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BEHAVIORAL STRATEGY IN
PERSPECTIVE
ADVANCES IN STRATEGIC
MANAGEMENT
Series Editor: Gino Cattani

Recent Volumes:

Volume 27: Globalization of Strategy Research Edited by: Joel A.C.

Baum and Joseph Lampel
Volume 28: Project-Based Organizing and Strategic Management
Edited by: Gino Cattani, Simone Ferriani, Lars
Frederiksen and Florian Taube
Volume 29: History and Strategy Edited by: Steven J. Kahl, Brian S.
Silverman and Michael A. Cusumano
Volume 30: Collaboration and Competition in Business Ecosystems
Edited by: Ron Adner, Joanne E. Oxley and Brian S.
Silverman
Volume 31: Finance and Strategy Edited by: Belén Villalonga
Volume 32: Cognition and Strategy Edited by: Giovanni Gavetti and
William Ocasio
Volume 33: Business Models and Modelling Edited by: Charles
Baden-Fuller and Vincent Mangematin
Volume 34: Strategy Beyond Markets Edited by: John M. De
Figueiredo, Michael Lenox, Felix Oberholzer-Gee and
Richard G. Vanden Bergh
Volume 35: Resource Redeployment and Corporate Strategy Edited
by: Timothy B. Folta, Constance E. Helfat and Samina
Karim
Volume 36: Geography, Location, and Strategy Edited by: Juan
Alcácer, Bruce Kogut, Catherine Thomas, Bernard Yin
Yeung
Volume 37: Entrepreneurship, Innovation, and Platforms Edited by:
Jeffrey Furman, Annabelle Gawer, Brian S. Silverman
and Scott Stern
Volume 38: Sustainability, Stakeholder Governance & Corporate
Social Responsibility Edited by: Sinziana Dorobantu,
Ruth V. Aguilera, Jiao Luo and Frances J. Milliken
ADVANCES IN STRATEGIC MANAGEMENT VOLUME
39

BEHAVIORAL STRATEGY IN
PERSPECTIVE
EDITED BY

MIE AUGIER
Naval Postgraduate School, USA

CHRISTINA FANG
Leonard N. Stern School of Business,
New York University, USA

VIOLINA P. RINDOVA
Marshall School of Business,
University of Southern California, USA
Emerald Publishing Limited
Howard House, Wagon Lane, Bingley BD16 1WA, UK

First edition 2018

Copyright © 2018 Emerald Publishing Limited

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their use.

British Library Cataloguing in Publication Data

A catalogue record for this book is available from the British Library

ISBN: 978-1-78756-348-3 (Print)

ISBN: 978-1-78756-347-6 (Online)
ISBN: 978-1-78756-349-0 (Epub)

ISSN: 0742-3322 (Series)

CONTENTS
List of Contributors

Introduction – Behavioral Strategy: A Quick Account

Mie Augier, Christina Fang and Violina P. Rindova

PART I
THE FIELD OF BEHAVIORAL STRATEGY AND
ITS EVOLUTION

Some Thoughts on the Development of Disciplines, with

Particular Attention to Behavioral Strategy
James G. March

A Strategy for Behavioral Strategy: Appraisal of Small,

Midsize, and Large Tent Conceptions of this Embryonic
Community
Donald C. Hambrick and Craig Crossland

Decoupling and Intergroup Dynamics in Behavioral Strategy,

and a More Integrative Alternative
James D. Westphal

A Behavioral (Simonian) Perspective on (Behavioral)

Strategic Management Research
Mie Augier and Nicholas Dew

PART II
PERSPECTIVES ON BEHAVIORAL STRATEGY
AND STRATEGIZING

From Strategy to Strategic Organization

Daniel A. Levinthal

The Organizational Foundations of Behavioral Strategy

Phanish Puranam

Where to Search
Henrich R. Greve

Organizational Sensing and the Occasions for Strategizing

Sidney G. Winter

Hierarchical Sensing and Strategic Decision-making

Elad Green and Zur Shapira

Bounded Rationality, Heuristics, Computational Complexity,

and Artificial Intelligence
Richard A. Bettis and Songcui Hu

Romantics, Mercenaries, and Behavioral Rationality

Thomas C. Powell

The Three Minds of the Strategist: Toward an Agentic

Perspective in Behavioral Strategy
Violina P. Rindova and Luis L. Martins

Praxis, Character, and Competence: From a Behavioral to a

Communitarian View of the Firm
Haridimos Tsoukas

PART III
BEHAVIORAL STRATEGY IN ACTION

Behavioral Strategy and Strategy Prescription

Philip Bromiley and Devaki Rau

Behavioral Strategy: An Alternative Account of Superior

Profitability?
Christina Fang and Chengwei Liu

Behavior in Behavioral Strategy: Capturing, Measuring,

Analyzing
Charlotte Reypens and Sheen S. Levine

Teaching Strategists to Take Advantage of What Happens

William Starbuck

PART IV
EPILOGUE

Confessions of a Behavioral Strategist!

Edward J. Zajac

Index
LIST OF CONTRIBUTORS
Mie Augier Naval Postgraduate School, USA
Richard A. University of North Carolina Chapel Hill - Kenan-
Bettis Flagler Business School, USA
Philip Bromiley University of California, Irvine, USA
Craig Crossland University of Notre Dame, USA
Nicholas Dew Naval Postgraduate School, USA
Christina Fang New York University, USA and CKGSB, China
Elad Green Google, Switzerland
Henrich R. INSEAD, Singapore
Greve
Donald C. Pennsylvania State University - Smeal College of
Hambrick Business, USA
Songcui Hu University of Arizona, USA
Sheen S. Levine University of Texas at Dallas, USA
Daniel A. Wharton School, University of Pennsylvania, USA
Levinthal
Chengwei Liu University of Warwick, UK
James G. March Stanford University, USA
Luis L. Martins University of Texas at Austin, USA
Thomas C. Said Business School, University of Oxford, UK
Powell
Phanish INSEAD, Singapore
Puranam
Devaki Rau Northern Illinois University, USA
Charlotte University of Warwick & University of Texas Dallas,
Reypens USA
Violina P. Marshall School of Business, University of Southern
Rindova California, USA
Zur Shapira New York University, USA
William Lindquist College at University of Oregon and NYU,
Starbuck USA
Haridimos University of Cyprus and University of Warwick,
Tsoukas Cyprus
James D. Stephen M. Ross School of Business, University of
Westphal Michigan, USA
Sidney G. Wharton School, University of Pennsylvania, USA
Winter
Edward J. Zajac Northwestern University - Kellogg School of
Management, USA
INTRODUCTION – BEHAVIORAL
STRATEGY: A QUICK ACCOUNT

Mie Augier, Christina Fang and Violina P. Rindova*

The origins of strategy are inherently behavioral. Some point to the

publication of Simon’s Administrative Behavior in 1947 as such an
origin point. Others point to the works of Andrews and his
colleagues that defined business policy, and later strategy, as “the
study of the functions and responsibilities of general management
and the problems which affect the character and success of the total
enterprise” from the viewpoint “of the chief executive or general
manager, whose primary responsibility is the enterprise as a whole”
(Learned, Christensen, Andrews, & Guth, 1965, p. 3).
The origins of strategy have also been undeniably organizational:
Simon’s Administrative Behavior (1947), March and Simon’s
Organizations (1958), and Cyert and March’s A Behavioral Theory of
the Firm (1963) focused attention on the study of organizational
policies, decisions, and structures as the contexts that shape how
decisions are made. In one account of the field’s history, the
behavioral roots of strategic management were set aside in favor of
adopting economic theories to legitimate the fledgling field, provide
its baseline equilibrium assumptions, and guide the adoption of
sophisticated econometrics methods (Rumelt, Schendel, & Teece,
1991). Michael Porter’s reorientation of the industrial-organization
economics research toward understanding the factors that limit
competition, along with the research on related diversification in the
1980s, provided the theoretical foundation of business-level and
corporate strategy.
Despite this shift toward economics, the behavioral roots of
strategy remained strong, and researchers working from a
behavioral perspective embraced “strategy process” approach. The
strategy field evolved as a multiparadigmatic field, divided between
strategy content and strategy process research. In the last 10 years,
as we recount below, the term “behavioral strategy” has come to the
forefront of strategic management (Bromiley & Rau, 2014; Gavetti,
2012; Levinthal, 2011; Powell, Lovallo, & Fox, 2011). On the one
hand, the term congealed relatively recently, following the footsteps
of sister disciplines which have already embraced behavioral
approaches (such as behavioral economics, behavioral finance, and
behavioral accounting). On the other hand, the assumptions,
theoretical ideas, and empirical work that pertain to the domain of
behavioral strategy have been developing since the emergence of
the field of strategy itself.
The goal of the volume is to bring together scholars from several
generations who have led and defined the behavioral approaches in
strategy to reflect on the past, present, and future of behavioral
strategy. Thus, rather than seeking empirical contributions that
would fill in research gaps and expand research in specific domains,
we endeavored to: (a) represent the diversity of perspectives that
inform behavioral research in strategic management; (b) open up a
space for reflection and provocation by scholars who are widely
recognized as thought leaders both in their respective strands of
behavioral strategy research and in the field as a whole; and (c)
offer a set of perspectives and directions for the field of behavioral
strategy at a pivotal moment in its evolution. We received
enthusiastic responses from many eminent scholars represented in
this volume, as they shared our premise that this is an opportune
moment for a collective reflection on the theoretical foundations, key
problems, varying approaches, and significance of behavioral
strategy, and that such a reflection could provide some signposts for
the diverse scholarly community it now encompasses.
 To provide some background on the evolution of the field, we
first offer a synoptic overview of what we see as behavioral
approaches to strategic management research and how they inform
the emerging field of behavioral strategy. We then briefly recount the
formative events that led to the delineation of behavioral strategy as
an identity domain and review extant definitions of the field, adding
the “big tent” view, which motivated the design of this volume. We
conclude by briefly summarizing the contributions to the volume.

THE DIVIDE IN STRATEGIC MANAGEMENT

RESEARCH
As Levinthal (2011, p. 1517) succinctly pointed out, “the
fundamental divide” in strategy research arises from the underlying
assumptions of scholars whether “choice processes are ‘rational’ and
adhere to the axioms of expected utility theory, or whether choice
processes are characterized by some ‘behavioral’ mechanisms.” The
shift from behavioral to economic approaches received a significant
boost from Michael Porter’s seminal paper titled “How Competitive
Forces Shape Strategy” (1979), and his follow-on book on
Competitive Strategy (1980).1 Moving away from the core concerns
with monopoly power in industrial economics, Porter’s work paved
the way to understanding how the conditions that interfere with free
competition enable the generation of abnormal returns. The
resulting Five Forces framework has been one of the most influential
ideas in strategy, and a core framework for both teaching and
research, focusing strategic management research efforts on the
structural characteristics of industries.
In the mid-1980s, the Resource Based View of the firm (the RBV)
directed attention to the firms’ internal resources and capabilities.
Barney (1986, 1995) argued that firms need to shift attention from
an external industry analysis, which rests on public information to an
internal analysis of resources which are valuable, rare, inimitable,
and nonsubstitutable. It is important to note that the RBV traced its
roots to Penrose (1959), who argued that “a firm is […] a collection
of productive resources the disposal of which between different uses
and over time is determined by administrative decision” (p. 24). She
also emphasized the heterogeneity of these resource collections and
their role in generating superior economic performance. Penrose,
however, further linked the superior performance effects of
resources – which she defined as physical things and people – to the
cognitive capacities of firm strategists. It is through their depth of
experience and resulting insights that strategists conceived of new
productive services from the resources to pursue growth
opportunities. Yet, most of the subsequent theory and research in
the RBV tradition generally adopted general equilibrium assumptions
emphasizing the economic rationales behind the rent-generating role
of resources (e.g., Peteraf, 1993). As a result, the behavioral
component in Penrose’s argument was lost and RBV was brought
into the fold of the “optimizing hyper-rationality” that characterizes
the economic perspectives in strategic management (Bromiley &
Rau, 2014: p. 6).
In addition to the development of RBV as an internal perspective
on competitive advantage, the strategy field witnessed several
prominent intellectual responses to the pre-dominance of economic
theory in strategy: the strategic leadership “upper echelons”
perspective (Hambrick & Mason, 1984); the Neo-Carnegie
perspective (Gavetti, Levinthal, & Ocasio, 2007; Nelson & Winter,
1982); and the cognitive perspective (Porac & Thomas, 1990; Porac,
Thomas, & Baden-Fuller, 1989; Reger & Huff, 1993). While their
theoretical positions, empirical approaches, and research agendas
are distinct and separate, each represented a different behavioral
reaction to the growing centrality of the economic paradigm and
provided the theoretical and empirical foundation of much of the
current work in behavioral strategy, beyond early classics noted at
the outset.
The strategic leadership perspective defined a major area of
inquiry within strategic management research (Finkelstein &
Hamrick, 1996) by focusing on the demographic and human capital
characteristics of CEOs and general managers, and the structure,
composition, and dynamics of top management teams and corporate
boards of directors. It offered a behavioral perspective that
explained strategic choices and firm performance as driven by the
individual characteristics and group properties. While this approach
did not focus on psychological processes in its early stages of
development, it pointed to the importance of individual and group
characteristics in shaping the underlying information processing
dynamics in strategic decision-making.
The Neo-Carnegie tradition provided another important influence
on the behavioral foundations of strategy (Gavetti et al., 2007).
Nelson and Winter (1982) evolutionary economics challenged the
economic orthodoxy by articulating that actors may not consider all
latent set of alternatives and instead rely upon routines and the
status quo. Relatedly, James March and his colleagues have built on
earlier classics such as Cyert and March (1963) and argued that
actors do not update priors and learn optimally. Rather, they learn
from experience in response to performance feedback. Such learning
leads to various behavioral “pathologies” such as the myopia of
learning, superstitious learning, and competency traps. In addition,
Levinthal (1997), and subsequent work utilizing the NK paradigm,
further showed how learning is complicated in interdependent
contexts. Thus, the Neo-Carnegie approaches articulated a
behavioral perspective of strategy as being driven by habits and
routines and learning from feedback that is ambiguous, delayed, or
otherwise misleading. March (2006) made further break from
traditional conceptions of rationality, arguing that the field of
strategic management can gain from exploring foolishness and
adaptive intelligence.
A third behavioral response was evident in the cognitive
perspective in strategy research that emerged in the late 1980s and
early 1990s. This perspective built on the growing interest in
interpretive processes in organizations to investigate their strategic
relevance for strategic management questions, such as threat-
opportunity framing (Jackson & Dutton, 1988), cognitive industry
boundaries (Fombrun & Zajac, 1987; Huff, 1982), competitive groups
(Porac & Thomas, 1990), dominant logic of the corporation
(Prahalad & Bettis, 1986), and industry knowledge recipes (Spender
1989). This work shifted the focus from the characteristics of the
environment to the interpretation of the environment by strategists,
as a function of their cognitive structures and processes. Porac et al.
(1989) influential paper on the Scottish knitwear industry showed
that managers not only interpret environments, but also construct
them by enacting their mental models into distinct patterns of
interactions and exchange. The cognitive perspective, therefore,
acknowledged that environmental complexity leads to partial
representations and potentially bounded rational decisions. However,
it took a step further highlighting the creative constructive power of
managerial perceptions, knowledge, and beliefs, presenting the
environments themselves as constructed through perceptions,
actions, and communications (Rindova & Fombrun, 1999).

ORGANIZING MOVES
The three different strands of behavioral approaches to strategy
briefly summarized earlier operated as distinct paradigms throughout
the 1990s and early 2000s. In 2007, Jerker Denrell and Christina
Fang organized a symposium at the Annual Meeting of the Academy
of Management titled “Behavioral Strategy: A Research Agenda” with
the goal of bringing together researchers dealing with strategy,
behavioral finance, and behavioral decision theory to examine core
strategic issues around strategic factor market efficiency. In 2010,
Daniel Lovallo (one of the panelists in the 2007 symposium) and
Oliver Sibony published an article in the McKinsey Quarterly titled
“The Case for Behavioral Strategy” calling for strategists to take into
account known heuristics and biases in decision-making and to
proactively seek to debias important decisions. The paper quickly
became one of the most downloaded papers from the McKinsey
Quarterly that year. Later that year, Sheen Levine and Shayne Gary
organized a professional development workshop at the Academy of
Management Annual Meetings with the explicit goal to rethink the
neoclassical economic paradigm as a foundation for strategic
management research and to advance behavioral and cognitive
research as an alternative theoretical foundation for strategy.2
Around this time, efforts to formalize behavioral strategy as a
distinct field accelerated. Thomas Powell, Daniel Lovallo, and Craig
Fox guest-edited a special issue at Strategic Management Journal
(SMJ) titled “Psychological Foundations of Strategic Management”
that called for papers examining the psychological foundations of
strategy. Thomas Powell and Richard Bettis, among others, also
spearheaded initiatives that led to the development of Behavioral
Strategy Interest Group within the Strategic Management Society.
Through these activities and publications, behavioral strategy
came to be defined as merging “cognitive and social psychology with
strategic management and practice” with the goal to “strengthen the
empirical integrity and practical usefulness of strategy theory by
grounding strategic management in realistic assumptions about
human cognition, emotion, and social interaction” (Powell et al.,
2011, p. 1371). The psychological foundations of behavioral strategy
are now widely recognized. For example, Gavetti (2012, p. 267)
argues that behavioral refers to psychological underpinnings of a
given phenomenon; and Bromiley and Rau (2014) further identify
behavioral strategy with the assumption of bounded rationality. This
approach has a number of advantages. First, its clear and distinct
theoretical foundation complements the predominant economic
approaches. Second, this approach also shares similar conceptual
and methodological apparatus with behavioral economics, thereby
creating opportunities for cross-fertilization among related sister
disciplines.
While we agree that human psychology should be at the center
of behavioral research in strategy, we see the need for two
fundamental augmentations of the approach and definitions of
behavioral strategy. First, we see a need for a broader understanding
of the human element in strategy and markets. As Powell (2017, p.
67) has recently argued, “Behavioral research shows that human
market participants do not behave like rational economic agents.
Real people are in many ways more impressive than economic
agents. They are capable of passion, benevolence, insight, and
perseverance. They have moral and esthetic ideals, and they exhibit
altruism, trust, reciprocity, compassion, justice, loyalty, and love.” We
therefore advocate for an inclusive approach to studying the people
who make, evaluate, or contribute to firm strategies. Such an
inclusive approach incorporates the study of values, emotions, social
and cultural embeddedness, identity, esthetic knowledge and
perception, and the lived experience. In terms of disciplinary
knowledge, it would welcome disciplinary transfusions not only from
cognitive and social psychology, but also from sociology, political
science, anthropology, philosophy, linguistics, communications, and
cultural and art studies.
Second, while bringing the focus back on the strategists – as
individuals and groups – is valuable, firms and even markets are not
simply collection of individuals. Indeed, the long-standing behavioral
roots of strategy are organizational. Further, the changing nature of
organizations, and the ways in which they incorporate customers
and other actors in their core productive activities, suggests that
there is an opportunity for behavioral strategy to broaden its
disciplinary foundations to account for collective-level behavioral
processes, such as those studied in sociology. The collective
contributions to this volume, discussed next, point in similar
directions. While reaffirming the psychological foundations of
behavioral strategy, they also open up new conversations about the
human experience and the role of organizations.

THE CONTRIBUTIONS IN THIS VOLUME

The contributions represent reflections on and theoretical extensions
of behavioral research in strategic management and are organized in
three sections and an epilogue. Each section addresses the goals for
this volume in different ways. The first set of papers discusses the
emergence and evolution of the field of behavioral strategy. The
papers in the second set offer a variety of perspectives on behavioral
strategizing, both extending core approaches and charting directions
for new ones. The papers in the final set spell out important
implications for empirical research, normative recommendations, the
study of firm performance, and teaching strategy. The epilogue
offers one personal narrative of navigating the emergence of the
field.
The first section opens up with Jim March’s contribution titled
“Some Thoughts on the Development of Disciplines, with Particular
Attention to Behavioral Strategy.” He reflects on the evolution of
fields in general, taking a close look at the evolution of the “sister
discipline” of organization studies. The development of the field, like
much of its content, is the result not of optimal, well-calculated
rational-choice decisions. Instead, it involves a diverse set of ideas,
intellectual traditions, and developments. The field of behavioral
strategy is in a precarious position of emergence, which is both an
opportunity and a challenge. Part of the opportunity, March
suggests, is to expand beyond its traditional emphasis on decision-
making and bounded rationality. In the next paper, titled “A Strategy
for Behavioral Strategy: Appraisal of Small, Midsize and Large Tent
Conceptions of this Embryonic Community,” Don Hambrick and Craig
Crossland highlight the ambiguity of the term of behavioral strategy
and its subfield, describing three alternative conceptions of
behavioral strategy, each having pros and cons, and each pointing to
different paths for the future of the field. A “small tent”
conceptualization essentially applies the logic of behavioral
economics to strategy; a “midsize tent” conceptualization includes
the application of cognitive and social psychology to strategic
management phenomena; and a “large tent” conceptualization
broadens up the behavioral foundations for strategy even further.
The authors offer rich discussion of vibrant streams of research
within the midsize and large tent definitions and offer their
expectations of how these alternative scenarios of field may unfold.
 In the third paper in this section, Jim Westphal makes a strong
case for the large tent view in his contribution titled “Decoupling and
Intergroup Dynamics in Behavioral Strategy, and a More Integrative
Alternative.” He builds on social categorization theory and personal
interviews with colleagues with different types of behavioral self-
identifications to warn the emerging field about the dangers of
ingroup/outgroup dynamics that marginalize perspectives that ought
to be included. He proposes an approach that takes
interdisciplinarity to heart and encourages collaboration across
subfields in order to increase both the contributions and the position
of behavioral strategy to the field, as well as creativity and the rate
of theoretical advancement in the field as a whole. The paper by Mie
Augier and Nick Dew concludes this section by reviewing larger
history of the field of strategic management “with Simon in mind”
and offering several guiding principled inspired by Simon’s work, on
which the field of behavioral strategy can build.
The second section, “Perspectives on Behavioral Strategy and
Strategizing” includes nine contributions, each taking a different
perspective on strategy making and the core processes of
strategizing. In the first contribution titled “From Strategy to
Strategic Organization,” Dan Levinthal reminds us that strategy
problems are fundamentally “complex” and “hard” and surveys
different approaches for dealing with this strategic recalcitrance. He
advocates a pragmatic approach to strategy that focused on the
actions and interactions of boundedly rational individuals to improve
the survival of the organization. Such an approach invites taking
strategic organizational design seriously and in an evolutionary and
behavioral way. Phanish Puranam’s contribution titled “The
Organizational Foundations of Behavioral Strategy” picks up on the
topic of organizations as contexts for strategic behavior and
articulates both its core components: goals, representations, and
choices, and the organizational designs that operate on them. The
paper makes a compelling case for behavioral strategy as
understanding individuals in context as complex systems nested
within larger complex systems.
 The next three papers focus on two core strategizing processes –
search and sensing. Henrich Greve poses the question “Where to
Search” and extends the theory of problemistic search by observing
that current approaches fail to explain “how a firm goes from a
falling short on a broad goal to taking a specific action in response.”
He considers a multiplicity of alternatives about how organizations
respond to broad goals, identifying goal multiplicity, power,
cognition, and the environment, all as having the capacity to direct
search.
Sid Winter’s essay on “Organizational Sensing and the Occasions
of Strategizing” presents a provocative perspective on importance of
deliberative thinking and the limits to its understanding, especially
under conditions of information asymmetries – “us” versus “them”
strategizing. Under such conditions, the keen attention to constraints
– and the development of a variety of sensors–are paramount to
strategizing. Sensing then is a complex system that bestows
organizations differential recognition of problems and opportunities.
Sensing is also the concept explored in Green and Shapira’s paper,
“Hierarchical Sensing and Strategic Decision Making”. They simulate
an organization as a complex signal detection system, comprising
individuals who perceive, interpret, and act upon information they
receive. They investigate the interaction between organizational
hierarchical structure and individuals’ attentional and cognitive
constraints on the organization’s capacity to identify changes in its
environment. The paper by Rich Bettis and Songcui Hu, “Bounded
Rationality, Heuristics, Computational Complexity and Artificial
Intelligence,” in turn, draws on Newell and Simon’s work to argue
that computational complexity provides a rigorous, important, and
largely ignored theoretical grounding for the study of bounded
rationality. The optimal solutions of many important problems in
economics, behavioral strategy, and other fields are simply
intractable and cannot be generally be computed by any known or
envisioned computer. As such, strategy is by necessity behavioral,
and heuristics rather than optimization should be the basis for
managerial and organizational decision-making.
 The three final papers in this section offer perspective that depart
from the current definitions and approaches in behavioral strategy.
Thomas Powell, in his paper titled “Romantics, Mercenaries, and
Behavioral Rationality” advances the notion of behavioral rationality,
which – in contrast to normative economic rationality – “draws on
human beliefs, passions and moral values to produce commitment-
maximizing courses of action.” The notion of behavioral rationality
implies two distinct dimensions for behavioral strategy: decisions –
which have received the bulk of prior research attention – and
actions – which depend on different psychological processes and
states that have been largely ignored. The success for the field of
behavioral strategy, he further argues, is less likely to result from
removing cognitive biases (debiasing) than from helping firms
understand how to generate strategic actions. Violina Rindova and
Luis Martins’ contribution titled, “The Three Minds of the Strategist:
Toward an Agentic Perspective on Behavioral Strategy” similarly
challenges the focus of behavioral research on decisions alone. They
build on the sociological work on agency to distinguish between
agentic and decision-making/information processing capacities.
Because agency unfolds across the temporal horizons of the past,
present, and future, an agentic perspective accounts for the different
sociocognitive processes that anchor strategists in the past, enabling
them to manage the present and support their envisioning of the
future. Strategists’ agentic capacities, in turn, depend on how they
combine these temporally oriented cognitive processes. Finally,
Haradimos Tsoukas, in his paper titled “Praxis, Character and
Competence: From a Behavioral to a Communitarian View of the
Firm” introduces a communitarian view of the firm that departs from
behavioral strategy’s limiting assumptions–namely, the assumptions
of methodological individualism and dualism. The communitarian
view accounts for praxis, and its formative effects on agents, as well
as for teleological action, and its implications for the value
commitments that firms make. These ideas implicate Selznick’s
notions of distinctive competences and organizational character,
pointing behavioral strategy toward recovering some of its normative
orientation roots.
 The final section, “Behavioral Strategy in Action,” contains four
papers that address the implications of behavioral strategy for core
domains of action, including theoretical prescription, firm
performance, empirical research, and teaching. In particular, the
paper by Phil Bromiley and Devaki Rau, “Behavioral Strategy and
Strategy Prescription” discusses and contributes to the
underdeveloped area of “strategy prescription.” While the idea of a
united prescriptive theory is “attractive but illusory,” they find
behavioral strategy insights can fruitfully be applied to strategy
prescription. They stipulate the foundations of such prescriptions:
bounded rationality (and satisficing) and decomposed problem-
solving, consistent with mid-range theories of organizations. Mid-
range theories of strategy, they suggest, emphasize near
decomposability and hierarchical organizations of human systems,
and can be useful for understanding organizations and processes
and structures for developing ways to frame problems and for
offering strategies. Christina Fang and Chengwei Liu concretize the
idea of prescriptive behavioral strategy in their contribution titled
“Behavioral Strategy: An Alternative Account of Superior Profitability,”
which proposes a normative theory of superior profitability. Building
on decades of research on sociocognitive biases, they argue that
strategists can exploit these generative mechanisms of behavioral
biases to outsmart and outcompete rivals in a systematic way.
Charlotte Reypens and Sheen Levine turn their attention to the
empirical tools that can advance research in behavioral strategy.
Their contribution titled “Behavior in Behavioral Strategy: Capturing,
Measuring, Analyzing” presents a toolbox of methods to capture
observed outcomes and expose underlying processes: experimental
tasks to simulate environments and capture behavior protocol
analysis and text analysis to understand cognitive processes,
applying them to collect data on two key topics: the exploration–
exploitation trade-off and strategic risk taking. Finally, Bill Starbuck,
in his paper titled “Teaching Strategists to Take Advantage of What
Happens” challenges our approach to teaching strategy and raises
the possibility for improving teaching effectiveness in the domain of
strategy by teaching students to recognize and effectively respond to
unexpected events. He discusses four helpful procedures that tend
to improve firms’ responses to strategic events: (1) future-perfect
framing, (2) scanning for relevant events, (3) experimenting with
strategic variations, and (4) watching experimental results carefully
and events opportunistically.
As a closing contribution to the volume, Ed Zajac offers a
personal narrative to discuss the mindset inclined to connect
intellectual worlds, such as behavioral and economic views on
strategy, that had been seen as divided […] well, it wouldn’t be fair if
we told his story, would it?

NOTES
* Authorship is alphabetical.
1. The book was voted the ninth most influential management
book of the twentieth century in a poll of the Fellows of the
Academy of Management.
2. This yearly workshop has attracted on average 200 attendees
and has involved speakers ranging from evolutionary biology to
computer science.

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 PART I
THE FIELD OF BEHAVIORAL
STRATEGY AND ITS EVOLUTION
SOME THOUGHTS ON THE
DEVELOPMENT OF DISCIPLINES,
WITH PARTICULAR ATTENTION TO
BEHAVIORAL STRATEGY

James G. March

ABSTRACT
The earliest contributors to discussions of strategy
were advisors to military leaders, and that model
was carried into early business schools, where the
teachers of strategy were, for the most part, people
with extensive experience as executives or advisors
to them. The key course materials were anecdotes
and cases, and the standard intellectual discourse
was organized around recollected episodes in
organizational history. The central contributions of
the early teaching of strategy were consciousness of
the complications introduced by complexity,
competition, and attention to the second-order
surprises of intentional action. There was neither a
pretense of theory nor a significant involvement in
research.
Although it shared in the onus of a general academic
skepticism about the academic legitimacy of research
on business, the “discipline” of strategy sought to
emulate the attributes of more established
disciplines. The new field was typified by an early
open interdisciplinary flavor that facilitated the
differentiation of a new field, and a subsequent
refinement that restricted access. By the start of the
twenty-first century, this process had run much of its
course, and the field of strategy had taken its place
as a reasonably respectable academic specialty. The
history of an emphasis on real organizations in real
situations led to an openness to anchors drawn from
sources other than conventional economics. These
included particularly the theory of games, the
evolutionary theory of the firm, and the behavioral
theory of organizations.

The struggle for respectability in economics was

repeatedly frustrated by the difficulty of discovering
a formulation that honored the litany of economics
while fitting the observations of real strategy making.
The future seems likely to be more of the same, a
combination of efforts to secure recognition through
emulation of the standards and barriers to entry that
characterize established disciplines, and of
exploratory gambits that are mostly destined to be
forgotten. The optimal balance is likely to be as
elusive as it is in other domains.

Keywords: Intellectual history; disciplines;

exploration; academic legitimacy; exploitation;
behavioral strategy
There are enough definitions of strategy available to make the
definition of “behavioral strategy” a particularly uncertain one. My
response to this uncertainty is to limit myself to a minimalist
definition by which strategy is a plan for future action that
emphasizes outcomes in the relatively distant future and outcomes
that depend to an important extent on the actions of adversaries
taken in response to one’s own. The study of strategy seeks for the
most part to discover an approach to strategy that leads to favorable
outcomes. The behavioral study of strategy seeks to understand the
processes by which strategies are adopted in organizations and the
implications of those behavioral characteristics for strategy making.

AN INCOMPLETE HISTORY
The teaching of strategy making in this sense quite easily lays claim
to relevance in the training of managers as a component of the
teaching of strategy. Although it is not yet admitted to the inner
sanctum of academe, “strategy” in business administration is now a
scholarly field somewhat remotely analogous to chemistry, philology,
and finance. It was not always so. The earliest contributors to
discussions of strategy were advisors to military leaders, and that
model was carried into early business schools, where the teachers of
strategy were, for the most part, people with extensive experience
as executives or advisors to them. The key course materials were
anecdotes and cases, and the standard intellectual discourse was
organized around recollected episodes in organizational history.
The teaching of strategy was embedded in schools of applied
economics or business. Economics has long been typified by some
uncertainty about whether its theory should be seen as a guide to
intelligent economic behavior or as a predictor of actual behavior. In
modern practice, the distinction is blurred by the assumption,
sometimes made explicit but sometimes not, that competition for
survival assures that the actual behavior of survivors will
approximate the optimal, and theories of optimal behavior can be
used to predict actual behavior. At least 90% of contemporary
economics follows that route.
The optimal behavior that is postulated in almost all modern
economics is some variation of means-end rationality, the idea that
intelligent individuals and organizations maximize expected
subjective returns. Such behavior is both recommended and
predicted for economic actors. The central contributions of the early
teaching of strategy were consciousness of the complications
introduced by complexity, competition, and attention to the second-
order surprises of intentional action. Clausewitz wrote of the “fog of
war.” There was neither a pretense of theory nor a significant
involvement in research.
This changed gradually during the last half of the twentieth
century and the first decade of the twenty-first. Although it shared in
the onus of a general academic skepticism about the academic
legitimacy of research on business, strategy became a recognized
subspecialty of business administration. As such a field, it claimed
authority to recruit and appoint new colleagues to professorial
positions. Scholarly journals appeared, as did scholarly associations.
Research dissertations purporting to contribute to strategy as a
scholarly field appeared at major research institutions.
The search for order was entirely conventional. The “discipline”
of strategy sought to emulate the attributes of more established
disciplines. It created scholarly journals with peer-review procedures
and (gradually) shared understandings of the characteristics of
quality research. As the journals differentiated themselves with
respect to standards, so also did individual scholars differentiate
themselves by publishing in them. The field laid claim to academic
positions that became the possession of properly anointed scholars
who gained control of entry to those positions and who began to
supervise the advanced training of new entrants to the field and to
positions within it. They created professional associations that
managed the careers of members of the field, establishing and
maintaining standards of merit.
 By the start of the twenty-first century, this process had run
much of its course, and the field of strategy had taken its place as a
reasonably respectable academic specialty. It had neither the cachet
nor the history of the great academic disciplines, but it had standing.
The story was very similar to the stories that could be told of
academic fledglings such as biochemistry and sociology. In each
case, the new field was typified by an early open interdisciplinary
flavor that facilitated the differentiation of a new field, and a
subsequent refinement that restricted access. Strategy had
substantially molded an understanding of a field with reasonably
well-established terms of discourse, career lines, and rights to
academic positions. That shaping of the field led strategy to a
position rather close to economics. An important part of this
development drew from the writings of Michael E. Porter and his five
forces analysis, which was an outgrowth of industrial organizational
economics.
It would be a mistake to see this history as primarily a
consequence of intended, rational choices. The directions of action
that have distinguished the development of strategy and behavioral
strategy as quasidisciplines has been little determined by conscious
rational choice, but have arisen from the somewhat less rational
ways decisions happen in organizations (March, 1994). In particular,
the field’s voyage in academic waters led to experimentation with
several possible academic anchors. There was an obvious link to
economics and to rational theories of choice, but the history of an
emphasis on real organizations in real situations led to an openness
to anchors drawn from other sources. Three quite varied deviant
theoretical gambits become particularly important. The first was the
theory of games (Von Neumann & Morgenstern, 1944). Game theory
provided a thoroughly respectable framework that had an emphasis
on rationality, competition, and the long run. The second anchor was
the evolutionary theory of the firm (Nelson & Winter, 1982).
Evolutionary theory, with its emphasis on ecological interactions,
complexity, and effects, led to greater awareness of industry-wide
and community-wide consequences of strategic choices made by
individual firms. The third anchor was the behavioral theory of
organizations (Cyert & March, 1963). By focusing on how decisions
actually occurred in modern organizations, this anchor moved the
field a bit from its normative premises and legitimized the term
“behavioral strategy” as focused on efforts to understand how
strategies evolved in actual organizations.

TALES OF SUCCESS AMONG TALES OF FAILURE

The story is extremely conventional and could be written with only
modest idiosyncratic features for virtually any established scholarly
discipline in the western world. Most of the survivors have followed
similar routes. They begin with relatively precarious combinations of
previously separate components, gradually build acceptance as
autonomous specialties, and then construct standards and rules that
provide barriers against entry. It is a process that typically has
involved building independent disciplinary status.
The most conspicuous success story within schools of business is
the story of finance. Finance has long been a field of instruction in
business schools, originally taught primarily by experienced financial
officers with a modicum of economics and numerous practical
examples. When, in the 1950s and 1960s, there was a flowering of
fundamental ideas, concepts, and models, including behavioral
finance, the field became a recognized specialty within economics,
with confirmation through Nobel prizes.
The story of finance is a story of success in academic terms. Not
all of the stories have been a success. Not all of the would-be
scholarly disciplines that have appeared have reached the pinnacle
of uniformity that allowed them to survive. Some scholarly start-ups
failed. A perusal of college catalogs over the 30 years following
World War II will reveal numerous academic initiatives that failed to
survive. For the most part, they are forgotten, and history remains
primarily a history of success and survival; but the failures are also
informative.
 Some of the unsuccessful post-World War II initiatives were
derivatives of the behavioral science movement in the 1940s, 1950s,
and 1960s. This movement, which captured a good deal of academic
and financial support, was built on war-time successes. It
emphasized interdisciplinary research and the use of mathematics in
data analysis and theory building. It shaped an era in which such
research was heavily favored and in which several new academic
bailiwicks became established, including policy schools and statistics
departments.
One conspicuous disciplinary initiative in the social sciences
during this period was the Harvard University Department of Social
Relations, an interdisciplinary collaboration among three of the social
science departments at Harvard University (anthropology,
psychology, and sociology) beginning in 1946. The department
flowered, had a distinguished history, but ultimately (by the 1970s)
died.
In most academic dimensions, the Harvard Department of Social
Relations was a success. The department was successful both in
recruiting distinguished faculty,1 and in attracting and graduating
productive and imaginative doctoral students.2 Despite these
successes, the program disaggregated into its component
departments around 1972, after about 25 years of distinction. There
are many possible explanations for the failure to survive; but most
conspicuously, the Harvard Department of Social Relations did not
create idiosyncratic conceptions of scholarly virtue and did not
replicate itself significantly as an organization. Its graduates found
themselves, for the most part, employed in conventional disciplinary
departments. As a result, there was no sustained development of a
differentiated discipline. The psychologists from the Department of
Social Relations thought of themselves as psychologists and found
jobs and published primarily in psychology; the sociologists in
sociology; and the anthropologists in anthropology. Departments of
Social Relations did not thrive away from Harvard.
A somewhat similar, though fuzzier, history can be found within
business schools. A quasidiscipline of organization studies arose in
the 1950s and 1960s, bringing together particularly scholars from
psychology and sociology but including also some recruited from
political science and economics. The quasidiscipline spawned
associations and journals that became important and established
itself as a recognized subcomponent of the business school
intelligentsia. In the process, the study of organizations merged with
and often replaced the study of management.
Organization studies had difficulty maintaining stability in the face
of three internal dynamics. First, the tendency of organizational
psychologists and organizational sociologists to differentiate
themselves from each other and seek separate disciplinary
validation, thereby undermining unity and exchange within
organization studies. Behavioral students of decision-making were
only to a minor degree associated with organization studies as a
unified discipline and sought and secured independent recognition in
psychology, sociology, and economics departments. However, they
thrived best in professional schools, particularly business schools.
Second, the tendency of students of organization to seek
legitimacy within business schools by finding practical uses of
organizational research. The financial and reputational attractions of
consultancy were powerful. Some of the more highly renowned
students of organizations were better known for their consulting
advice (often provided in books) than for the contributions of their
scholarly research.
Third, the tendency of students of organization to see the
domain of strategy as a domain attractive to invasion. It became
clear that organization studies, built heavily on behavioral ideas of
organizational decision-making and under pressure to be relevant,
would not long ignore strategy making as one aspect of decision-
making, and that some activists in strategy studies would welcome
the linkage as a possible intellectual counterforce to economics.
In combination, these three dynamics tended to destabilize
organization studies. Organization studies became a success, but a
fragile one.
 THE FUTURE OF BEHAVIORAL STRATEGY
Behavioral strategy sits in a precarious position similar to that of
organization studies. Located almost exclusively in academic schools
of applied economics, it is marked by elements of a non-academic
tradition and only a half-hearted acknowledgment of any claim to
being an established discipline. Its domain is conspicuously
important to business education, but its intellectual contribution is
unclear. It relies on questions much more than answers. There is no
agreed-upon body of received truth to identify the possession or
acquisition of knowledge. Lacking a strong research involvement, it
relies heavily on links to domains such as organization studies, which
suffer from some of the same weaknesses.
Like all precarious positions, behavioral strategy provides an
opportunity. The field is relatively open to ideas and to scholars
associated with other fields. In behavioral strategy, as in other fields,
this openness will almost certainly give way gradually to the erection
of standards and boundaries that isolate the discipline from late
arrivals and build a structure of beliefs and knowledge that will
differentiate legitimate behavioral strategy scholars from others. The
opportunity is two-fold: (1) to open the field to ideas and (2) to
provide closure around them.
The twenty-first century history of behavioral strategy has been
one in which both opportunities were pursued to a limited extent. A
few academic leaders of behavioral strategy were identified and
located in senior professorships in top universities. Journals open to
papers in behavioral strategy sought and achieved academic
respectability. Communities of behavioral strategy professors gained
status in business schools. These clusters attracted renegade
scholars from other disciplines. The most obvious forecast of the
future is that the process will continue, and the field of behavioral
strategy will become more firmly established as an independent
academic domain within departments of strategy in business
schools.
 There are, of course, numerous reasons why the opportunity
might not be taken. The study of strategy has fed on external
traditions of ideas, most notably industrial economics, game theory,
evolutionary theory, and theories of bounded rationality, but it does
not always find the feast to be easily digested. For example,
although many of the basic ideas of intentional bounded rationality
are manifest in behavioral treatments of strategy, more recent ideas
of ambiguity, imitation, adaptation, and endogenous values have
proven less ripe for adoption. It is possible that the difference
mirrors a difference in fundamental utility, but it is also possible that
the difference stems from an incompatibility of the latter ideas with
important elements of the pursuit of respectability. Such an
incompatibility limits the absorptive capacity of thinking about
strategy, a limitation that inhibits new developments along this line.
It is largely taken for granted that success requires a balance
between exploiting what is already known and exploring what might
come to be known, a balance that is complicated by exchanges
among organizations and over extended periods of time. The
mélange of efforts to refine the study of behavioral strategy has led
to a persistent sense of disorganization and a persistent search for
methodological and substantive order. This drive for disciplinary
closure, when successful, normally strengthens a field within
academic battlegrounds, but it is usually primarily exploitation. A
field becomes a discipline by tightening standards for research and
discussion and for training, by improving and standardizing the
research and teaching product. The past 25–50 years have
witnessed that process both in organization studies and in strategy.
But legitimacy gained by standardization simultaneously reduces the
stimulation of novelty and the amalgamation of ideas from other
fields, which have been some of the main contributions of behavioral
strategy. On the whole, it seems likely that the pursuit of legitimacy
and standing is more likely to inhibit than to stimulate exploration.
There is no special magic associated with behavioral strategy
studies. Most conspicuously, strategy is a field that is unavoidably
linked to the giving of advice, a practice that is often (though not
always) reviled in studies of organizations. Many students of
organizations treasure their roles as scientific observers, rather than
engineers. They provide a list of more or less established ways of
looking at behavioral strategy. Organizations do what they do
because of intentional, but bounded, rationality, because of survival
and replication, because of learning through trial and error, and
because of networks of imitation and diffusion. Each of these
provides a perspective for understanding behavioral strategy and a
frame for integrating studies of behavioral strategy and behavioral
studies of decision-making, an integration that will strengthen the
academic position of behavioral strategy insofar as the studies of
decision-making are recognized generally as providing progress.
Neither history nor good sense, however, suggests that
exploration in behavioral strategy will be, or should be, limited to
studies of decision-making. It is almost certainly too early to close
down exploration.

RATIONALIZING THE LEVEL OF FOOLISHNESS

The tension between exploitation and exploration, and the apparent
tendency for the pursuit of short-run success to favor exploitation
that effectively limits exploration has led to various efforts to
rationalize the level of exploration. It is argued that rationality
requires elements of nonrationality to assure experiments with
novelty, and that the precise character and amount of nonrationality
can be determined rationally. The interpretation is tempting, for it
seeks to specify the gains and losses from foolish exploration and
invites analyses that seek to determine the balance point between
foolishness and sensibility in the names of intelligence and creativity.
It seems inevitable that the conceits of rationality will lead us to
try to specify the optimum time, place, and magnitude of foolishness
in the name of intelligence. However, those who have tried to define
and impose optimal foolishness have persistently discovered that
both the definition and the imposition are problematic. In the end,
the attempt to rationalize and operationalize foolishness may be an
unfortunate manifestation of a consequentialist paradigm. When
Sancho Panza challenges Don Quixote to provide a consequential
justification for his foolishness, Quixote (I, 25) replies: “Que volverse
loco un caballero andante con causa, ni grado ni gracias.El toque
está en desatinar sin ocasión.” “For a knight errant to make himself
crazy for a reason merits neither credit nor thanks. The point is to be
foolish without justification.” For Quixote, anticipating Max Weber,
rationality is an iron cage in which humans are enslaved by reasons
imposed by their environments. A free soul ignores rationality in the
name of an identity without any attempt to justify the action in
consequential ways.
It is possible that the human condition is better served by
Quixote’s exhortation to make foolishness an obligation of
humanness, not an action justified by its consequences. And if,
occasionally, it turns out that such foolishness serves intelligence or
progress, that happy happenstance is to be acknowledged with
wonder and gratitude, not pride. As behavioral strategy seeks to
discover and shape its destiny, it may want to consider foolishness
without justification as an element of its program, but perhaps only
after reading Don Quixote often enough to appreciate both the
reality and the irrelevance of the downside of foolishness.

ACKNOWLEDGMENT
I am grateful for the comments of Mie Augier and Roderic March.

NOTES
1. Including anthropologists Clyde Kluckhohn, Florence Rockwood
Kluckhohn, Evon Z. Vogt, Beatrice Blyth Whiting, and John Whiting;
psychologists Gordon Allport, Roger Brown, Jerome Bruner, Gardner
Lindzey, Eleanor Maccoby, and Robert W. White; sociologists George
Homans, Alex Inkeles, Seymour Martin Lipset, Barrington Moore,
Talcott Parsons, David Riesman, Charles Tilly, and Harrison C. White.
2. Including anthropolgists Roy G. D’Andrade, Carol R. Ember,
Clifford Geertz, Michelle Zimbalist Rosaldo, Renato Rosaldo, Richard
A. Shweder, and Marc J. Swartz; psychologists Bertram J. Cohler,
Janellen Huttenlocher, Edward E. Jones, Leon Kamin, Nathan Kogan,
Jean Lipman-Blumen, Jean Mandler, Dan P. McAdams, Stanley
Milgram, Barbara Rogoff, Abby Stewart, and Michael Wallach;
sociologists Robert N. Bellah, Zelda Gamson, Harold Garfinkel, Mark
Granovetter, Edward O. Laumann, Neil Smelser, Fred L. Strodtbeck,
Charles Tilly, and Barry Wellman.

REFERENCES
Cyert, R. M., & March, J.G. (1963). A Behavioral Theory of the Firm.
Englewood Cliffs, N.J: Prentice Hall.
March, J. G. (1994). A Primer on Decision Making: How Decisions
Happen. New York, NY: The Free Press.
Nelson, R., & Winter, S. (1982). An Evolutionary Theory of Economic
Change. Cambridge, MA: Harvard University Press.
Von Neumann, J., & Morgenstern, O. (1944). The Theory of Games
and Economic Behavior. Princeton, NJ: Princeton University
Press.
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The affinities with the order Stolonifera are clearly seen in the genera
Xenia and Telesto. Some species of Xenia form flattened or domed
colonies attached to stones or corals, with non-retractile anthocodiae
and body-walls united for only a short distance at the base. Young
Xenia colonies are in fact Stolonifera in all essential characters. In
Telesto prolifera we find a network of stolons encrusting coral
branches and other objects after the manner of the stolons of many
species of Clavularia, although the zooids do not arise from these
stolons singly, but in groups, with their body-walls fused together for
a certain distance. In Telesto rubra the spicules of the body-walls are
fused together to form a series of perforated tubes very similar in
some respects to the tubes of Tubipora.

A remarkable genus is Coelogorgia. Here we find a branching colony

arising from a basal stolon, and the axis of the main stem and of
each branch consists of a single very much elongated zooid bearing
on its thickened walls the branches of the next series and other
zooids. It is true that in this genus there is very little fusion of
neighbouring zooids, and the amount of true coenenchym is so small
that it can hardly be said to exist at all. Bourne[373] has united this
genus with Telesto into a family Asiphonacea, which he joins with the
Pennatulida in the order Stelechotokea; but their affinities seem to
be closer with the Alcyonacea than with the Pennatulacea, from
which they differ in many important characters.
 Fig. 154.—Alcyonium digitatum, a single-lobed specimen, with some of the
zooids expanded.

The genus Alcyonium not only contains the commonest British

Alcyonarian (A. digitatum), but it is one of the most widely distributed
genera of all Alcyonaria that occur in shallow water.

The genera Sarcophytum and Lobophytum occur in shallow water in

the tropics of the old world. The former frequently consists of huge
toad-stool shaped masses, soft and spongy in consistency, of a
green, brown, or yellow colour. On some reefs the colonies of
Sarcophytum form a very conspicuous feature, and from their very
slimy, slippery surface, add to the minor dangers of wading in these
regions. Both genera are dimorphic. Some species of the genus
Sclerophytum,[374] which occur in the Indian Ocean, are so hard and
brittle that they might readily be mistaken for a Zoantharian coral.
This character is due to the enormous number of tightly packed
spicules borne by the coenenchym. Some of these spicules in S.
querciforme are 7 mm. × 1.7 mm.; the largest, though not the longest
(vide p. 335) of any spicules occurring in the order.

Another very important genus occurring on coral reefs, and of very

wide distribution, is Spongodes. This genus forms bushy and rather
brittle colonies of an endless variety of beautiful shapes and colours.
Arising from the neck of each anthocodia there are one or two long,
sharp, projecting spicules, which give the surface a very spiny or
prickly character.

The genera Siphonogorgia and Chironephthya form large brittle,

branching colonies which might readily be mistaken for Gorgonians.
The strength of the branches, however, is mainly due to the large,
densely packed, spindle-shaped spicules at the surface of the
coenenchym, the long coelenteric cavities of the zooids penetrating
the axis of both stem and branches. Siphonogorgia is usually
uniformly red or yellow in colour. Chironephthya, on the other hand,
exhibits a great variety of colour in specimens from the same reef,
and indeed in different branches of the same colony.

Fam. 1. Xeniidae.—Alcyonacea with non-retractile zooids. Spicules

very small discs, usually containing a relatively small proportion of
lime.

Xenia, Savigny; Indian Ocean and Torres Straits. Heteroxenia,

Kölliker; Red Sea, Cape of Good Hope, and Torres Straits.

Fam. 2. Telestidae.—Colonies arising from an encrusting

membranous or branching stolon. The erect stem and branches are
formed by the body-walls of two or three zooids only, from which
secondary zooids and branches of the next order arise.

Telesto, Lamouroux, widely distributed in warm waters of the

Atlantic, Pacific, and Indian Oceans. The genus Fascicularia, Viguier,
from the coast of Algiers, seems to be related to Telesto, but the
groups of zooids are short, and do not give rise to branches.

Fam. 3. Coelogorgiidae.—The colony arborescent, attached by

stolon-like processes. The stem formed by an axial zooid with
thickened body-walls. Branches formed by axial zooids of the
second order, and branchlets by axial zooids of the third order, borne
either on two sides or in spirals by the main stem. Genus
Coelogorgia, Zanzibar.

Fam. 4. Alcyoniidae.—The colonies of this family are usually soft

and fleshy, and the spicules, evenly distributed throughout the
coenenchym, do not usually fuse or interlock to form a continuous
solid skeleton. They may be unbranched or lobed, never dendritic in
form. The principal genera are:—Alcyonium, Linnaeus,
cosmopolitan, but principally distributed in temperate and cold
waters. Alcyonium digitatum is the commonest British Alcyonarian. It
is found in shallow water, from the pools left at low spring tides to
depths of 40 or 50 fathoms, at most places on the British shores. It is
stated by Koehler to descend into depths of over 300 fathoms in the
Bay of Biscay. There are two principal varieties; one is white or pale
pink in the living condition, and the other yellow. In some localities
the two varieties may be found in the same pools. Another species,
Alcyonium glomeratum, placed in a distinct genus (Rhodophyton) by
Gray, and distinguished from the common species by its red colour
and long digitate lobes, is found only off the coast of Cornwall.
Paralcyonium, Milne Edwards; Mediterranean. Sclerophytum, Pratt;
sometimes dimorphic, Indian Ocean. Sarcophytum, Lesson;
dimorphic, principally tropical. Lolophytum, Marenzeller; dimorphic,
tropical. Anthomastus, Verrill; dimorphic, Atlantic Ocean, deep water.
Acrophytum, Hickson; dimorphic, Cape of Good Hope.

Fam. 5. Nephthyidae.—Colonies dendritic. Usually soft and flexible

in consistency. Nephthya, Savigny; Indian and Pacific Oceans.
Spongodes, Lesson; widely distributed in the Indian and Pacific
Oceans.

Fam. 6. Siphonogorgiidae.—Colonies often of considerable size.

Dendritic. Spicules usually large and abundant, giving a stiff, brittle
consistency to the stem and branches. Siphonogorgia, Kölliker; Red
Sea, Indian, and Pacific tropics. Chironephthya, Wright and Studer;
Indian and Pacific Oceans. Lemnalia, Gray; Zanzibar. Agaricoides,
Simpson;[375] Indian Ocean, 400 fathoms.

Order IV. Gorgonacea.

This order contains a very large number of dendritic and usually
flexible corals occurring in nearly all seas and extending from
shallow waters to the very great depths of the ocean. A large
proportion of them are brightly coloured, and as the principal
pigments are fixed in the spicules, and are therefore preserved when
the corals are dead and dried, they afford some of the most
attractive and graceful objects of a natural history museum.
The only character that separates them from the Alcyonacea is that
they possess a skeletal axis that is not perforated by the coelenteric
cavities of the zooids. The coelenteric cavities are usually short. The
order may conveniently be divided into two sub-orders.

Sub-Order 1. Pseudaxonia.
The axis in this sub-order consists of numerous spicules tightly
packed together, or cemented together by a substance which is
probably allied to horn in its chemical composition. This substance
may be considerable in amount, in which case it remains after
decalcification as a spongy, porous residue; or it may be so small in
amount, as in Corallium, that the axis appears to be composed of
solid carbonate of lime. The statement is usually made that the axis
is penetrated by nutritive canals in certain genera, but the evidence
upon which this is based is unsatisfactory and in some cases
unfounded. There can be no doubt, however, that in some genera
the axis is porous and in others it is not, and this forms a useful
character for the separation of genera.

Fam. 1. Briareidae.—The medullary substance consists of closely

packed but separate spicules embedded in a soft horny matrix,
which is uniform in character throughout its course. Nearly all the
genera form dendritic colonies of considerable size.

The principal genera are:—Solenocaulon, Gray; Indian Ocean and

North Australia. Many of the specimens of this genus have fistulose
stems and branches. The tubular character of the stem and
branches is probably caused by the activity of a Crustacean,
Alpheus, and may be regarded as of the nature of a gall-formation.
[376] Paragorgia, M. Edwards; Norwegian fjords, in deep water. This
genus forms very large tree-like colonies of a ruby-red or white
colour. It is perhaps the largest of the dendritic Alcyonarians. It is
dimorphic. Spongioderma, Kölliker; Cape of Good Hope. The surface
of this form is always covered by an encrusting sponge. Iciligorgia,
Ridley; Torres Straits. The stem and branches are compressed and
irregular in section.

Fam. 2. Sclerogorgiidae.—The medullary mass forms a distinct

axis consisting of closely packed elongate spicules with dense horny
sheaths.

Suberogorgia, Gray, has a wide distribution in the Pacific Ocean,

Indian Ocean, and the West Indies. Keroeides, W. and S., comes
from Japan.

Fam. 3. Melitodidae.—The axis in this family exhibits a series of

nodes and internodes (Fig. 155), the former consisting of pads
formed of a horny substance with embedded spicules, the latter of a
calcareous substance with only traces of a horny matrix. The
internodes are quite rigid, the nodes however give a certain degree
of flexibility to the colony as a whole. Neither the nodes nor the
internodes are penetrated by nutritive canals, but when dried the
nodes are porous.
 Fig. 155.—Melitodes dichotoma, showing the swollen nodes and the internodes.

The principal genera are:—Melitodes, Verrill; widely distributed in the

Indian and Pacific Oceans, Cape of Good Hope, etc. This genus is in
some localities extremely abundant and exhibits great brilliancy and
variety of colour. The branching is usually dichotomous at the nodes.
Wrightella, Gray. This is a delicate dwarf form from Mauritius and the
coast of South Africa. Parisis, Verrill; Pacific Ocean from Formosa to
Australia but not very common. One species from Mauritius. The
branches arise from the internodes.

Fam. 4. Coralliidae.—The axis is formed by the fusion of spicules

into a dense, solid, inflexible, calcareous core.

Corallium, Lamarck. Corallium nobile, Pallas, the "precious coral,"

occurs in the Mediterranean, chiefly off the coast of North Africa, but
also on the coasts of Italy, Corsica, Sardinia, and it extends to the
Cape Verde Islands in the Atlantic Ocean. C. japonicum, Kishinouye,
called Akasango by the fishermen, occurs off the coast of Japan,
and C. reginae, Hickson, has recently been described from deep
water off the coast of Timor.[377] The genus Pleurocorallium, Gray, is
regarded by some authors as distinct, but the characters that are
supposed to distinguish it, namely, the presence of peculiar "opera-
glass-shaped spicules," and the occurrence of the verrucae on one
side of the branches only, are not very satisfactory. The following
species are therefore placed by Kishinouye[378] in the genus
Corallium:—C. elatius, Ridley (Momoirosango); C. konojoi,
Kishinouye (Shirosango); C. boshuensis, K.; C. sulcatum, K.; C.
inutile, K.; and C. pusillum, K.,—all from the coast of Japan. Of the
coral obtained from these species, the best kinds of Momoirosango
vary in price from £30 per pound downwards according to the quality.
The Shirosango is the least valuable of the kinds that are brought
into the market, and is rarely exported.[379] Three species of
Corallium (Pleurocorallium) have been described from Madeira,[380]
and one of these, C. johnsoni, has recently been found in 388
fathoms off the coast of Ireland.[381] Other species are C.
stylasteroides, from Mauritius; C. confusum, Moroff,[382] from
Sagami Bay in Japan; and an undescribed species obtained by the
"Siboga," off Djilolo. These corals range from shallow water to
depths of 300-500 fathoms. Pleurocoralloides, Moroff, differs from
the others in having very prominent verrucae and in the character of
the large spindle-shaped and scale-like spicules. It was found in
Sagami Bay, Japan. Specimens attributed to the genus
Pleurocorallium have been found fossil in the white chalk of France,
but Corallium has been found only in the tertiaries.[383]

Sub-Order 2. Axifera.
The axis in this sub-order may be horny, or horny with a core of
calcium carbonate, or composed of horn impregnated with calcium
carbonate, or of nodes of horn alternating with internodes of calcium
carbonate. It may be distinguished from the axis of the Pseudaxonia
by the fact that in no case have definite spicules been observed to
take part in its formation. It has been suggested that as the Axifera
represent a line of descent distinct from that of the Pseudaxonia they
should be placed in a separate order. Apart from the character of the
axis, however, the two sub-orders show so many affinities in their
general anatomy that it is better to regard the two lines of descent as
united within the Gorgonacean limit. It is very improbable that the
two groups sprang independently from a stoloniferous ancestor.
Fam. 1. Isidae.—This family includes all those Axifera in which the
axis is composed of alternate nodes of horn and internodes of
calcareous substance.

There can be little doubt of the close affinities of many of the genera
of this family with the Melitodidae among the Pseudaxonia. In both
the coenenchym is thin and the coelenteric cavities short. No
important differences have been observed between the structure of
the zooids of the two families, and now that we know that the
"nutritive canals" of Melitodes do not perforate the nodes there is no
important difference left between the coenosarcal canal systems.
The structure and method of calcification of the internodes of the two
families are very similar. The main difference between them is that
the nodes of the Isidae are purely horny, whereas in the Melitodidae
the horny substance of the nodes contains calcareous spicules.

The principal genera are:—Isis, Linnaeus; Pacific Ocean. This genus

forms substantial fan-shaped colonies with, relatively, a thick
coenenchym, short stout internodes and black horny nodes.
Mopsea, Lamouroux; Coast of Australia. The verrucae are club-
shaped and are arranged in spiral rows round the stem. Acanella,
Gray; principally found in deep water in the Atlantic Ocean but also
in the Pacific. The internodes are long and the branches arise from
the nodes. Most of the species occur in deep water, some in very
deep water (A. simplex, 1600 to 1700 fathoms). In this and the
following genera the coenenchym is thin and the zooids imperfectly
or not retractile. Ceratoisis, Wright; Atlantic Ocean, extending from
shallow to deep water. The branches arise from the nodes.
Chelidonisis, Studer; deep water off the Azores. Isidella, Gray;
Mediterranean Sea. Bathygorgia, Wright; off Yokohama, 2300
fathoms. This genus is unbranched, with very long internodes and
short nodes. The zooids are arranged on one side only of the stem.

Fam. 2. Primnoidae.—This is a well-marked family. The axis of the

colonies is horny and calcareous. The coenenchym and the non-
retractile zooids are protected by scale-like spicules, which usually
overlap and form a complete armour for the protection of the soft
parts. On the aboral side of the base of each tentacle there is a
specialised scale, and these fit together, when the tentacles are
folded over the peristome, to form an operculum.

The principal genera are:—Primnoa, Lamouroux; Atlantic Ocean,

occurring also in the Norwegian fjords. This genus is usually found in
moderately deep water, 100 to 500 fathoms. Primnoella, Gray. This
genus seems to be confined to the temperate seas of the southern
hemisphere. It is unbranched. The zooids are arranged in whorls
round the long whip-like stem. Plumarella, Gray; southern
hemisphere, in moderately deep water. This is branched pinnately in
one plane. The zooids are small and arise at considerable intervals
alternately on the sides of the branches. Stenella, Gray; widely
distributed in deep water. The zooids are large and are arranged in
whorls of three situated at considerable distances apart. Stachyodes,
W. and S.; Fiji, Kermadecs, Azores, in deep water. Colony feebly
branched. Zooids in regular whorls of five. Other genera belonging to
this group of Primnoidae are Thouarella, Gray, and Amphilaphis,
Antarctic seas.

The following genera are placed in separate sub-families:—

Callozostron, Wright; Antarctic Sea, 1670 fathoms. The axis is
procumbent and the zooids are thickly set in rows on its upper
surface. The zooids are protected by large imbricate scales, of which
those of the last row are continued into long spine-like processes.
Calyptrophora, Gray; Pacific Ocean, in deep water. The base of the
zooids is protected by two remarkably large scales. Primnoides, W.
and S.; Southern Ocean. The opercular scales are not distinctly
differentiated and the calyx is therefore imperfectly protected.

Fam. 3. Chrysogorgiidae.[384]—The axis in this family is composed

of a horny fibrous substance with interstratified calcareous particles,
and it springs from a calcareous plate, which sometimes gives off
root-like processes. It may be unbranched or branched in such a
way that the branches of the second, third, and subsequent orders
assume in turn the direction of the base of the main axis. The axis is
frequently of a metallic iridescent appearance. The zooids usually
arise in a single straight or spiral row on the branches, and are not
retractile. The coenenchym is thin. The spicules vary considerably,
but in a very large proportion of the species they are thin, oval, or
hour-glass plates (Fig. 149, 10, p. 336).

By some authors this family is considered to be the simplest and

most primitive of the Axifera; but the delicate character of the axis of
the main stem and branches, the thinness of the coenenchym, the
position of the zooids on one side of the branches only, and the
tenuity of the calcareous spicules may be all accounted not as
primitive characters, but as special adaptations to the life in the slow
uniform currents of deep water.

The principal genera are:—Lepidogorgia, Verrill; Atlantic and Pacific

Oceans, 300 to 1600 fathoms. Axis unbranched. Zooids large and
arranged in a single row. Trichogorgia, Hickson; Cape of Good Hope,
56 fathoms. Colony branching in one plane. Zooids numerous and
on all sides of the branches. Chrysogorgia, D. and M.; deep water.
Axis branched. Spicules on the zooids always large. Metallogorgia,
Versluys; Atlantic Ocean, 400 to 900 fathoms. Basal part of the stem
unbranched (monopodial). Iridogorgia, Verrill. Spiral stem and
branches. Pleurogorgia, Versluys. Axis branched in one plane.
Coenenchym thick. Riisea, D. and M. Monopodial stem and thick
coenenchym.

Fam. 4. Muriceidae.—This is a large family, exhibiting very great

variety of habit. The spicules are often very spiny, and project
beyond the surface of the ectoderm, giving the colony a rough
appearance. A great number of genera have been described, but
none of them are very well known. The family requires careful
revision.

The more important genera are:—Acanthogorgia, Gray; principally in

deep water in the Atlantic Ocean. The calices are large, cylindrical,
and spiny. Villogorgia, D. and M.; widely distributed. Delicate,
graceful forms, with thin coenenchym. Echinomuricea, Verrill;
Muricea, Lamouroux; Paramuricea, Köll; Acamptogorgia, W. and S.;
Bebryce, Philippi.

Fam. 5. Plexauridae.—In this family we find some of the largest and

most substantial Gorgonids. The axis is usually black, but its horny
substance may be impregnated with lime, particularly at the base.
The coenenchym is thick, and the zooids are usually completely
retractile, and the surface smooth. The species of the family are
principally found in shallow water in warm or tropical regions.

The principal genera are:—Eunicea, Lamouroux. The calices are

prominent, and not retractile. Plexaura, Lamouroux; Euplexaura,
Verrill. Eunicella, Verrill. With an outer layer of peculiar torch-shaped
spicules. The only British species of this order is Eunicella cavolini
(formerly called Gorgonia verrucosa). It is found in depths of 10 to 20
fathoms off the coast of the English Channel and west of Scotland.
Occasionally specimens are found in which a gall-like malformation
with a circular aperture is seen, containing a Barnacle. Such gall
formations, common enough in some species of Madreporaria, are
rarely found in Alcyonaria.

Fig. 156.—Eunicella cavolini. Some branches of a large dried specimen, showing

a gall formed by a Cirripede.

Fam. 6. Gorgoniidae.—This family contains some of the

commonest and best-known genera of the order. They usually form
large flexible branched colonies with delicate horny axes and thin
coenenchym. The zooids are usually completely retractile.

The principal genera are:—Gorgonia, Linn. This genus includes

Gorgonia (Rhipidogorgia) flabellum, the well-known fan Gorgonia
with intimately anastomosing branches, from the warm waters of the
Atlantic Ocean. The genera Eugorgia, Verrill, and Leptogorgia, Milne
Edwards, differ from Gorgonia in the character of the spicules. In
Xiphigorgia, Milne Edwards, from the West Indies, the branches are
much compressed, forming at the edges wing-like ridges, which bear
the zoopores in rows. Malacogorgia, Hickson, has no spicules. Cape
of Good Hope.

Fam. 7. Gorgonellidae.—In this family the horny axis is

impregnated with lime. The surface of the coenenchym is usually
smooth, and the spicules small. The colonies are sometimes
unbranched (Juncella). In the branching forms the axis of the
terminal branches is often very fine and thread-like in dimensions.

Fig. 157.—Verrucella guadaloupensis, with an epizoic Brittle star (Oph.) of similar

colour.

The principal genera are:—Gorgonella, with a ramified flabelliform

axis; Ctenocella, with a peculiar double-comb manner of branching;
and Juncella, which forms very long unbranched or slightly branched
colonies, with club-shaped spicules. All these genera are found in
shallow water in the tropical or semi-tropical regions of the world.
Verrucella is a genus with delicate anastomosing branches found
principally in the shallow tropical waters of the Atlantic shores. Like
many of the Gorgonacea, with branches disposed in one plane
(flabelliform) Verrucella frequently carries a considerable number of
epizoic Brittle stars, which wind their flexible arms round the
branches, and thus obtain a firm attachment to their host. There is
no reason to suppose that these Brittle stars are in any sense
parasitic, as a specimen that bears many such forms shows no sign
of injury or degeneration, and it is possible they may even be of
service to the Verrucella by preying upon other organisms that might
be injurious. An interesting feature of the association is that the
Brittle stars are of the same colour as the host, and the knob-like
plates on their aboral surface have a close resemblance to the
verrucae (Fig. 157).

Order V. Pennatulacea.
The Sea-pens form a very distinct order of the Alcyonaria. They are
the only Alcyonarians that are not strictly sedentary in habit, that are
capable of independent movement as a whole, and exhibit a bilateral
symmetry of the colony. No genera have yet been discovered that
can be regarded as connecting links between the Pennatulacea and
the other orders of the Alcyonaria. Their position, therefore, is an
isolated one, and their relationships obscure.

The peculiarities of the order are due to the great growth and
modification in structure of the first formed zooid of the colony. This
zooid (Oozooid, Hauptpolyp, or Axial zooid) increases greatly in
length, develops very thick fleshy walls, usually loses its tentacles,
digestive organs, and frequently its mouth, exhibits profound
modification of its system of mesenteries, and in other ways
becomes adapted to its function of supporting the whole colony.
 Fig. 158.—Diagram of a Sea-pen. L, leaves composed of a row of autozooids; R,
rachis; St, stalk; T, anthocodia of the axial zooid, usually suppressed. (After
Jungersen.)

The axial zooid shows from an early stage of development a division

into two regions: a distal region which produces by gemmation on
the body-wall numerous secondary zooids, and becomes the rachis
of the colony; and a proximal region which becomes the stalk or
peduncle, and does not produce buds (Fig. 158). The secondary
zooids are of two kinds: the autozooids and the siphonozooids. The
former have the ordinary characters of an Alcyonarian zooid, and
produce sexual cells; the latter have no tentacles, a reduced
mesenteric system, and a stomodaeum provided with a very wide
siphonoglyph.

The arrangement of the autozooids and siphonozooids upon the

axial zooid is subject to great modifications, and affords the principal
character for the classification of the order. In the Pennatuleae the
autozooids are arranged in two bilaterally disposed rows on the
rachis, forming the leaves or pinnae of the colony (Fig. 158). The
number in each leaf increases during the growth of the colony by the
addition of new zooids in regular succession from the dorsal to the
ventral side of the rachis[385] (Fig. 159). In other Pennatulacea the
autozooids are arranged in rows which do not unite to form leaves
(Funiculina), in a tuft at the extremity of a long peduncle (Umbellula),
scattered on the dorsal side of the rachis (Renilla, Fig. 160), or
scattered on all sides of the rachis (Cavernularia, Fig. 161). In those
forms in which the autozooids are scattered the bilateral symmetry of
the colony as a whole becomes obscured. The siphonozooids may
be found on the leaves (Pteroeides), but more frequently between
the leaves or rows of autozooids, or scattered irregularly among the
autozooids. Usually the siphonozooids are of one kind only, but in
Pennatula murrayi there is one specially modified siphonozooid at
the base of each leaf,[386] which appears to have some special but
unknown function.

Fig. 159.—Diagram of a portion of a rachis of a Sea-pen, aut, The rows of

autozooids; 1-6, the order of age of the autozooids composing a leaf; D, the
dorsal side of the rachis; Si, the siphonozooids; V, the ventral side of the
rachis. (After Jungersen.)

In Umbellula gracilis each siphonozooid bears a single pinnate

tentacle, and in some other species of the same genus there is a
tentacle which is not pinnate.[387]

The zooids and coenenchym are usually protected by a crust of

coloured or colourless, long, smooth, needle-like, calcareous
spicules, situated principally in the superficial layer, so as to leave
the subjacent tissues soft and spongy in texture. In some cases the
spicules are smooth double clubs, rods, discs, or irregular granules,
and in Sarcophyllum, Chunella, some species of Umbellula and
others, there is no calcareous skeleton. The tuberculated spindles,
so common in other Alcyonaria, are not found in any species. In
most genera a horny, or calcified horny rod is embedded in the
central part of the axial polyp, serving as a backbone or support for
its muscles. It is absent, however, in Renilla, and reduced or absent
in Cavernularia.
The sexual organs are borne by the mesenteries of the autozooids
only, and each colony is either male or female. There is no record of
hermaphroditism in the order. The eggs contain a considerable
amount of yolk, and fertilisation is effected in the sea-water after their
discharge. The segmentation is irregular, and the free-swimming
ciliated larva (of Renilla) shows the rudiments of the first buds from
the axial polyp before it settles down in the mud.

The Sea-pens are usually found on muddy or sandy sea-bottoms,

from a depth of a few fathoms to the greatest depths of the ocean. It
is generally assumed that their normal position is one with the
peduncle embedded in the mud and the rachis erect. Positive
evidence of this was given by Rumphius, writing in 1741, in the case
of Virgularia rumphii and V. juncea at Amboina,[388] and by Darwin in
the case of Stylatula darwinii at Bahia Blanca.[389]

"At low water," writes Darwin, "hundreds of these zoophytes might be

seen projecting like stubble, with the truncate end upwards, a few
inches above the surface of the muddy sand. When touched or
pulled they suddenly drew themselves in with force so as nearly or
quite to disappear."

It is not known whether the Pennatulids have the power of moving

from place to place when the local conditions become unfavourable.
It is quite probable that they have this power, but the accounts given
of the Sea-pens lying flat on the sand do not appear to be founded
on direct observation. The fable of Pennatula swimming freely "with
all its delicate transparent polypi expanded, and emitting their usual
brilliant phosphorescent light, sailing through the still and dark abyss
by the regular and synchronous pulsations of the minute fringed
arms of the whole polypi," appears to be based on a statement made
by Bohadsch in 1761, and picturesque though it be, is undoubtedly
erroneous.
The brilliant phosphorescence of many species of Pennatulacea has
been observed by many naturalists, and it is very probable that they
all exhibit this property to some degree. The phosphorescence
appears to be emitted by the mesenteric filaments of the autozooids,
but it is not yet determined whether the phenomenon is confined to
these organs or is more generally distributed.

The Pennatulacea are usually devoid of epizoites, but occasionally

the parasitic or semi-parasitic Entomostracan Lamippe is found in
the zooids. A small crab is also frequently found between the large
leaves of species of Pteroeides. The most remarkable case of
symbiosis, however, has recently been observed in the form of an
encrusting Gymnoblastic Hydroid[390] living on the free edge of the
leaves of a species of Ptilosarcus.

The order Pennatulacea is divided into four sections.

Sect. 1. Pennatuleae.—In this section the colony is distinctly

bilaterally symmetrical, and the autozooids are arranged in rows with
their body-walls fused to form leaves.

The genus Pteroeides, the representative genus of the family

Pteroeididae, is a fleshy Sea-pen found in shallow sea water in the
warm waters of the Pacific Ocean and in the Mediterranean. It has
large leaves with long spiny, projecting spicules, and the
siphonozooids are borne by the leaves. Pennatula, the
representative genus of the family Pennatulidae, has a wider
distribution in area and in depth. Pennatula phosphorea is a common
British species, found in depths of 10 to 20 fathoms in many
localities off our coasts. It is about 5 inches in length. There are
several varieties of this species distributed in Atlantic waters.
Pennatula grandis is a magnificent species found in Norwegian
fjords, in the Faeroe Channel, and off the northern coasts of N.
America, in depths of from 50 to 1255 fathoms. Specimens have
been obtained no less than 2½ feet in length. P. murrayi and P.
naresi are species of the genus found at depths of a few hundred
fathoms in tropical seas.

The genus Virgularia, belonging to the family Virgulariidae, is

represented in the British seas by V. mirabilis, a long slender Sea-
pen found in many localities off the Scottish coasts.

Sect. 2. Spicatae.—This section includes those Sea-pens in which

the autozooids are arranged bilaterally on the axial zooid in rows or
more irregularly, but do not unite to form leaves. It is a large section
and contains many widely divergent genera.

The family Funiculinidae is represented on our coasts by Funiculina

quadrangularis, a long and slender Sea-pen 2 to 3 feet in length. The
autozooids are arranged in oblique rows, and the siphonozooids are
on the ventral side of the rachis. There is one point of special interest
in this genus. The siphonozooids appear to change as the colony
grows and to become autozooids. If this is the case it may be more
correct to describe the genus as devoid of true siphonozooids.

The family Anthoptilidae contains the species Anthoptilum

grandiflorum, which has a wide distribution in depths of 130 to 500
fathoms in the N. and S. Atlantic Ocean. It is perhaps the largest of
all the Pennatulacea, specimens having been obtained from the
Cape of Good Hope over 4 feet long with expanded autozooids,
each more than half an inch in length.

The family Kophobelemnonidae contains a number of forms with

remarkably large autozooids arranged in irregular rows on the two
sides of the rachis. The siphonozooids are numerous and scattered,
and their position is indicated by small papilliform calices on the
coenenchym. The surface of these pens is usually rough, owing to
the presence of numerous coarse projecting spicules.
Kophobelemnon occurs in the Mediterranean in deep water, off the
coasts of Ireland and Scotland, and in other regions.

The family Umbellulidae contains some of the most remarkable and

interesting examples of the deep-sea fauna. The peduncle is very
long and the rachis stunted and expanded. The autozooids are of
great size, non-retractile, and arranged in a cluster or rosette on the
terminal rachis. There is a wide structural range between the
species. Some species have numerous large spicules, others have
none. In some species the siphonozooids have a single pinnate or
digitate tentacle, in others the siphonozooids are of the usual type.
Umbellula appears to be a somewhat rare but cosmopolitan genus in
deep water, extending from the Arctic to the Antarctic region in water
ranging from 200 to 2500 fathoms.

The interesting genus Chunella was discovered by the German

"Valdivia" Expedition at a depth of about 420 fathoms off the coast of
E. Africa, and subsequently by the Dutch "Siboga" Expedition at a
depth of about 500 fathoms in the Malay Archipelago. According to
Kükenthal,[391] this genus with another closely allied genus
Amphianthus should form a new section of Pennatulacea, the
Verticilladeae. Chunella has a long and very delicate rachis and
peduncle, and the former terminates in a single autozooid and has
five or six whorls of three autozooids, situated at considerable
distances from one another. Spicules are absent. The full description
of this genus has not yet been published, but it is clear that it
occupies a very isolated position in the order.

Fig. 160.—Renilla reniformis, a small specimen (34 mm.), showing the dorsal
side of the expanded rachis. A, autozooid; H, the mouth of the axial zooid;