Table Of ContentEvolutionary Biology
of Primitive Fishes
Evolutionary Biology
of Primitive Fishes
Edited by
R. E. Foreman
Bamfield Marine Station
Bamfield, British Columbia, Canada
A. Gorbman
University of Washington
Seattle, Washington
J. M. Dodd
University College of North Wales
Bangor, United Kingdom
and
R. Olsson
University of Stockholm
Stockholm, Sweden
Plenum Press
New York and London
Published in cooperation with NATO Scientific Affairs Division
Proceedings of a NATO Advanced Resean:h Workshop on
Evolutionary Biology of Primitive Fishes,
held April 14-17, 1985,
at Bamfield Marine Station, Bamfield, British Columbia, Canada,
co-sponsored by the Natural Sciences
and Engineering Research Council of Canada
and the Western Canadian Universities
Marine Biological Society
Library of Congress Cataloging in Publication Data
NATO Advanced Research Workshop on Evolutionary Biology of Primitive Fishes
(1985: Bamfield, B.C.)
Evolutionary biology of primitive fishes.
(NATO ASI series. Series A, Life sciences; v. 103)
"Proceedings of a NATO Advanced Research Workshop on Evolutionary
Biology of Primitive Fishes, held April 14-17, 1985, at Bamfield Marine Station,
Bamfield, British Columbia, Canada"-T.p. verso.
"Published in cooperation with NATO SCientific Affairs Division."
Includes bibliographies and index.
1. Fishes-Evolution-Congresses. 2. Fishes, Fossil-Congresses. I.
Foreman, R. E. (Ronald Eugene), date. II. North Atlantic Treaty Organization.
Scientific Affairs Division. III. Title. IV. Series.
QL618.2.N37 1985 597'.038 85-31168
ISBN-13: 978-1-4615-9455-0 e-ISBN-13: 978-1-4615-9453-6
001: 10.1007/978-1-4615-9453-6
© 1985 Plenum Press, New York
Softcover reprint of the hardcover I st edition 1985
A Division of Plenum Publishing Corporation
233 Spring Street, New York, N.Y. 10013
All rights reserved. No part of this book may be reproduced, stored in a retrieval system,
or transmitted in any form or by any means, electronic, mechanical, photocopying,
microfilming, recording, or otherwise, without written permission from the Publisher
PREFACE
What, precisely, is a primitive fish? Most biologists would agree that the living
cyclostomes, selachians, crossopterygians, etc. cannot be considered truly primitive.
However, they and the fossil record have served to provide the information which
forms the basis for speculation concerning the nature of the original vertebrates.
This symposium of biologists from a variety of disciplines was called together to
create collectively, from the best available current evidence, a picture of the
probable line of evolution of the prototype primitive fishes. The symposium was
designed to follow one that took place in Stockholm in 1967, convened for a similar
purpose, with about the same number of participants. It is a matter of interest that
almost the entire 1967 symposium (Nobel Symposium 4) dealt only with the hard
tissues, whether fossil or modern.
In charting the course of the present symposium it was felt that the intervening
years have produced numerous lines of new evidence that could be employed in the
same way that a navigator determines his position. Each field, be it adult
morphology, geology, ecology, biochemistry, development or physiology, generates
evidence that can be extrapolated backward from existing vertebrate forms and
forward from invertebrate forms. If the intersect of only two lines of evidence
produces a navigational "fix" of rather low reliability, then an intersect, however
unfocussed, of multiple guidelines from more numerous disciplines might provide a
better position from which to judge early vertebrate history. Accordingly, we have
recorded here information from behaviorist-ecologists who have tried to see the
early vertebrates in their own prevailing environments and subject to the exacting
restraints of these environments. The biochemists and immunocytochemists can
trace particular peptide combinations from the established vertebrate taxa clear
across the terra (or mare) incogllita into the invertebrate domain. Developmental
information hopefully supplies even more insight than adult structure, and we have
tried to obtain as much of this as is available and appropriate. The large body of
accumulated physiological lore now can be drawn upon to give us some
understanding of how the early vertebrates may have solved their adaptive
functional problems.
At any rate, we have here as good a picture as is currently available of the
prevailing circumstances and probable structural and functional features of the truly
primitive fishes, if that is what these aquatic creatures may be called. For whatever
it is worth, we present this crystallization, or condensation, of current information
as it applies to early vertebrate phylogeny.
v
vi PREFACE
We gratefully acknowledge the editorial assistance of Dr. William S. Wheeler and
Dr. Christine E. Milliken, and typing assistance of Miss Sandra Kschischang and
Mrs. Linda Mather. The technical software support group of Lexisoft, Inc. of
Davis, California, was always there when we needed their assistance, for which we
are very appreciative.
We are most grateful to Dr. M. di Lullo, Director of NATO ARW Programme,
and the North Atlantic Treaty Organization for their support of the workshop.
Financial support was also provided by the Natural Sciences and Engineering
Research Council of Canada and the Western Canadian Universities Marine
Biological Society. The sponsoring organizations greatly contributed to the success
of the workshop, as did Mrs. Linda Mather, Miss Sabina Leader and Miss Anne
Bergey.
R. E. Foreman
A. Gorbman
J. M. Dodd
R. Olsson
September, 1985
CONTENTS
Scenarios: Why?
C. Gans
General Ecology of Primitive Fishes. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . II
G. Bertmar
Facts and Thoughts on Piscine Phylogeny. . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 31
H. C. Bjerring
Reconstructing the Life Cycle and the Feeding of
Ancestral Vertebrates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 59
J. Mallatt
Habitat, Phylogeny and the Evolution of Osmoregulatory
Strategies in Primitive Fishes .... . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 69
R. W. Griffith
The Brain and Sense Organs of the Earliest Vertebrates:
Reconstruction of a Morphotype . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
R. G. Northcutt
The Lateral Line System of Cyclostomes .............................. 113
B. Fernholm
Freshwater Parasitic Lamprey on Vancouver Island and
a Theory of the Evolution of the Freshwater
Parasitic and Nonparasitic Life History Types . .. . . .. .. . . .. . . .. . . .. 123
R. J. Beamish
Organ Development and Specialization in Lamprey Species. . . . . . . . . . . . . . .. 141
J. H. Youson
Early Development of Oral, Olfactory and Adenohypophyseal
Structures of Agnathans and Its Evolutionary Implications ............ 165
A. Gorbman and A. Tamarin
vii
viii INDEX
Immunologic Relations Among Fish Groups ........................... 187
G. T. Rijkers
Evolution of Temperature Regulation and of Constancy of
Function (Homeokinesis) at Different Temperatures ................. 203
C. L. Prosser
Respiration in Phyletically Ancient Fishes ......................... 217
W. Burggren, K. Johansen and B. McMahon
Regulation of Blood and Body Fluids in Primitive Fish Groups ......... 253
R. Hinge
Evolution of the Renin-Angiotensin System and Its Role
in Control of Cardiovascular Function in Fishes . . . . . . . . . . .. 275
H. Nishimura
Evolutionary Aspects of Reproduction in Cyclostomes and
Cartilaginous Fishes .................................... 295
J. M. Dodd and M. H. I. Dodd
On Urea Formation in Primitive Fishes ......................... 321
G. W. Brown, Jr. and S. G. Brown
Some Aspects of Hormonal Regulation of Metabolism in Agnathans .... .. 339
E. Plisetskaya
Hagfish Insulin: Evolution of Insulin . . . . . . . . . . . . . . . . . . . . . . . . . .. 363
S. O. Emdin, D. F. Steiner, S. J. Chan and S. Falkmer
Evolution of Gastro-Entero-Pancreatic Endocrine Systems
in Lower Vertebrates ................................... 379
M. C. Thorndyke and S. Falkmer
Functional Evolution of Gastrointestinal Hormones ................. 40 I
S. R. Vigna
Hormonal Peptide Evolution ................................. 413
Y. A. Fontaine
Tissue Distribution of Hormonal Peptides in Primitive Fishes .......... 433
M. Nozaki
Group Photo 455
Contributors and Participants 457
Index ................ . 461
SCENARIOS: WHY?
Carl Gans
Division of Biological Sciences
The University of Michigan
Ann Arbor. Michigan 48109-1048
Introduction
In beginning this symposium on the patterns disclosed by primitive fishes, we
should perhaps ask what it is that we are trying to achieve. Some of us hope to
generate schemes for the taxonomic placement of certain Recent animals and for
mapping the shifting pattern of their phenotypes through time. However, another
part of our agenda might involve the generation of scenarios, i.e. efforts to
reconstruct the functional changes producing the process of vertebrate evolution.
Obviously, there are two sets of evidence that can be used to achieve these aims.
The first derives from fossils and was emphasized in the previous symposium. The
second derives from the Recent and is the topic here.
At the start, we may wish to remind ourselves about the guidelines for making
decisions about the past events that we wish to reconstruct. Specifically, what is it
that we can observe and what is it that we may conclude from it? Obviously,
comparison is the key of this symposium, and we hope to enhance the potential for
it by assembling specialists on different groups of organisms and of distinct organ
systems. We operate less by absolute determinations of conditions in one kind of
system or animal, than by analyzing the similarities and differences disclosed by
observation of diverse species. Hence, the rules of comparison apply to our system
as well as to most others in biology, and we must keep them in mind.
Aspects in comparison
Animals provide us with three sets of data, their adult phenotype, the pattern
by which the phenotype develops after fertilization and the way the phenotype is
utilized by the organism. Thus, structure, development and function form three
sets of descriptors of currently observable phenomena (Fig. I). These descriptors
2 C.GANS
No
Fig. 1. Venn diagram to show the three kinds of current similarities
likely to be exhibited by organisms and the combined states that
these may display. Note that similarity of function is not an
aspect generated by the phenotype, but one that is permitted by
the phenotype. (After Gans, 1985, Amer. Zool., in press).
may be compared, and the similarities and differences thus disclosed may be refined
by incorporating more specimens and species and improving the reliability of the
observations.
However, the similarities and differences obtained by the study of individuals
of any single or pair of species may have multiple causes and permit only limited
conclusions about affinity. More is needed, because similarities of morphology and
physiology may reflect closeness of derivation from a common ancestor, matching
of different phenotypes to a common environment or both. Hence analysis of
affinity must take into account the potential pitfalls posed by the convergence of
characters. This difficulty requires matching our findings against phylogenetic
schemes, or cladograms. These let us test whether a particular condition seen in
several species represents a single invention (i.e. appears in a single evolutionary
line) or whether it has been invented independently a number of times (i.e. it
appears in several separate evolutionary lines; Fig. 2).
The origin of vertebrates involves another set of complicating problems. There
are few obvious "outgroups" against which the vertebrates, and for that matter the
chordates, may be compared; too many of the presumably intermediate forms
appear to have become extinct. Also, there are no immediately obvious. "ancestors"
from which "descendents" are derivable in a single, or relatively few steps. Indeed,
we see an array of vertebrate characteristics, all of which are shared and derived,
and all of which appear in the fossil record almost simultaneously, not shared with
any other group. Such a situation suggests either that much time has passed (with
the intermediate forms having all become extinct) or that there may be a common
denominator for some of the set.
SCENARIOS: WHY? 3
2 3 4
A
2 3 4
B
2 3 4
c
f
TIME
I
2 3 4
1
D
TIME
I
Fig. 2. A set of cladograms labelled and modified to show the pattern of
character states of recent organisms (A), and the way this pattern
would be modified if the vertical axis is explicitly made to
represent time and fossils mapped into the appropriate places
(C). The ecological, behavioral, physiological states seen in the
Recent have been mapped onto one cladogram (B); whereas, the
time-associated diagram indicates the potential for functional (as
well as phenotypic) shifts in each line leading to members of a
pair of sister groups (D). Obviously, selection for the transition
must have been effected by the function-associated demands
existing at the time of bifurcation.
Common denominators are of interest because it is obviously easier to posit
evolution by means of a limited number of changes than by positing multiple,
coincident, but supposedly independent, events. Incidentally, this is one reason for
