occur
immediately prior to the planned start of training could result in removal from par-
ticipation while further investigations are performed. To lessen the likelihood of an
athlete needing to be withheld from their training program for sports that have des-
ignated start periods (e.g., high school or collegiate sports), it is recommended that
exams occur several weeks prior to the anticipated start date for that sport. As men-
tioned above, communication with consultants in cardiology and radiology should
take place in advance so that they will be prepared to examine athletes who have had
a concern raised based upon the history and exam.

Setting and Implementation

If the PPE is conducted in a location other than the office of the athlete’s personal
physician, the organizers should arrange a setting that ensures privacy, is comfort-
able for the athletes, and is conducive to maximizing the ability to perform the
examination. For CV screening of groups of athletes from a school, program, or
team, securing the use of patient examination areas in a medical facility is ideal. An
individual exam room is preferred for reasons of privacy and the ability to have
quiet space for auscultation. The use of gymnasiums, auditoriums, locker rooms,
and other non-private areas is not recommended. Attempting to create a level of
separation within a large room by using a “pipe and drape” setup is likewise not
recommended.
In order to conduct a thorough exam, an appropriate amount of time should be
allocated for each athlete being screened. The amount of time needed to conduct an
4 J. DiFiori et al.

exam for an individual athlete, the total number of athletes needing to be screened
in a given time period, and the number of available examination rooms should be
determined in advance. This will indicate the number of examiners needed and the
total time required to perform a complete CV history and exam for a group of
athletes.
Other factors to consider include whether an online questionnaire was completed
in advance or a hard copy was completed on site. Online questionnaires must be
completed on a secure website, and then viewed within an electronic medical record,
or uploaded or printed and scanned to become part of the athlete’s official medical
record. If an online questionnaire or hard copy is to be completed on site, a private
space should be provided for the athlete to complete the document. In either case,
the athlete (or parent/guardian) must sign and date the questionnaire attesting to its
accuracy.
Although uncommon, an athlete (or his parent/guardian) may withhold or mis-
represent important medical information due to a concern that providing such infor-
mation could jeopardize medical clearance for sport participation. Thus, it is
important that the physician confirm that the acknowledgment is signed. In some
cases, an athlete may view the history and exam as an unnecessary burden or “rub-
ber stamp” process prior to the beginning of training. In these circumstances, an
athlete may choose to select negative responses throughout the questionnaire in
order to expedite the screening. This leads to substandard screening that could place
the athlete at risk. In order to recognize if an athlete is not reading and responding
to each question individually, and simply checking the “no column,” it may be help-
ful to embed a question that requires a positive response. An example of such a
question is “have you ever played a competitive sport?” Should the clinician feel
that the athlete is providing inaccurate information, they should proceed to perform
the history using a primary “interview” format, asking each question and clarifying
each response verbally.

Personal and Family History

A detailed history and physical examination have been the cornerstones of the pre-
participation evaluation of athletes in the United States for decades. However, given
the high degree of variability and lack of standardization of cardiovascular assess-
ment, the American Heart Association (AHA) convened an expert panel in 1996 to
make recommendations for a standardized process for this component of the prepar-
ticipation evaluation [14] with an updated review of the recommendations in 2007
and 2014 [2, 11]. The result of this work was the development of a 14-point evalu-
ation, which has now been widely embraced for the cardiovascular preparticipation
screening of athletes (Table 1.1).
Perhaps the most important component of this 14-point evaluation is the personal
and family history, since athletes with underlying yet undetected cardiovascular
disease may present with warning signs (e.g., syncope or chest pain during exercise)
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the inside of the shell, parallel with the rows of ambulacral pores. Lign.
103, fig. 3, is a remarkable example, in which flint occupies the base
of the shell, while the upper surface is lined with crystals of calcareous
spar.

Lign. 107. Micraster (Spatangus) cor-anguinum.

Chalk. Lewes.
Fig. 1. View from above, showing the petaloid
— ambulacra.
2.
View of the base, with the mouth.
—

Micraster cor-anguinum (Snake-heart). Lign. 107.—Of this genus

there are many species in the Chalk. This type of Spatangidæ are
more or less oval, elongated, and heart-shaped, wider before than
behind, with a sulcus, or furrow, in front. The shell is fragile, and
composed of large polygonal plates; the tubercles small and irregularly
distributed; the spines are short. The mouth is transverse, situated
anteriorly, and protected by a strong projection of the odd
interambulacrum, which is named the lip. The vent is terminal, and
placed above the margin. There are but four ambulacra, and these are
incomplete, comparatively of small extent, and situated in deep
furrows. A large and new species of Micraster (M. cor-bovis, of Prof. E.
Forbes), from the Sussex Chalk, is figured in Dixon's Fossils, pl. xxiv.
figs. 3, 4, p. 342.
TOXASTER COMPLANATU

Lign. 108. Toxaster complanatus.

Greensand. Switzerland.
Fig. 1.
Profile.
—
2.
View of the summit, showing the Vent at the side; e.
—
3. View of the base, displaying the situation of the mouth,
— and the union of the five ambulacra; their pores are not
introduced in figs. 2 and 3.
a. The narrow porous divisions of the shell, termed
Ambulacra.
b. Interambulacral spaces.
c. Areæ, or spaces covered by the wide plates.
d. The Mouth.
e. The Vent, or Outlet.
 Toxaster (Spatangus) complanatus. Lign. 108.—In this form of
Spatangus (constituting the genus Toxaster Agass.), the ambulacra are
not depressed or furrowed, as in the preceding echinites, nor petaloid
(leaf-shaped), as in those which M. Agassiz denominates true
Spatangi, but converge to a point on the summit, as is shown in fig. 2;
the external rows of pores are elongated horizontally, and form a kind
of furrow. The odd ambulacrum is in a deep groove. The mouth is
transverse, fig. 3, d; and at the anterior part of the inferior face there
is a depression, which results from the convergence of the ambulacral
areæ towards that point. The vent is in the posterior face. This species
is from the Neocomian strata of France; I introduce it to illustrate the
characters of several other echinites, which the French geologists
suppose to be confined to the so-called Neocomian formation; but
which also occur in the Upper Greensand of Blackdown.
Holaster is another genus of Spatangidæ established by M. Agassiz,
for those echinites that are heart-shaped, with simple ambulacra
converging towards the summit. The mouth is elongated transversely;
the outlet is on the posterior face. A specimen first described in my
Foss. S. D. (pl. xvii. fig. 9, 21), as Spatangus planus, is common in the
Lower Chalk, and Chalk-marl, and abundant in the Firestone Malm-
rock.

Our limits will not allow of a more extended notice of the fossil
Echinidæ. The student should consult the Memoirs on the genera, now
in course of publication at the Government School of Mines, by
Professor Edward Forbes; the plates are exquisite, as works of art, and
the descriptions all that can be desired.
Mr. Dixon's work contains three excellent plates of cretaceous
Echinites. Several chalk species are figured in my Foss. South Down.
The numerous coloured figures of fossil sea-urchins in the Pictorial
Atlas of Organic Remains, have already been mentioned.
Geological Distribution of Echinites.—No vestiges of this order of
radiata have been discovered in the Silurian deposits: the earliest
known occurrence of any type is in the Carboniferous formation. The
most ancient Echinidæ, according to the present state of our
knowledge, are the Cidares, in the modified form previously noticed,—
Archæocidaridæ (ante, p. 322.).
In the Trias another type appears, Hemicidaris, which holds an
intermediate place between the Cidarites properly so called and the
Diadema.
In the Oolite, and Jurassic formations, numerous forms are for the
first time met with, constituting the genera Echinus, Clypeus, Disaster,
Holectypus, Diadema, Nucleolites, &c.
The Cretaceous seas swarmed with echini belonging to genera of
which no traces have been found in earlier rocks; viz. Holaster, Salenia,
Micraster, Salerites, Discoidea, Ananchytes, Cassidulus, &c.
In the Tertiary formations, Spatangus, Scutella, Clypeaster, and
other new genera appear, and many of the ancient ones are absent; or
at least have not been observed. Of the genera printed above in italics,
no living species are known.
On collecting and developing fossil ON COLLECTING FOSSIL ECHINODERMAT

Echinodermata.—In the previous remarks on the fossil remains of

radiated animals, we have pointed out those remains that are the most
important and instructive, and should be sought for by the student.
Thus, in collecting Crinoids, the receptacle or body should be the
principal object of research, and if only detached plates can be
extracted from the rock, their relative position should be carefully
noted, and the specimens glued to a card or board, in their natural
order; and some of the ossicula of the column, and of the arms, or
tentacula, be placed with them.
Mr. Miller dissected specimens of every genus, and has figured the
separate plates or bones that enter into the composition of the
receptacle,[302] and arms. Traces of the tentacula, and their
subdivisions, must be sought for, and if discovered, should be removed
with the stone to which they are attached, and the block be afterwards
reduced in size by a mason's saw, and not by the blows of a hammer,
which might displace the ossicula.
 [302] Natural History of the Crinoidea.

If imbedded in Lias shale, or other fragile material, a thick slab

should be removed, for greater safety in conveyance; this, when
reduced to a convenient size and thinness, may be imbedded in a tray
with plaster of Paris, or glued to a piece of thin, well-seasoned
mahogany, or deal. The specimens of the Pear Encrinite of Bradford,
and of the Pentacrinites from Lyme Regis, in the British Museum,[303]
were prepared in this manner.
[303] Petrifactions, p. 78.

The crinoideal remains in Chalk belong but to few genera; they

merely require the usual manipulation of cretaceous fossils. The
collector, however, should remember that the ossicula and plates of the
receptacle (as for instance of the Marsupites), are but slightly adherent
to each other, and the chalk must not be wholly removed, or these
parts will become detached.
The receptacles of the Apiocrinites of the chalk are rarely found
with more than a few joints of the column attached; and I believe no
vestiges of the arms have been observed: these parts are therefore
desiderata, and should be diligently sought for: the radicles of these
crinoideans are long, articulated, and branching, and without due
caution may be mistaken for the arms, or for another species. The first
remark will also apply to the Marsupite; any specimens with but a few
ossicula of the arms are very precious. I may observe that there is yet
much to learn as to the number of species and genera, and the
peculiar characters of the Crinoidea of the chalk, and that any addition
to our knowledge on this subject will be valuable.
The Asteridæ are so simple in form and structure, that it is
unnecessary to offer any suggestions for their development; of course
they must not be removed from the stone.
Among the detached ossicula dispersed through the chalk, the
student will remember that the large madrepore-like tubercle of the
Star-fishes, (ante, p. 304.,) may often occur. It may easily be mistaken
for an encrinital body, or for a coral, but an accurate inspection will
show that it is not composed of anchylosed plates, like the receptacle
of an Apiocrinite, but has surfaces for attachment to other ossicula;
while the ends, which in a crinoideal column would have radiated
surfaces, are rounded and entire.
In collecting Echinites, much caution is required in dissecting
specimens surrounded by spines. If imbedded in hard limestone, or in
laminated clay, it is scarcely possible to preserve the spines in
connexion with the shell; but it often happens that the Cidarites of the
Oolite are attached by the base to the solid limestone, and the case
with the spines is imbedded in sandy, friable aggregate, not difficult of
removal. A specimen in my cabinet, with upwards of fifty spines
attached to the shell, was obtained under such circumstances.[304]
[304] Now in the British Museum.

The Chalk Echinites will be found to possess spines more frequently

than is commonly supposed, if care be taken to explore the
surrounding chalk before it be removed. T have often procured
Cidarites with spines, when there were no apparent vestiges of these
appendages, by carefully scraping away the surrounding mass until the
extremity of a spine appeared, and then tracing it up to its connexion
with the shell; another point was discovered by further removal, and
that was developed in the same manner; and at length a Cidaris with
several spines was obtained. The chalk around the mouth should
always be cautiously removed in the dentated species, in the hope of
preserving the teeth, as in the specimens, Lign. 101, fig. 1, and Lign.
104, fig. 1.
As the shells of Chalk Echinites, when hollow, are often lined with
crystals (see Lign. 103, fig. 3), it is worth while to break all indifferent
specimens of the common species, with the chance of obtaining an
example of this kind.
The chalk must not be scraped off from the crust or shell of the
Echinites, or the minute granulations and papillæ will be injured or
removed; it should be flaked off with a blunt point.
 In friable arenaceous strata, as in some of the Maestricht and
Tertiary deposits, the Echinites may be extricated in as perfect a
condition as if fresh from the sea; it is, indeed, probable, from the
habit of these animals of burrowing in mud and sand, that in many
instances they were entombed alive by the sediment in which their
fossil remains are imbedded.
Beautiful Cidarites and their spines may be collected in the Oolitic
strata at Calne, Chippenham, Bath, &c.; and in the coralline Oolite near
Faringdon; and of Saleniæ, in the Greensand gravel-pits near that
town.
The Upper Greensand near Warminster, and at Chute Farm, near
Heytesbury, abounds in small Cidaritidæ and other echinites. The large
sinuated Clypeus is found in great perfection in the Oolite at Malton,
Cheltenham, Gloucester, &c.
The cretaceous echinites are to be met with in most localities of the
white chalk. The chalk-pits in Kent, especially at Gravesend, Northfleet,
Chatham, &c. are rich in Cidarites, and their spines. The Galerites, and
Ananchytes, are also very fine and numerous; and the softness of the
chalk renders their extrication from the stone a delightful task for the
young geologist.
Specimens of the common kinds of fossil Echinoderms may be
obtained at moderate prices of the dealers named in the Appendix.
There is a matchless suite of fossil Echinidæ in the British Museum,
which has been arranged and named by Mr. S. P. Woodward, and is
now the most instructive and interesting collection extant. It contains
examples of the following genera, viz., Ananchytes, Echinocorys,
Echinolampas, Holaster, Galerites, Cidaris, Diadema, Acrosalenia,
Glypticus, Disaster, Pygurus, Clypeaster, Scutella, Salmasis,
Echinocyanus, &c. There is also a good series of echinital spines.
 CHAPTER X.
FOSSIL FORAMINIFERA—MICROSCOPICAL
EXAMINATION OF CHALK AND FLINT.

"Where is the dust that has not been alive?"

Young.

That those infinitesimal forms of animal existence which swarm

throughout the waters of the ocean, but whose presence can only be
made manifest by the aid of the microscope, are preserved in a fossil
state,—that their durable remains constitute mountain ranges, and
form the subsoil of extensive regions,—and that the most stupendous
monuments erected by man are constructed of the petrified relics of
beings invisible to the unassisted eye,—are facts not the least
astounding of those which modern Geology has revealed.
This interesting field of research, which the labours of that eminent
observer M. Ehrenberg first made known, has since been explored by
other naturalists, and in every part of the world many of the Tertiary
and Secondary deposits have been found to contain microscopic
organisms in profusion. At present this branch of palæontology is in its
infancy, and it offers to the young student an inexhaustible and most
attractive path of scientific investigation; it possesses, too, this great
advantage over many others, that it can be pursued at home, and the
materials for its prosecution are everywhere at hand. Unlike my
explorations in the Wealden, in which a few fragments of bones, or
teeth, scattered at wide intervals through the rocks, and in localities
many miles apart, were often the only reward of a day's labour, here,
in the quiet of my study, I may discover in a few atoms of flint, or
grains of chalk, picked up by the road-side, the fossil remains of beings
as interesting and extraordinary as the extinct colossal reptiles of
Tilgate Forest.
The microphytes, or fossil Diatomaceæ, described in a MICROPHYTE
previous chapter, (ante, p. 93.,) were formerly classed with the
organisms that now claim our attention, under the name Infusoria;
from the belief that generally prevailed among naturalists, of their
animal origin. In fact, some eminent microscopic observers, while
admitting the vegetable character of Xanthidium, Micrasterias, &c.
consider the Naviculæ, Ennotiæ, &c. as belonging to the animal
kingdom.
Thus Dr. J, W. Bailey, in a late "Memoir on the Microscopic
Organisms in Various Localities of the United States," divides these
bodies into three groups; viz. Infusoria, Desmidieæ, and Diatomaceæ;
with the remark, that he has separated the two latter tribes from the
Infusoria, because so many distinguished naturalists consider them
decidedly to belong to the vegetable kingdom: "but," he adds, "while I
believe that no positive line of separation can be drawn between
certain animals and vegetables, I am yet disposed to regard the
Desmidieæ, from the sum of all their characters, as most nearly allied
to admitted vegetables; while the Diatomaceæ, notwithstanding Mr.
Thwaites's interesting observations on their conjunction,[305] still seem
to me, as they have always done, to be true animals. There is such
apparent volition in their movements, such an abundance of nitrogen
in the composition of their soft parts, and such resemblance between
the stipitate Gomphonematæ and some of the Vorticellæ, that I should
be still disposed to class them as animals, even if Ehrenberg's
observations of the retractile threads and snail-like feet of some of the
Naviculæ should not be confirmed."[306]
[305] The mode of fructification, or conjunction, as it is termed,
in the Algæ, consists in the adhesion of two cells or frustules
together, and their fusion into one; from their united contents a
mass of granular substance is produced, that becomes consolidated
and forms the spore or fruit, which, when arrived at maturity, is set
free by the bursting of the cell. Mr. Thwaites has ascertained that
the fructification is similar in the Diatomaceæ.
 [306] Smithsonian Contributions, vol. ii. p. 34.

Thus, whilst referring Closterium, Arthrodesma, Euastrum,

Xanthidium, Micrasterium, &c. as vegetables, to the Desmidieæ, Dr.
Bailey places, Actinocyclus, Campilodiscus, Coscinodiscus, Ennotia,
Navicula, Gomphonema, Pinnularia, Triceratium, &c. among the
Diatomaceæ, as animals.
Of the animal nature of the microscopic objects which now require
our attention, there is however no question, although the zoological
position and affinities of many of the organisms included in this survey
are still but imperfectly determined.

The animals designated Foraminifera,[307] or Rhizopoda,[308] are of

a more simple structure than the Polypifera and Echinodermata
described in the previous chapters; yet as their relics are for the most
part presented to the notice of the geologist as aggregations of shells,
forming extensive beds of limestone, it will be convenient to treat of
them in this place.
[307] Foraminifera, i.e. bearing foramina—a name derived from
the minute openings in the shells and their septa.
[308] Rhizopoda; root-like feet; from the long fibrous processes,
or pseudopodia.

The fact that the fossil remains of Foraminifera, and of Mollusca,

alike consist of shells, and constitute strata identical in mineral
characters, and deposited under like physical conditions, renders the
examination of these Microzoa[309] a fit introduction to the study of
the durable remains of the higher order—the Mollusca.
[309] Microzoa; from μικρος, mikros, small, and ζωον, zoon,
animal; a convenient term to denote minute animal organisms
whose forms can only be defined by the aid of the microscope.

It is scarcely more than a hundred and twenty years ago, that the
existence of this numerous order of microzoa was first made known to
naturalists by Beccarius, who detected a considerable number of
species in the sand on the shores of the Adriatic. But the structure of
the animals that secreted these shells is a discovery of comparatively
modern date. The early collectors classed these microscopic bodies
with the shells of true mollusca; and even M. D'Orbigny, whose
elaborate researches justly constitute him a high authority in this
branch of natural history, in his first memoir, in 1825, described the
involuted discoidal forms as Cephalopoda. This error was corrected by
the investigations of M. Dujardin, who in 1835 satisfactorily
demonstrated that the Foraminifera are animals of the most simple
structure, and entirely separated by their organization from the
Mollusca.
But the true nature of this class is so little understood by British
collectors of fossil shells,—of course I mean the uninitiated, and the
amateur naturalist, for whose use these unpretending pages are
designed,—that in order to invest the study of the fossil species with
the interest which a knowledge of the structure and economy of the
living originals can alone impart, I must give a history of the recent
forms somewhat in detail, taking M. D'Orbigny as my chief authority.
[310]

[310] The best work for the student to consult is M. D'Orbigny's

"Foraminifères Fossiles du Basse Tertiaire de Vienne, Autriche."
Paris, 1846. 1 vol. 4to. with plates. I rejoice to learn that a
Monograph on the British Foraminifera is in preparation by Dr.
Carpenter and Professor Williamson: than whom there are none
more competent.

The Foraminifera are marine animals of low organization, FORAMINIFER

and, with but few exceptions, extremely minute: in an ounce of sea-
sand between three and four millions have been distinctly enumerated.
When living, they are not aggregated, but always individually distinct;
they are composed of a body, or vital mass, of a gelatinous
consistence, which is either entire and round, or divided into
segments, placed either on a simple or alternate line, or coiled in a
spiral, or involuted round an axis. This body is covered with an
envelope or shell, which is generally testaceous, rarely cartilaginous, is
modelled on the segments, and follows all the modifications of form
and contour of the body. From the extremity of the last segment there
issue sometimes from one, sometimes from several openings of the
shell, or through the numerous pores or foramina, very elongated,
slender, contractile, colourless, filaments, more or less divided and
ramified, serving for prehension, and capable of entirely investing the
shell.
The body varies in colour, but is always identical in individuals of
the same species; it is yellow, ochreous, red, violet, blue, &c.
Its consistence is variable; it is composed of minute globules, the
aggregation of which determines the general tint. It is sometimes
entire, round, and without segments, as in Gromia, Orbulina, &c.,
which represent at all ages the embryonic state of all the other genera.
The animal increases by gemmation, each segment being essentially
distinct, but connected with the preceding one by a tube or neck.
When the body is divided by lobes or segments, the primary lobe, as in
the permanent condition of the Gromia, is at first round or oval,
according to the genus; once formed it never enlarges, but is
enveloped externally by testaceous matter.
The segments, which successively appear, are agglomerated
together in seven different ways, and these modifications are the basis
of M. D'Orbigny's classification. The discoidal forms, as the Rotalia,
Rosalina, Cristellaria, &c. are involute, like the Nautilus, and divided by
septa or partitions, which, like the enveloping shell, are perforated.
The lobes of the body occupy contemporaneously every chamber, and
are connected by a tube that extends through the entire series. In the
spiral form, as the Textularia, &c., the same structure is apparent.
Whatever the form of the body, the filaments always consist of a
colourless transparent matter; they are capable of being elongated to
six times the diameter of the shell. They often divide and subdivide, so
as to appear branched; and though alike in form in the different
genera, vary much in their position. In some species they form a
bundle which issues from a single aperture, and is withdrawn into the
same by contraction; in others, the filaments project only through each
of the pores in that portion of the shell which covers the last segment:
in many they issue from both the large aperture and the foramina.
These filaments or pseudopodia fulfil in these animals the functions of
the numerous tentacula in the Star-fishes; serving as instruments of
locomotion and attachment.
Neither organs of nutriment nor of reproduction have been
discovered. In the genera having one large aperture from which the
filaments issue and retract, we can conceive nutriment to be absorbed
by that opening; but this cannot be the case in the species which have
the last cell closed up; in these the filaments issuing through the
foramina are probably also organs of nutrition. M. D'Orbigny considers
the Foraminifera as constituting a distinct class in zoology; though less
complicated than the Echinoderms and the Polypifera in their internal
organization, they have the mode of locomotion of the first; while by
their free, individual existence, they are more advanced in the scale of
being than the aggregated and immovably fixed animals of the latter
class.
But though I consider the animal of the Foraminifera as single, and
the additional lobes, or segments, as the continuous growth of the
same individual, I must state that some eminent naturalists regard the
entire structure as a series of distinct individuals, developed by
gemmation from the first formed segment, like the clusters of the
compound Tunicata; and not as a single aggregated organism, made
up of an assemblage of similar parts indefinitely repeated. In a
palæontological point of view, it matters not which opinion is adopted.
[311]

[311] See a masterly paper on the structure of Nummulina and

Orbitoides, by Dr. Carpenter; Geol. Journal, vol. vi. pp. 21-39, with
admirable representations of the structural details.
 Lign. 109. Foraminifera.
Chalk. Charing.
Fig. 1. Globigerina cretacea;
— the original is 1/60 of an inch in
diameter.
2.
Textularia Globulosa; 1/40.
—
3.
Verneuilina tricarinata; 1/30.
—
4.
Cristellaria rotulata; 1/40.
—
5.
Rosalina Lorneiana; 1/40.
—

Classification of the Foraminifera.—The CLASSIFICATION OF THE FORAMINIFER

number of genera is so great, that I can only attempt to convey a very
general idea of the principles of classification adopted by M. D'Orbigny,
and give a few illustrations of some of the most abundant fossil
species.
That the reader may be cognizant of the usual aspect of these
shells five specimens from the Chalk, belonging to as many genera,
are here represented (Lign. 109); the deposit whence they were
obtained will be described hereafter.
As the mode in which the growth of the body, and consequently of
the shell, takes place, differs greatly in certain groups, an obvious and
natural arrangement is suggested, by which the class is divided into
seven orders:—
1. The primary, or simplest type; one segment or cell; as in
Orbulina.
2. The segments arranged in circular lines; as in Orbitolina.
3. Segments disposed in a straight or arched single line;
successively increasing from the first to the last cell; as in Nodosaria,
Lign. 111.
4. Segments, spirally and discoidally disposed, on the same plane,
like cells of the Nautilus; as in Cristellaria, Lign. 109.
The same type, but coiled obliquely, and inequilaterally, like the
shells of Gasteropoda; as in Globigerina, Lign. 109, Rosalina, Lign.
109.
5. Segments developed alternately on the right and left of the first,
and successively on each side the longitudinal axis; as in Textularia,
Lign. 109.
6. Arrangement of the segments combining the modes of 4 and 5;
that is, the segments are formed alternately, but the whole are coiled
spirally, either obliquely, or on the same plane; as in Amphistigena.
7. Segments round a common axis, on two, three, four, or five,
opposite faces, returning after each entire revolution; the new cells
being placed exactly on the preceding series; as in Quinqueloculina,
Pict. Atlas, pl. lxii. fig. 12.
The Foraminifera vary considerably in magnitude; by far the greater
number of species are invisible to the unassisted eye, and the aid of a
lens or microscope is required to define the structure even of the
largest; yet many are of sufficient size to be recognized, as for
example the Spirolinæ, Lign. 112. A few genera are from a quarter of
an inch to nearly an inch in diameter, as the Orbitoides Mantelli of the
tertiary formations of North America,[312] and the Nummulina,
commonly termed Nummulites, of Europe, Lign. 110.
 We will now describe the genera selected for illustration,
commencing with the large and well-known type, whose aggregated
remains form extensive beds of crystalline limestone in the Alps, and in
Asia, and Africa.[313]
[312] Dr. Morton's Synopsis of the Organic Remains of the
Cretaceous group of the United States. Philadelphia, 1824, p. 45, pl.
v. fig. 9.
[313] Geol. Journal, vol. v.

Lign. 110. Nummulites, or Nummulina; nat.

From the Great Pyramid of Egypt.
Fig. 1.— Transverse section of a Nummulite,
showing the
form and arrangement of the cells.
2, 3. Specimens with part of the external
— plate removed.

Nummulina lævigata. Lign. 110.—The shell is of a discoidal or

lenticular form, composed of numerous cells, concentrically arranged
round an axis on the same plane; both sides of the disk covered by a
smooth thick plate.
Under the name of Nummulites, from their resemblance NUMMULITE
to a piece of money, the fossil shells of this genus of Foraminifera have
long been known to naturalists, and are figured in many of the early
works on petrifactions. They occur in immense quantities in certain
rocks, and are of all sizes, from a mere point, to disks an inch and a
half in diameter; thus exceeding in magnitude all other animals of this
class.
Perfect specimens appear as a calcareous solid circular body, of a
lenticular shape; smooth, and slightly convex on both sides, and
without any visible structure. On splitting the fossil transversely, or
rubbing down one of the convex planes, a series of minute cells,
arranged in a discoidal spire, is brought to view, as shown in Lign. 110,
fig. 1. But this description gives a very inadequate idea of the
complicated and exquisite structure of the original, which has been
admirably worked out by Dr. Carpenter. This eminent physiologist has
shown that each chamber was occupied by a living segment,
connected with other segments by one or more tubular prolongations,
which absorbed nutrition from without, by means of filamentous
pseudopodia, that projected through a system of passages leading
from the medial plane to the external surface.[314] A figure of the
supposed form of a living Nummulina is given in Pict. Atlas, p. 187.
[314] Geol. Journal, vol. vi. p. 21. See also a paper by Prof.
Williamson, "On the minute Structure of the Calcareous Shells of
some recent species of Foraminifera." Trans. Microscop. Soc. vol. iii.
p. 105.

The specimens figured in Lign. 110 are from the limestone that
forms the foundation rock of the Great Pyramid of Egypt, and of which
that structure is in great part composed. Strabo alludes to the
Nummulites of the Pyramids, as lentils which had been scattered about
by the artificers employed on those stupendous monuments, and
become stone.[315] Silicified masses of Nummulites are occasionally
met with; polished slices of such specimens are richly figured by the
sections of the inclosed Foraminifera.
[315] An interesting fact was communicated to me by a friend
who lately descended the Nile; the Nummulitic limestone rocks are
in some parts of the course of the river washed into the stream, and
becoming disintegrated, the Nummulites are set free, and re-
deposited in the recent mud of the Nile.
 The Nummulitic limestones are of the Eocene or ancient. Tertiary
epoch, as the labours of Sir Roderick Murchison in the Alps, Apennines,
and Carpathians first established: Nummulites are unknown in the
Secondary formations.[316]
[316] Geol. Journal, vol. v.

Orbitoides.—The fossil bodies thus named are disciform, like the

Nummulites; and one species, which forms the constituent substance
of ranges of limestone mountains, 300 feet high, near Suggsville, in
North America, was first described by my lamented friend, the late Dr.
Morton, of Philadelphia, as N. Mantelli, in his work on the Cretaceous
Fossils of the United States.
The discovery that the Nummulitic deposits of the Old World were
of the tertiary period, directed attention to their supposed geological
equivalents in America; and on a careful examination of their fossils,
the rocks proved to be tertiary, and the shells true Foraminifera allied
to the Nummulites, but generically distinct.[317] A reference to Dr.
Carpenters memoir, previously cited, is necessary to comprehend the
complicated structure of these fossils.[318]
[317] Manual of Geology, p. 208.
[318] Geol. Journal, vol. vi.

Siderolina, or Siderolites, is a genus of Foraminifera, which may be

described as Nummulites, in which the turns of the spire are
intercepted by elongated appendages, that project beyond the
periphery of the disk, and produce a stellated figure. These fossils
abound in the cretaceous strata of Maestricht.
Fusulina.—The shell is fusiform, being elongated transversely to the
axis; the cells are divided internally by constrictions. Only one species
is known, (F. cylindrica,) and this is confined to the Carboniferous
formation; it is the most ancient or earliest type of the class, according
to the present state of our knowledge.
 Lign. 111. Foraminifera, &c.
Fig. 1 and 2. Campilodiscus; a perfect frustule, and
— part of another, seen from above,
Tertiary; Egra in Bohemia.
See ante, p. 96.
3a.— Lituola nautilotdea. Chalk, Chichester.
(By Mr. Walter Mantell.) Side view, × 8.
3b.— Front view of the last cell of 3d, to show
the foramina with which it is pierced.
3c.— The last cell of 3a.
3d.— Side view of a young shell, before the
produced, or straight part appears, ×
20.
4a.— Flabellina Baudonina. Chalk. (M.
D'Orbigny.) A young individual seen in
profile, × 12.
4b.— The same, viewed laterally, shows the
oblique arrangement of the cells.
5.— Nodosaria. Chalk, Chichester. (By Mr.
Walter Mantell.) The line below indicates
the natural size.

NODOSARIA. LITUOL
 Nodosaria. Lign. 111, fig. 5.—Straight, elongated cells placed end to
end, separated by constrictions; the last formed cell has a round
central aperture. Several beautiful species abound in the Chalk;
specimens often occur adhering to the surface of the flint nodules.
Nodosariæ are abundant in tertiary deposits. Mr. Walter Mantell
discovered Foraminifera of this genus in the blue clay of Kakaunui, in
New Zealand.
Cristellaria. Lign. 109, fig. 4.—The shell is in the form of a
compressed Nautilus; it has a single aperture, which is situated at the
angle of the keel; the cells are oblique.
This genus comprises seventy fossil species, which occur in the
Lias, Oolite, and Chalk. Living species are abundant in almost every
sea.
Flabellina. Lign. 111, fig. 4a.—In a young state, this shell, like the
preceding, resembles that of a Nautilus, and the cells are oblique; but
in the adult, are of a zigzag (chevron) form. It has a single round
aperture. Fig. 4a shows a young individual, seen in profile; fig. 4b a
lateral view, exhibiting the obliquity of the cells.
Species of Flabellina are often found in cretaceous strata. The
genus is not known in a recent state.
Polystomella.—In its general form this genus resembles the other
nautiloid shells above described, but its structure differs essentially; for
there are several apertures along the side of the shell, as well as the
opening in the last segment. The cells are simple, and each is a single
cavity. The figures and details of structure, given by Professor
Williamson, must be referred to, for an insight into the organization of
this beautiful and complicated type of Foraminifera.[319] One species of
Polystomella is said to occur in the Chalk; I have not detected this
genus in our cretaceous deposits. Recent species swarm in our seas;
and may be easily obtained from the mud and sand on the shores at
Brighton.[320]
[319] Trans. Microscop. Society, vol. ii.
 [320] Mr. Poulton has specimens of the shells, and the bodies of
the animals deprived of the shell, mounted for the microscope.

Lituola. Lign. 111, fig. 3a.—In a young state the shell is nautiloid,
as in fig. 3b, 3d; but becomes produced by age, and assumes a
crosier-like form, as in fig. 3a. The cells are filled with a porous
testaceous tissue, as shown in figs. 3b, 3c; which also illustrate the
foraminiferous character of the shells in this class of animals; for both
the external testaceous covering, and the septa of the cells, exhibit
perforations.[321]
[321] The perforations are omitted, by mistake, in the figure of
the adult shell, fig. 3a.
 Lign. 112. Spirolinites in Flint.
Chalk. Sussex.
(By the late Marquis of Northampton.)
The specimens are only sections of the shells
Imbedded in flint, and seen as opaque objects with a
lens of moderate power. The small figures denote the
size of the originals.

Spirolina. Lign. 112.—The general form resembles that of SPIROLIN

Lituola: the young shell being a discoidal, involute, and becoming
produced by age; but the internal structure is different; the cells are
simple cavities.
The chalk and flints of Sussex abound in these crosier-like shells,
whose existence in the cretaceous rocks was first made known by my
deeply lamented friend, the late Marquis of Northampton. The annexed
lignograph, from drawings by his lordship, shows the form and
structure as displayed by sections in fractured flints. Four species were
named by Lord Northampton (see Wond. p. 32-5); but it is doubtful
whether all the specimens belong to more than one species; the
apparent diversity of structure may arise simply from the different
planes in which the sections happen to have been made.
Globigerina. Lign. 109, fig. 1.—The shell is turbinated, the cells are
spheroidal, and the last, or terminal one, has a semilunar aperture at
the umbilical angle. Several fossil species abound in the Chalk and in
the tertiary deposits; and many living species swarm in our seas.

Lign. 113. Nonionina Germanica, × 290.

A recent specimen of the body of the
animal; the shell removed by acids.
From the North Sea, Cuxhaven. (M.
Ehrenberg.)
a, a.—Naviculæ and other organisms in
the segments of the animal.

Nonionina.—Lign. 113.—A nautiloid shell, with simple cells; the last

cell has a single narrow aperture placed transversely over the dorsal
aspect of the spire. One species occurs in the chalk formation of
Germany; several in tertiary deposits, and in the existing seas.
The figure, Lign. 113, represents the body of the animal deprived
of its shell, to illustrate the nature of certain fossils from the Chalk.
 Lign. 114. Foraminifera in Chalk and Flint. × ×.
(Seen by transmitted light.)
Fig. 1, 2, 3, 6.
Different forms of Rotalia.
—
2.— Resembles the recent Rotalia stigma;
Ehrenberg;> from the North Sea,
near Cuxhaven.
4.— Portion of a Nautilus, showing five
chambers, partially separated, each
pierced by the siphunculus: in Flint,
from
Ireland.
5.— The body of a Rotalia, in Flint; the
shell is not apparent.

Rotalia. Lign. 114.—The shell, though nautiloid in its ROTALIA—ROSALIN

contour, is regularly turbinated, the cells not globular; the last cell has
a central, semilunar, transverse, aperture. There are fifty fossil species.
The Rotaliæ appear in the Lias, Oolite, and Chalk, in immense
numbers, and swarm in the present seas.
Rosalina. Lign. 109, fig. 5.—The shell is depressed; the spire
apparent on one side; the aperture is a prolonged slit extending from
one cell to another, and opening on the umbilicus; that is, on the side
opposite to the spire. There are eighteen fossil, and many recent
species of this genus.
Textularia. Lign. 109.—This, and the following genus, belong to that
order of Foraminifera in which the segments or cells are arranged in
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