Towards a Theory of Ecosystems

Abstract

Research Summary
The recent surge of interest in “ecosystems” in strategy research and practice has mainly focused on what ecosystems are and how they operate. We complement this literature by considering when and why ecosystems emerge, and what makes them distinct from other governance forms. We argue that modularity enables ecosystem emergence, as it allows a set of distinct yet interdependent organizations to coordinate without full hierarchical fiat. We show how ecosystems address multilateral dependences based on various types of complementarities ‐ supermodular or unique, unidirectional or bidirectional, which determine the ecosystem's value‐add. We argue that at the core of ecosystems lie non‐generic complementarities, and the creation of sets of roles that face similar rules. We conclude with implications for mainstream strategy and suggestions for future research.

Managerial summary
We consider what makes ecosystems different from other business constellations, including markets, alliances or hierarchically managed supply chains. Ecosystems, we posit, are interacting organizations, enabled by modularity, not hierarchically managed, bound together by the non‐redeployability of their collective investment elsewhere. Ecosystems add value as they allow managers to coordinate their multilateral dependence through sets of roles that face similar rules, thus obviating the need to enter into customized contractual agreements with each partner. We explain how different types of complementarities (unique or supermodular, generic or specific, uni‐ or bi‐directional) shape ecosystems, and offer a “theory of ecosystems” that can explain what they are, when they emerge and why alignment occurs. Finally, we outline the critical factors affecting ecosystem emergence, evolution, and success ‐‐ or failure.

Full text

Towards a Theory of Ecosystems
Michael G. Jacobides, London Business School, mjacobides@london.edu *
Carmelo Cennamo, Bocconi University, carmelo.cennamo@unibocconi.it
Annabelle Gawer, University of Surrey, a.gawer@surrey.ac.uk
in Strategic Management Journal (2018) 39: 2255-2276
Acknowledgments: We want to thank the Editor and two anonymous Reviewers for their
detailed and constructive feedback during the revision of the manuscript. Carliss Baldwin has
provided thorough and insightful comments on multiple rounds of revision. Her suggestions,
especially on complementarities, have helped structure our argumentation. Sid Winter,
Michael Cusumano, Ron Adner, Rob Grant, Erkko Autio, Vangelis Souitaris, Brice Dattee,
Fernando Suarez and Charles Baden-Fuller also gave us valuable comments, and Tom
Albrighton helped with copy editing. We also acknowledge feedback from seminar
participants at the Wharton School, Stanford U., MIT’s Initiative for the Digital Economy,
London Business School, EPFL, Judge Business School (Cambridge U.), Cass Business
School (City U.), Warwick Business School, Bocconi U., LUISS Business School (Rome) and
participants of the Imperial College Business School Innovation and Entrepreneurship
Conversations, the Academy of Management Meetings and the Future of Mobility Paris
Meetings (ESSEC/Supelec). Jacobides acknowledges financial support from London Business
School’s RMD Fund.

Towards a Theory of Ecosystems
Abstract
The recent surge of interest in “ecosystems” in strategy research and practice has mainly focused
on what ecosystems are and how they operate. We complement this literature by considering
when and why ecosystems emerge, and what makes them distinct from other governance forms.
We argue that modularity enables ecosystem emergence, as it allows a set of distinct yet
interdependent organizations to coordinate without full hierarchical fiat. We show how
ecosystems address multilateral dependences based on various types of complementarities -
supermodular or unique, unidirectional or bidirectional, which determine the ecosystem’s
value-add. We argue that at the core of ecosystems lie non-generic complementarities, and the
creation of sets of roles that face similar rules. We conclude with implications for mainstream
strategy and suggestions for future research.
Managerial Abstract
We consider what makes ecosystems different from other business constellations, including
markets, alliances or hierarchically managed supply chains. Ecosystems, we posit, are
interacting organizations, enabled by modularity, not hierarchically managed, bound together
by the non-redeployability of their collective investment elsewhere. Ecosystems add value as
they allow managers to coordinate their multilateral dependence through sets of roles that face
similar rules, thus obviating the need to enter into customized contractual agreements with each
partner. We explain how different types of complementarities (unique or supermodular, generic
or specific, uni- or bi-directional) shape ecosystems, and offer a “theory of ecosystems” that
can explain what they are, when they emerge and why alignment occurs. Finally, we outline the
critical factors affecting ecosystem emergence, evolution, and success -- or failure.

1
INTRODUCTION
Over the last few years, there has been a surge of interest in the concept of “ecosystems” as a
new way to depict the competitive environment. Practitioners share this enthusiasm: In the
2014 prospectus for the world’s largest IPO to date, Alibaba, the word “ecosystem” appears
no fewer than 160 times. The term is entering the vocabulary not only of technology firms but
also of more established sectors, from financial services to manufacturing (Deloitte, 2015).
Beyond practice and the popular business press, ecosystems have also been eagerly adopted in
the field of strategy, with Teece (2014:1) suggesting that “the concept of ecosystem might
now substitute for the industry for performing analysis.” While ecosystems have been
considered in our field for some time (Dhanaraj & Parkhe, 2006; Iansiti & Levien, 2004;
Moore, 1993), the last few years have seen a boom. Searching the keyword “ecosystem” in the
title or abstract of the top strategy journals shows that its frequency has increased sevenfold
over the last five years.
Unarguably, this research has articulated some important competitive, collaborative, and
organizational challenges faced by firms. Adner (2017) offers a view on how ecosystem
research relates to established views, and offers a useful guide to the differences in
phenomenological emphasis between ecosystem research and other streams. This paper takes
a step further, synthesizing existing research to elucidate the key mechanisms behind the
emergence and dynamics of ecosystems, and why we have seen such a rise in interest. As a
first step toward a positive theory, we consider the conditions necessary for ecosystems to
emerge—modularity in particular—and the interactions that make them interesting—
specifically, the coexistence of different types of complementarities. We further argue that
examining the nature, directionality, and intensity of these complementarities, and what firms
do to influence them, thus shaping ecosystem formation and structure, can help explain the
distinct value creation and capture dynamics within and between ecosystems.

2
Our objective is to complement the literature’s interest in what ecosystems are, by analyzing
how and why they differ from other phenomena, and by clarifying what unique mechanisms
they have for value creation and value capture. We reflect on when we might expect to see
ecosystems displace traditional market-based arrangements or vertically integrated supply
chains. We propose that ecosystems are distinct forms of organizing economic activities that
are linked by specific types of complementarities. We also sharpen the distinction between the
structures of ecosystems and the behaviors they give rise to, building on the fundamental
distinctions between different types of complementarities to create clearly distinct sets of
ecosystems, with particular strategic dynamics. We conclude with the resulting research
agenda.
THEORETICAL BACKGROUND AND THE GROWTH OF ECOSYSTEMS
Borrowed from biology, the term “ecosystem” generally refers to a group of interacting firms
that depend on each other’s activities. Scholars have emphasized different aspects of an
ecosystem depending on the unit of analysis. In reviewing the literature, we identified three
broad groups of papers:1 a “business ecosystem” stream, which centers on a firm and its
environment; an “innovation ecosystem” stream, focused around a particular innovation or
new value proposition and the constellation of actors that support it; and a “platform
ecosystem” stream, which considers how actors organize around a platform.
1 Our literature review consisted of looking at all papers with “ecosystem” in their title and abstract, published in
the top management journals. Having read them, we created three tentative categories. These categories were
refined, and further validated, by textual analysis (available upon request) using NVivo, which confirmed the
existence of different themes and topic emphases in each of these three streams. While these categories are not
meant to be mutually exclusive and collectively exhaustive, the exercise did help create groupings, which were
further validated by asking a research assistant to assign papers into each of the categories reported in the text.

3
The first stream focuses on an individual firm or new venture2, and views the ecosystem as a
“community of organizations, institutions, and individuals that impact the enterprise and the
enterprise’s customers and supplies” (Teece, 2007: 1325). Here, the ecosystem is conceived as
an economic community of interacting actors that all affect each other through their activities,
considering all relevant actors beyond the boundaries of a single industry. For Teece (2007),
the ecosystem represents the environment that the firm must monitor and react to, which
affects its dynamic capabilities and thus its ability to build sustainable competitive advantage.
Others stress the “shared fate” of the community as a whole (Iansiti & Levien, 2004: 69)—
individual members’ performance is tied to the overall performance of the ecosystem. Despite
the emphasis on co-evolution of firm capabilities, there is little explanation of how firms
mutually adapt. Authors such as Iansiti and Levien (2004) stress the role of ecosystem
managers—“hub” or “keystone” firms—as the providers of stability. As Dhanaraj and Parkhe
(2006) argue, hub firms manage knowledge mobility, innovation appropriability, and network
stability. These mechanisms have rarely been studied (for exceptions, see Azzam et al., 2016
or Pellinen et al., 2012) and empirical support remains limited even within these studies.
The second set of studies focuses on a focal innovation and the set of components (upstream)
and complements (downstream) that support it, and views the ecosystem as “the collaborative
arrangements through which firms combine their individual offerings into a coherent,
customer-facing solution” (Adner, 2006: 98). The emphasis is on understanding how
interdependent players interact to create and commercialize innovations that benefit the end
customer—with the corollary that if coordination within the ecosystem is inadequate,
2 Studies focusing specifically on new ventures tend to see the ecosystem as a location-specific cluster of firms,
entities, and individuals; consider the new venture as the unit of interest of the analysis; and focus on new
venture creation and related entrepreneurial issues, including knowledge/innovation spillovers, growth, access to
resources, and markets (e.g., McGregor & Madsen, 2013; Pitelis, 2012; Zahra & Nambisan, 2012; Zacharakis et
al., 2003). Empirical studies in this area have mainly examined start-ups in internet, high-tech, and ICT sectors.

4
innovations will fail (e.g., Adner & Kapoor, 2010; Adner, 2012; Kapoor & Lee, 2013). Here,
the anchoring point is the system of innovations that allows customers to use the end product,
rather than the firm. Accordingly, the ecosystem concept is intended to capture the link
between a core product, its components, and its complementary products/services
(“complements”), which jointly add value for customers. The firm(s) producing the focal
innovation may or may not be directly connected to complement providers
(“complementors”); the extent to which firms align through different arrangements will affect
their capacity to create value for the end customer (Adner 2017). Here, the ecosystem casts a
net around the “virtual network[s]” (Iyer et al., 2006) or “complex entities of group-related
actors” (Brusoni & Prencipe, 2013) that offer focal and complementary innovations. Research
has considered how different collaborative arrangements between the innovator and its
complementors affect both groups’ ability to coordinate investments into a new technology
and its commercialization (e.g., Kapoor & Lee, 2013; Leten et al., 2013); how knowledge
sharing affects the strength of inter-firm relationships and thus the development of the
ecosystem (e.g., Alexy et al., 2013; Brusoni & Prencipe, 2013; Frankort, 2013); or the health
and survival of the ecosystem (Leten et al., 2013; West & Wood, 2013).
The third set of studies focuses on a specific class of technologies—platforms—and the
interdependence between platform sponsors and their complementors. In this view, the
ecosystem comprises the platform’s sponsor plus all providers of complements that make the
platform more valuable to consumers (Ceccagnoli et al., 2012: 263; Gawer & Cusumano,
2008: 28)3. The platform ecosystem takes a “hub and spoke” form, with an array of peripheral
firms connected to the central platform via shared or open-source technologies and/or
3 The platform concept has gained extensive traction in its own right, spurring a whole literature looking at the
peculiar network-externality dynamics characterizing so-called “platform two-sided markets” (e.g., Hagiu, 2006;
Parker & Van Alstyne, 2005; Rochet & Tirole, 2003). While we explicitly considered the platform in relation to
the ecosystem of specialized complementary goods/services, a broader review is outside this paper’s scope.

5
technical standards (which, for IT-related platforms, can be programming interfaces or
software development kits). By connecting to the platform, complementors can not only
generate complementary innovation, but also gain access, directly or indirectly, to the
platform’s customers—as in the examples of independent software vendors affiliating to SAP
(Ceccagnoli et al., 2012) or developers producing videogames for specific consoles (Cennamo
& Santaló, 2013). Accordingly, platform ecosystems are seen as “semi-regulated
marketplaces” that foster entrepreneurial action under the coordination and direction of the
platform sponsor (Wareham et al., 2014: 1211), or as “multisided markets” enabling
transactions between distinct groups of users (Cennamo & Santaló, 2013).4 See McIntyre and
Srinivasan (2017) for a recent critical review.
Ecosystems as new structures of economic relationships. While these views might reflect
differences in research focus, they emphasize aspects of the ecosystem that overlap in the real
world. It is broadly agreed that ecosystems require providers of complementary innovations,
products, or services, who might belong to different industries and need not be bound by
contractual arrangements—but have significant interdependence nonetheless. In this sense,
ecosystems do not fit into the classical firm-supplier relationship, Porter’s (1980) value
system, or a firm’s strategic networks; neither are they integrated hierarchies.
Studies taking the firm as the unit of interest consider the ecosystem to include the ties that the
firm has to the actors that affect, or are affected by, its activities. Those taking the innovation
as the unit of interest have considered interconnected innovations upstream (i.e., components)
4 Despite the greater consistency among ecosystem platform scholars, they still vary in terms of the elements
posited as constituting an ecosystem. While some studies examine platforms as means for complementors to
access customers (e.g., Ceccagnoli et al., 2012; Cennamo & Santaló, 2013; Cennamo, 2016; Wareham et al.,
2014), others focus more on technology evolution through the interaction between the platform owner and its
complementors, and in relation to competing ecosystems, but do not regard final customers as central to such
dynamics (e.g., Gawer & Cusumano, 2002; Gawer & Henderson, 2007).

6
and downstream (i.e., complements) in the same industry (e.g., Adner & Kapoor, 2010;
Frankort, 2013), connections running through sub-industries (e.g., Makinen & Dedehayir,
2013), firm-complementor dyads (e.g., Kapoor & Lee, 2013), or multi-party collaboration
(e.g., Leten et al., 2013; West & Wood, 2013). Studies on platform-based ecosystems have
considered connections between the platform sponsor and its complementors (Ceccagnoli et
al., 2012; Cennamo & Santaló, 2013; Gawer & Henderson, 2007) established through
standards and platform interfaces (Gawer 2014), the leadership role of the platform at the
industry level (e.g., Gawer & Cusumano, 2002), the impact of the platform’s technological
complexity on complementors’ innovation capacity (e.g., Kapoor & Agarwal 2017), or
rivalries between competing platform ecosystems (e.g., Cennamo & Santaló, 2013).
Most studies consider deliberate intent of specific actors to be important, and focus on the role
of the hub—dubbed the “lead firm” (Williamson & De Meyer, 2012), “keystone” organization
(Iansiti & Levien, 2004), or “ecosystem captain” (Teece, 2014)—in shaping the emergence of
an ecosystem (Moore 1993, Teece, 2007). According to Gulati et al. (2012), the presence of
an “architect,” who sets a system-level goal, defines the hierarchical differentiation of
members’ roles, and establishes standards and interfaces, is an essential and distinguishing
feature of an ecosystem (also see Teece, 2014).5 Studies tend to concur that ecosystems are
not hierarchically managed, but few have specifically looked at the rules governing
membership and relationships. Gulati et al. (2012) consider ecosystem membership to be
“open”—i.e., not granted bilaterally between hub and prospective member, but based on self-
selection. However, recent work paints a more nuanced picture, suggesting that formal
mechanisms, including the management of standards and interfaces (Baldwin, 2012; Teece,
5 Some contend that the hub is not necessarily the largest or most resource-rich member of the ecosystem, but
rather the one that uses “smart power” (Williamson & De Meyer, 2012), “problem framing” (Brusoni &
Prencipe, 2013), or “informal authority” based on knowledge, status, or control over key resources or technology
(Gulati et al., 2012). Also see Gawer and Phillips (2013) on “institutional work.”

7
2014), platform governance (Cennamo & Santaló, 2013; Wareham et al., 2014), or intellectual
property rights and other contractual forums, are key tools that hubs use to discipline and
motivate ecosystem members (Alexy et al, 2013; Brusoni & Prencipe, 2013; Leten et al,
2013; Ritala et al., 2013).
Research in the platform ecosystem tradition also considers how technological interfaces (and
which parts of the technology are “open” or “closed”) or governance (such as membership and
participation rules) shape collective outcomes (Ceccagnoli et al., 2012; Cennamo, 2016;
Gawer & Cusumano, 2002; Gawer, 2014; Wareham et al., 2014). Balancing the trade-offs
involved in controlling the core technology is one of the main goals of platform ecosystem
governance (Cennamo & Santaló, 2013; Cennamo, 2016; Wareham et al., 2014), also
identified as a key problem of organization design (Baldwin, 2008; 2012).
An effort to consolidate progress, and questions still unanswered. Clearly, ecosystem
research has not grown in isolation from the mainstream literature. Yet it is characterized by a
heavy emphasis on what is distinctive or novel about ecosystems. Only a handful of studies
have explicitly tried to bridge existing perspectives (e.g., network analysis; alliance research)
and ecosystems; even then, they have taken “ecosystems” as a given and examined them from
the perspective of a given theory. Venkatraman and Lee (2004) and Iyer et al. (2006), for
instance, have taken a “central” firm, in the network sense, to be the hub in an ecosystem.
While this yields an interesting map, there is less attention on how the extra complexity added
by the ecosystem terminology provides fresh insights. Likewise, Texeira et al. (2015) consider
network densities and collaboration in ecosystems, but don’t focus on what we learn from the
fact that ecosystems do have network structures. McIntyre and Srinivasan (2017) provide a
thorough and critical overview of platform and network-related ecosystems, articulating
questions to be considered by future research and linking with earlier work.

8
An important contribution here is made by Adner (2017), who proposes that “the ecosystem is
defined by the alignment structure of the multilateral set of partners that need to interact in
order for a focal value proposition to materialize” (2017: 42). “Alignment structure,” defined
as “the extent to which there is mutual agreement among the members regarding positions and
flows,” becomes the objective, pursued through a firm’s “ecosystem strategy” to “secure its
role in a competitive ecosystem” (2017: 47).
Taken together, prior research leaves open intriguing questions regarding the factors that make
ecosystems—as opposed to integrated supply chains or arm’s-length relationships—
“increasingly critical” (Adner 2017: 53); the factors that enable (or inhibit) alignment via an
ecosystem; and what ecosystem thinking can tell us that other literature streams cannot. The
remainder of this paper offers some proposals in this direction and identifies the key
contingencies that shape different types of ecosystems.
TOWARDS A THEORY OF BUSINESS ECOSYSTEMS
Theoretical primers: modularity and coordination in ecosystems. An important but neglected
characteristic of ecosystems is that they help coordinate interrelated organizations that have
significant autonomy. In nearly all the empirical cases we know of (from both literature and
experience), this is enabled by a modular architecture (Baldwin & Clark, 2000), where the
distinct parts of the ecosystem represent organizations that are separated by “thin crossing
points” (Baldwin, 2008), i.e. discrete parts of the production process. Technological
modularity allows interdependent components of a system to be produced by different
producers, with limited coordination required. While the overarching architecture design
parameters may be set by a hub, organizations have a large degree of autonomy in how they
design, price, and operate their respective modules, as long as they interconnect with others in

9
agreed and predefined ways.6 While new coordination issues always arise, ecosystems provide
processes and rules on how to resolve them, and encourage alignment through rules of
engagement, standards, and codified interfaces. The presence of modularity is also the
condition that allows a hub to forego at least some degree of explicit coordination. Thus,
modularity (though not necessarily open interoperability) creates the conditions for an
ecosystem to emerge.7
More modularization has been associated with a greater prevalence of ecosystems in a number
of sectors, from telecommunications to financial services to mobility. Many of the sectors that
have been studied in the context of ecosystems—IT, telecommunications, videogames, etc.—
tend to be more modular, suggesting that ecosystems may well be a distinct solution to the
problem of inter-firm coordination, distinct from the use of alliances, supply-chains or market-
based interactions. So, we posit that modularity allows for coordination of independent yet
interdependent firms through ecosystems. Yet while modularity may be necessary for
ecosystems to function, it is clearly not sufficient. As Baldwin (2008), Langlois (2003), and
Jacobides & Winter (2005) argue, modularization and the subsequent reduction of frictional
transaction costs is more likely to lead to the emergence of markets. For ecosystems to be
useful, there must also exist a significant need for coordination that cannot be dealt with in
markets, but which also does not require the fiat and authority structure of a central actor.
6 In most ecosystems, firms (e.g. app producers for iOS) are able to set the price and freely decide content of their
products, even if there is a set of clear specs. In others, such as Uber, drivers can set their desired availability, and
choose the quality they offer (which affects rider reviews and thus demand), and price can vary—albeit
uniformly for all drivers in an area. All these independent yet interdependent structures entail some modularity.
7 It is important here to distinguish between modularity as we define it and its more colloquial uses. Modularity is
here meant to denote the separability along a production (or production and consumption) chain. This does not
entail “plug and play” interoperability and free entry. Conditions of participation in each module or part of the
ecosystem may be exclusive. For instance, Apple may have a modular system, but part of it is open only to parts
of its own organization; parts of it are open to suppliers; parts of it are open (with stringent criteria) to
complementors; and parts of it are open with few restrictions. Modularity does not necessarily entail openness.

10
This, in turn, arises due to different types of complementarities.
Ecosystems, we posit, are distinctive both because of their structure, and because of the way
in which they allow the coordination challenge to be resolved. Figure 1 below illustrates how
ecosystems differ in terms of structure, when compared to either market-based transactions or
supplier-mediated arrangements (including those through a system integrator, or an integrated
firm). What sets them apart from an aggregate of buyer-supplier relations is that in ecosystems
final customers can choose among the components (or elements of offering) that are supplied
by each participant, and can also, in some cases, choose how they are combined. For
example, the end user in the Android phones ecosystem decides which apps they buy, and
from which provider, instead of buying a single, combined offering (“as is”) provided by a
single firm. What sets ecosystems apart from market-based arrangements is that end
customers choose from a set of producers or complementors who are bound together through
some interdependencies—by all adhering to certain standards, for instance. In this sense,
ecosystems differ from networks (Powell, 2003), and represent webs of standardized formal
or informal alliances between participants, where, e.g., complementors can choose from a set
menu of options and are treated similarly. In ecosystems, even customers themselves must
“affiliate” (Hagiu and Wright 2015) with one group or platform to be able to use its specific
complements like in the case of apps. But what distinguishes this group of “affiliated”
participants, and what is so different about ecosystem-mediated complementarities? To
understand this, we must first briefly consider the various types of complementarities in
economic relationships.

11
Types of complementarity: understanding what underpins ecosystems. As Teece (2017: 17)
notes, “the literature on complements is both confused and complex.”8 For our purposes, we
focus on two types of complementarities that can be expressed unambiguously in
mathematical terms, and characterize relationships between actors within ecosystems.
First, we have unique complementarities. The strict version is that “A doesn’t “function”
without B,” where A and B can be specific items—like the two ends of a pipeline (see Hart &
Moore, 1990)—steps, or activities. The more general version is that the value of A is
maximized with B (as opposed to B’). Such complementarity may also be a matter of degree,
with a continuum extending from strict or strong (where A requires B) to specific (where A
requires B to be customized to it to be productive) to generic.9 Unique complementarity can be
one-way: Activity or component A requires a particular (asset-specific) activity or component
B10, but not vice versa. Or it can be two-way, where A and B both require each other, which is
what underpins the idea of co-specialization (Teece, 1986).11
8 Teece suggests that Samuelson’s quote (1974: 1255) that “the time is ripe for a fresh, modern look at the
concept of complementarity … this ancient preoccupation of literary and mathematical economists. The simplest
things are often the most complicated to understand fully” is still relevant today. We concur.
9 Hart & Moore’s (1990) definition of unique (strict) complementarity implies that the two assets are
unproductive unless they are used together, which makes coordination of investments in the two assets critical to
maximize the marginal return on investment.
10 By A “needs” B, here we mean: the use of A and B together will help achieve the system’s overall purpose.
Note that most of the theory, especially transaction cost economics (TCE), considers that we can substitute
activities and components that are not specialized—using C rather than B—and the result will still be functional,
albeit less efficient. This trade-off between efficiency and the risks of specificity underpins TCE.
11 To put this in theoretical context, TCE focuses on unique complementarities that arise as a result of asset
specificity, which itself is a managerial choice, requiring the appropriate governance choices to protect against
attendant behavioral risks (Argyres & Zenger, 2012; Williamson, 1985). As such, unique complementarity that
arises because of asset specificity makes integration attractive. TCE focuses on unique complementarity in
production, though there can be unique complementarity in use, where a consumer needs to “assemble” different
components that only work together.

12
However, as Teece (1986) observed, complementarities may also be generic. That is, while a
particular good or service may be needed for the production of a complex value proposition or
innovation, that good or service may be generic (i.e., standardized) enough for firms to draw
on it with little concern for governance structure or risks of misappropriation. The use of
generic complements, discussed in detail by Helfat & Lieberman (2002), is an important and
common way to facilitate production while safeguarding against contractual hazards. The
same principle applies for an ecosystem analysis. To illustrate, electricity is needed for almost
everything, but the fact that it can be purchased in generic terms means that this
complementarity does not give rise to particular issues of economic organization12, and as
such, it can take place in markets instead (Adner, 2017). In other words, the generic nature of
this complement means that there is no need to coordinate in specific ways (i.e., no need to
create a specific alignment structure) between the economic actors. A teacup, boiling water,
and a tea bag may all be needed to make a cup of tea, but the complementarities are generic,
not specific. While consumers derive utility by combining these elements into a “product
system” (i.e., a cup of tea), producers do not need to coordinate their investments through
structures to enable such value. Consumers can thus buy them separately in the market and
combine them on their own. This is not to say that generic complements are not relevant for
economic actors; it is to say that generic complementarities have a different impact,
understood elsewhere (e.g., Rosenberg, 1969), and require no special new label.
The second category of complementarity we consider is supermodular or “Edgeworth”
complementarity. This concept has been developed by Milgrom and Roberts (1990), building
12 This highlights the key point that assets that are more fungible across applications along a production (and
consumption) chain are generic in nature (see Helfat & Lieberman, 2002 for an extended discussion.)

13
on Topkis (1978; 1998) and can be summarized as “more of A makes B more valuable,”
where A and B are two different products, assets, or activities.13 It can be found in both
production and in consumption. In production, it is manifested when coordinated investments
in both A and B yield higher returns than uncoordinated equivalents, or yield lower costs than
the sum of costs of independent investments into A and B (e.g., Arora & Gambardella, 1990;
Cassiman & Veugelers, 2006; Lee et al., 2010). Supermodularity in consumption, more
commonly associated with Edgeworth, is famously the basis of both direct and indirect
network effects (e.g., Farrell & Saloner, 1985; Parker & Van Alstyne, 2005)14, and can be
one-way or two-way. These different types of complementarities may also coexist. In the
example of an OS platform/app ecosystem, the app and the platform have a unique
complementarity in the sense that the app does not function without the OS (unique
complementarity, unidirectional, as the OS operates without most apps); and supermodular
complementarity, as the presence of apps increases the value of the OS, and (possibly) the
breadth of the OS installation increases the value of the app.
While there are additional ways to organize our understanding of complementarities,15 the
13 There is a long history of this type of complementarity, starting with F.Y. Edgeworth, who considered it in the
context of consumption of goods—looking at how changes in demand of one affects the demand of another (see
Samuelson, 1974, for a more recent analytical treatment and Weber, 2005, for a historical account). Milgrom and
Roberts shifted the analysis, and popularized Topkis, using lattice theory, and employing complementarities less
on consumption, than on production—with their quintessential example being the Japanese production system,
where the value of one practice depends on the existence of another, per the analytics of Topkis (1978). This is
why we use the formal term “supercomplementarity.” To provide the analytical definition, Milgrom and Roberts
(1994: 6) note that “a group of activities are (Edgeworth) complements if doing more of any subset of them
increases the returns to doing more of any subset of the remaining activities.”
14 Direct network effects emerge when consumers value the fact that other consumers use a particular product or
service- like in the case of users of a fax machine. Indirect network effects occur when the existence of a variety
of complements creates value to other complements- suggesting that some final customer values variety (not
volume).
15 Teece (2017), for instance, considers a different set of complementarities, which include some we don’t
consider, such as Hicksian complements, “when a decrease in the price of one factor leads to an increase in the
quantity used of its complements in production”, or Hirshleifer (Asset Price) Complementarity to denote how
innovation in one segment affects asset prices in another. He also considers technological complementarities,

14
discussion of these two types, and of their different directions, gives us our foundation here.
These complementarities can apply to both production and consumption.
As we define them, then, ecosystems are groups of firms that must deal with either unique or
supermodular complementarities that are non-generic, requiring the creation of a specific
structure of relationships and alignment to create value. The strength of ecosystems, and their
distinctive feature, is that they provide a structure within which complementarities (of all
types) in production and/or consumption can be contained and coordinated without the need
for vertical integration.16 From this perspective, ecosystems allow for some degree of
coordination without requiring hierarchical governance, precisely because of the ability to use
some standards or base requirements that allow complementors to make their own decisions
(in terms of design, prices, etc.), while still allowing for a complex interdependent product or
service to be produced.
Ecosystems as the result of a (partly designed) process. Ecosystems, of course, do not just
“emerge” spontaneously. They are at least in part the result of deliberate experimentation and
engineering from different parties. For instance, a firm may choose to modularize a process, or
may opt not to procure generic complements available on the market not only because non-
fungibility may give it additional design options, but because it wants to set up an ecosystem
to create or extract more value. Overall, powerful firms (especially hubs, or hub contenders)
craft rules and shape the process of ecosystem development to tie in complements and make
which occur when “the full benefit (or even any benefit) of the innovation cannot be achieved until some other,
complementary technology (which, on its own, has only lower value uses) has been created or re-engineered”—a
condition that we consider to be very frequent in the settings covered by ecosystems, and as such encompassed
by either of our proposed categories.
16 Adner and Kapoor (2010) intriguingly find that the role of upstream complements differs from the role of
downstream complements, when we consider their impact on ecosystem health. This arises, we would argue,
from the fundamentally different role of unique complementarity (dominant in production) from
Edgeworth/supermodular complementarity (dominant in consumption, i.e. downstream).

15
complementors abide to them. To give a specific example, even within the Google/Android
ecosystem, with Google as the hub and clear rules for complementors, some key handset
manufacturers such as Samsung and Motorola are starting to create sub-ecosystems. They
allow key app developers to connect via APIs in ways that are specific to their device, so as to
“lock them in” with non-fungible investments.
Ecosystem design is becoming ever more important as the question of what drives customer
value, and how firms can capture this value by monetizing it in some way, becomes an open
question in a technologically mediated world, where regulation provides some loose contours
of how sectors can work and what actors can legitimately sell (Parker, Van Alstyne, and Jiang,
2016). Firms identify what drives value to users (B2C), but do not always charge the users for
it; often they charge other clients (B2B), who are willing to fund a venture to acquire its client
information or access, or to show that they are affiliated with value-adding services (B2B2C).
This requires the formation of ecosystems, where rules and roles, and monetization, as well as
how players are connected, become an essential part of the business model design.17
Finally, while modularity, the nature of complementarities, and fungibility may all be partly
designed, the process is not always driven by foresight. Some firms—especially those who
allow modular technologies—may unwittingly form ecosystems. Arguably, this happened to
inkjet printer makers, who faced entry by unauthorized ink-cartridge producers. More famous,
17 Consider Traipse, an app providing a geo-located “treasure hunt” experience via smartphones, which serves
the dedicated group of puzzle-solvers who also like to engage with and explore local communities. Its B2C
technology allows individuals to challenge themselves in tours of historic districts as they solve riddles. Most of
its revenues come from B2B Chambers of Commerce or tourist bureaus that want to promote their local shops
and sights to this dynamic demographic, that fund the cost of developing tours; local businesses also fund the
venture by offering “prizes” as discounts in their stores, potentially giving a commission to Traipse for each sale.
Traipse also uses a cryptocurrency which is used as an exchange means and creates local stickiness, based on
smartoken.com—yet another venture which is B2B and connects with businesses giving it commissions for
creating its blockchain technology to power localized ways of exchanging funds. So user and client are distinct in
such models.

16
when hackers developed the first apps for Apple’s iOS, they wound up in court—until Steve
Jobs realized he could turn an unwitting ecosystem into a regulated one, and profit from it.
Characteristics of ecosystems. To understand how ecosystems operate, we first need to fully
define them. We do so by drawing on pragmatism (see James, 1975; Dewey et al., 1999).18
Our approach differs from existing ones by focusing on the nature of complementarities
between ecosystem participants, and the fungibility of their investments, as opposed to the
resulting collaborative (or alignment) structures. It is motivated by the desire to focus on the
mechanisms that result from complementarity type, and also by our desire to avoid issues that,
while potentially important, have been addressed by prior work.19 Concretely, we suggest that:
An ecosystem is a set of actors with varying degrees of multi-lateral, non-generic
complementarities that are not fully hierarchically controlled.
This encapsulates three crucial attributes of an ecosystem. First, “multi-lateral, non-generic
complementarities” are either unique complementarities (which essentially lead to some
degree of co-specialization), or supermodular/Edgeworth complementarities (often found in
complements-in-use). Our narrow definition delimits the scope of the ecosystem to specific
complements. So, while boiling water may complement tea bags and teacups, they are generic
complements in consumption and thus are not parts of an “ecosystem” by our definition. We
exclude generic complementarities because they do not give the parties any vested interest to
align and act as a group. While any focal actor would do well to consider all its
18 That is, we do not start from the realist premise that there is “some” truth about ecosystems in the world, which
we try to approximate, but rather view constructs as vehicles to understand the world in a pragmatic sense,
judged by their usefulness in helping us do so.
19 We do appreciate the potential merit and utility of definitions that also include generic complementarities.
Indeed, an inclusive approach may be the most appropriate way to go, if our intent is to warn managers against
underestimating the role of external alignment when delivering an interdependent value proposition or launching
an innovation, for instance (Adner, 2012, 2017).

17
complementarities, we do not think that generic complementarities can usefully capture what
is unique about an ecosystem. This uniqueness, we argue, lies in the non-generic nature of
complementarities, which also entails some degree of customization. It is precisely this
attribute that underpins the particularities of ecosystems.
We also posit that the (“multi-lateral”) complementarities exist at the level of the sets of roles
(Adner, 2017) that link the different parties together—e.g., hub(s), suppliers, or different types
of complementors. What makes ecosystems unique is that the interdependencies tend to be
standardized within each role, which creates the need for a new set of skills in terms of
designing ecosystems (Helfat and Raubitschek 2018).20 Thus, some of the relationships
between sets of actors will be unique, some supermodular, some generic, and others specific.
But, regardless, the relations can be described at the level of the roles or groups of actors as
opposed to the dyad, which is a fundamental shift from the usual mode of analysis—e.g. in
TCE (Williamson, 1985). While the arrangements entered in by ecosystem members might be
seen as webs of alliances, these are standardized and set for each role in an ecosystem.
Our analysis also makes headway in extending the useful notion of co-specialization (Teece,
1986, 2017) to explore the nature of mutual dependencies. These depend on the relative
fungibility of investments to operate in ecosystems, and relationships within them. They
define the cost to “re-tool” and “re-customize,” pitted against the benefit that ecosystems
bring. For unique dependencies, the benefit is the creation of a dedicated set of partners who
can fulfill the requirements needed and supply, or buy, what is offered. Thus, members of
20 For instance, all apps in the Android or iOS ecosystems are treated identically, and the interdependencies are
not between Apple and its millions of app developers individually, but across the group as a whole. That said, the
nature and definition of each role, as well as the conditions they are faced with is a matter of strategic (research)
design. That creates the need for a new set of skills in terms of designing ecosystems, which may parallel what
has been called for by Helfat and Raubitschek (2018).

18
ecosystems, rather than being stuck in individual sets of relationships, each fraught with their
own risks, can benefit from a greater set of options. For supermodular complementarities, the
benefit also comes from the value for actor X from the additional availability of input or
complement Y.21
This analysis presents a different canon to the one used in TCE, which is based on the issues
of risk mitigation in a dyadic relationship. The focus here is on how to maximize the benefits
by engaging (or being part of) a group of firms with complementary roles, or how to design
the best ecosystem structure (which will vary depending whose perspective we take). Unlike
in supply relationships explored by TCE, in ecosystems, neither prices nor qualities are fixed;
they are left to vary, and to be chosen (by design!) as a function of the choice of a final user,
and often the objective is to coalesce with other firms in securing more final users and
customers for the group.
Because of the complementarities, connecting to an ecosystem involves some investment that
is not fully fungible—that is, the investment, or assets in place, cannot be easily redeployed
elsewhere without cost. This cost may derive from product/offering configuration adjustments
(e.g., Kapoor and Agarwal, 2017) that require new investments, adjustments to the
membership and transaction rules of other ecosystems (e.g., Claussen et al. 2013), or
coordination costs with other members’ activities. This is, in our view, a fundamental
structural feature that makes within- and across-ecosystem interactions strategically distinct.
The degree to which a participant’s effort is tied to one ecosystem, and cannot be recoupled in
any other setting, determines the economic basis of their attachment to that ecosystem (see
21 This, however, is not symmetrical between two parties; it may be that the marginal value of X increases in Y
but this doesn’t mean that the marginal value of Y increases in X. Also, the benefits of supermodularity are not
exogenously driven: They will vary, and may decline, e.g., as the ecosystem grows and the benefit of more X
goes down due to saturation.

19
e.g., Cennamo, Ozalp and Kretschmer 2018).
More broadly, we argue that the nature and direction of the dependencies; the extent of the
underlying complementarity; and the question of whether they are unique, supermodular, or
both, as well as the fungibility of investments to participate all become important descriptors
of an ecosystem that can help us understand when and why alignment occurs—or fails. By
looking at the nature of complementarities, and describing whether they happen for
consumption or production, we arrive at different types of ecosystems. Figure 2 illustrates and
provides real-world examples of different types of ecosystems—including producer- and
platform-based ecosystems and multi-sided platforms. 22
Insert Figures 1 and 2 about here
Finally, our definition suggests that ecosystems are not unilaterally hierarchically controlled.
For all the power a hub (if there is one) may wield, ecosystems, as we define them here, lack
the hierarchical controls of traditional firm groupings, quasi-captive systems such as Keiretsus
or Chaebols, or supply networks. What we think is analytically distinct in ecosystems is that
their members all retain residual control and claims over their assets: no one party can
unilaterally set the terms for, e.g., prices and quantities, in addition to standards. That is, we
posit that ecosystems need to be both de jure and de facto run with decision-making processes
22 Multi-sided platforms (“MSPs”) (e.g., Hagiu, 2006; Hagiu and Wright, 2015; Parker and Van Alstyne, 2005),
i.e. marketplaces such as Amazon Marketplace (ecommerce marketplace), Match.com (online dating
marketplace), or Just Eat (takeaway food marketplace), may create their own ecosystems. In our view, a number
of MSPs are not necessarily, in and of themselves, ecosystems, inasmuch as they do not require any non-fungible
investment and require only generic supermodular complementarity. (Some, of course, require affiliation that
does lead to a type of non-fungible relational investment). Platform hubs, to be able to strengthen their position,
may choose to require some complementors to invest in non-fungible ways. For instance, the way a product is
promoted, sold and shipped to the final customer through Amazon Marketplace becomes unique to the Amazon
marketplace and different than in other two-sided markets to the extent that providers specialize into the specific
interface requirements of Amazon (e.g., Amazon’s product stocking and shipping requirements that align with
“Amazon Prime” service). Thus Amazon has an MSP that sustains an ecosystem with Amazon at its core.

20
that are to some extent distributed, and without all decisions (especially on both prices and
quantities) being hierarchically set—even though standards, rules, and interfaces are often set
by a “hub.”23 This allows us to distinguish between ecosystems and supply chains, since in
supply chains the hub (OEM, or buying firm) has hierarchical control—not by owning its
suppliers, but by fully determining what is supplied and at what cost.24
THEORY IMPLICATIONS: A RESEARCH AGENDA ON ECOSYSTEM DYNAMICS
Our approach complements existing research by moving beyond the description of how
players align to consider why and when they align; it also offers a set of predictions of when
ecosystems, as opposed to vertically integrated firms or supply networks, will dominate. Our
focus on different types of complementarity, as they interact with modularity to drive
ecosystem dynamics, can deepen and extend existing and emerging work on coordination,
collaboration, and value creation/capture.
Ecosystem coordination. We posited that modularity is a critical facilitator of ecosystem
emergence. This leads to a straightforward empirical prediction, which can be tested by
longitudinal, within-industry or cross-industry research linking changes in modularization
with the emergence and growth of ecosystems. Yet while modularity helps generate
ecosystems, it is not an exogenous factor. It results from the agency of key industry
23 There is significant variance in terms of how open and democratic rule-setting is, especially with regards to
standards and interfaces used. These vary from ecosystems like Apple’s, with interfaces that are strictly and fully
controlled, to Linux, with the involvement of various ecosystem participants. We return to this later.
24 Thus Toyota, which is at the center of a group of co-dependent suppliers that occupy different parts of the
value chain and co-specialize with it (Nishiguchi, 1994), unilaterally decides what it will procure, from whom,
and at what cost. Toyota is not, by our definition, the keystone of an ecosystem. Apple, on the other hand, with
its App Store, is a keystone. It manages participation criteria, standards, and rules, which define to a great extent
the type of members that participate in the ecosystem, and how they interact, but does not decide what
specifically they contribute to the ecosystem (which app they should produce), thus how many apps will be
published or downloaded; nor does it set prices, beyond setting an acceptable range.

21
participants—whether they are far-sighted or not (see Jacobides et al, 2016). Whereas firms
that want to encourage ecosystems are likely to push for modular structures with clear
interfaces, an ecosystem may coalesce even without the focal firm’s desire to open up, as
happened with IBM’s system 360, or Apple’s early iPhone ecosystem of unauthorized apps.
The understanding of “accidental” (or even illicit) ecosystems and the way they evolve is a
fascinating area for future research.25
Ecosystem collaboration. Why do some forms of inter-organizational collaborations happen
in ecosystems rather than in other forms, such as supply chains or alliances? And what kinds
of collaboration and coordination behaviors are we likely to observe within ecosystems? Our
framework suggests that depending on the type of complementarity, we will get a different set
of behaviors—and, likely, organizing structures too. Dynamics in nascent sectors, which have
received increasing attention of late (e.g., Gurses & Ozcan, 2014; Hannah, 2014), may
contrast with mature settings. As we move to increasingly dynamic settings, understanding
which attitudes and approaches enable the identification and then success of new ecosystems,
and which might lead to their demise (West and Wood 2013), will be an intriguing area of
new research.26
Likewise, there is benefit to a comparative analysis of different approaches that firms take to
similar problems, with some engaging with or even creating different types of ecosystems,
others choosing to rely on markets, and yet others becoming system integrators and vertically
25 It is also worth noting that for ecosystems to operate, we need some standards and rules; and often ecosystems
are most needed in emerging areas where coordination problems are rife, and these rules are sorely lacking.
26 Study of “live experiments” such as IDEO’s Co-Lab, wherein firms from different sectors come together to
structure new propositions, where rules and roles are “designed” in real time can yield promising new directions.

22
integrated providers. Is there an inherent advantage to each of these solutions? 27 To what
extent are there firm-specific skills in leveraging capabilities, or knowing how to run
ecosystems, as Helfat and Raubitschek (2018) argue—and which ones matter most?
Our analysis of complementarities also offers some guidance on managing ecosystems. Under
unique complementarities, we expect participants to care about ecosystem health only
inasmuch as its demise would eliminate the demand, whereas supermodularity would also
increase the very attractiveness of the product or service offered by ecosystem participants.
This should increase collaboration propensity.28 Furthermore, requisite investment fungibility
helps shape the appropriate strategies for managing ecosystems. The greater the
supermodularity and the lower the fungibility, the easier it will be to align effort for current
participant members. This might be because the less fungible the effort required to participate
in an ecosystem, the keener a participant will be to see the common enterprise succeed, as the
cost of redeployment increases. However, the lower the fungibility the harder it will be to
recruit ecosystem participants, who may fear being locked in. This implies what is likely to be
a recurrent strategic conundrum to the question of ecosystem design. A better understanding
of tactics and governance mechanisms that hub firms use to recruit, motivate, and retain
participants will be helpful.
Finally, our framework suggests that we need to consider ecosystems in their competitive
context. Fungibility of assets and relationships, for instance, is a function of how easy it is to
27 Consider, for instance, the challenge of electric vehicles. Different firms put together a variety of approaches,
both in terms of how integrated they are, and, if they use ecosystems, what the nature of these ecosystems is (see
Chen et al., 2017). Some, like Tesla, are vertically integrated and concurrently use supply networks. Others, like
Nissan, use the open market. Others, like Wanxiang, use ecosystems—with distinct and incompatible ways of
charging, and protected by different standards, but where participants have significant autonomy (Weiller et al,
2015).
28 This also offers a cautionary note on our interpretation of the increasing number of ecosystem studies, as the
findings of both mechanisms and outcomes will heavily depend on the nature of interdependencies. This affects
finding generalizability, especially for platform-based studies (see McIntyre and Srinivasan, 2017).

23
redeploy them; and the attractiveness of conditions in one ecosystem is a comparative
assessment of what the alternatives are. This requires a shift in empirical focus from within-
ecosystem to across-ecosystem dynamics, as they are likely to influence each other.
Ecosystem value creation/capture. Assessing how the different types of complementarity
play out can also highlight some of the underlying mechanisms of value creation and capture
in and across ecosystems. Consider, for instance, the role of directionality of co-
specialization. Whether fungibility is one-way or bilateral, and whether it is symmetric or
stronger in one direction or another, will affect both the behavior of the actors of an
ecosystem—in terms of their preference to cooperate vs. expropriate—and the recruitment of
new members. The very things that make it easy to capture value within an ecosystem make it
harder to recruit (and, less so, retain) members. This becomes even more important when
ecosystems compete for members, so that members may decide to shift to another ecosystem
if the conditions no longer favor them. A further interaction is that the more an ecosystem is
driven by supermodular complementarities, the more hubs will initially try to focus on
attracting members; yet as an ecosystem becomes dominant, recruitment takes care of itself,
so that value distribution may become more lopsided.
Our framework could also help explain competitive dynamics. How do the rules and expected
fungibility of participating in one ecosystem change as a result of actions in another? That is,
how does the growth of ecosystems such as Android affect the rules, required commitments,
and standards of competing ecosystems such as iOS? And how do the hub firms and
participants respond?29 Also, what role is played by regulators or social pressure groups? We
29 We will find it hard to understand one ecosystem and its rules without direct reference to the other: to
understand Hailo, we need to understand Uber, and local taxi dispatch structures too. Strategic dynamics become
even more complicated when firms participate in rival ecosystems, such as Microsoft developing its MS Office
Suite for Apple’s Mac OS, while simultaneously trying to advocate its own software ecosystem against that of
Apple.

24
can readily understand that ecosystems characterized by strong supermodular
complementarities will want to become established, and then profit on the basis of the network
externalities they generate—but we should also expect that regulators (and prospective
ecosystem members) will want to see the exact opposite, pushing for interchangeable
standards and generic complementarities that allow free entry and exit.30
We would expect ecosystems with supermodular complementarities to be more resilient than
those resting purely on unique complementarities, but even these can be overturned through
competition. This raises interesting questions that may explain, for instance, how Symbian,
with 67% of the smartphone OS market (and against the predictions of most economic or
strategy models), lost ground to the young upstart Android. As Pon et al. (2014) note, much of
this is due to differences in organizational efficiency, governance, and nature of co-
specialization.
Ecosystem governance and regulation. To understand such strategic dynamics, we need a
clearer sense of how ecosystems are structured and governed. Behavior in an ecosystem, and
ultimately its success, is affected by the rules of engagement and the nature of standards and
interfaces—open vs. closed; imposed vs. emergent. Some ecosystems have clear, possibly de
jure defined standards, especially if they have many members. Others, especially those not
based on technology, might have de facto expectations in terms of the rules of engagement.
Relatedly, we need to compare and contrast open and closed ecosystems, with several shades
of gray in between. Some ecosystems accept any participant who agrees to a minimal set of
30 This may also have significant welfare implications, with final customers or ecosystem members benefiting in
the short term, even though it might discourage ecosystem formation from hubs anxious about expropriation in
the longer term. A new set of policy and regulatory questions could thus come into play—as we have seen with
recent discussions on the power of Apple, Google, Facebook, and Amazon, or electric cars.

25
rules, whereas elsewhere membership is strictly controlled, whether by committee or by the
hub—if there is one.31 Rules pertaining to hierarchy or membership may change over time, as
with Facebook (Claussen et al., 2013). We need to understand how membership rules vary,
what drives this variation (and its competitive impact), and how this relates to standards (open
vs. closed; proprietary or sector-wide), modularity, and the nature of complementarities.
Our framework provides a starting point for understanding the underlying forces operating in
an ecosystem, but several questions remain. What determines the level and form of control in
an ecosystem? Which control mechanisms can a hub use, and when does control become so
unilateral that the ecosystem becomes a supply chain? Do the mechanisms governing the
ecosystem change as a function of the shifting nature of modularity, of complementarities, or
other factors? Where, in particular, do we see the emergence and success of distributed
governance in ecosystems, such as in the open-source movement (O’Mahony & Bechky,
2008)? Looking past the shiny success stories of strong hubs such as Apple, we should also
ask what we can learn from firms that tried to become hubs but failed. We should also
remember that most ecosystem members are complementors (for instance, in July 2014 there
were 2.3 million individuals working as app developers), with very limited power (Pon,
2016). While research has started to consider their plight (Ceccagnoli et al., 2012; Kapoor
2013; Selander et al., 2013), it has mostly examined firms facing tactical decisions such as
multi-homing (Bresnahan et al., 2014; Mantena et al., 2010). However, it remains to be
understood how these firms achieve complementarities at the ecosystem level, and how
participating in multiple ecosystems influences the type and intensity of such
31 Consider different videogame consoles’ ecosystems. Historically, Nintendo has set strict rules for participation,
imposing exclusivity clauses and limiting the number of complements members can develop for its systems.
Rival ecosystems, such as those sponsored by Sony or Microsoft, have adopted rather more laissez-faire policies.

26
complementarities, and thus the benefits for the participants of an ecosystem and its sponsor
(e.g., Cennamo et al. 2018).
Back to the strategy literature. Our Appendix provides some thoughts on how ecosystem
research can benefit, and contribute to, mainstream strategy research, complementing the
discussion of Adner (2017). Beyond these general links, we think that the answers to the
questions posed in this section can push the research agenda to revisit some of the
fundamental concerns of strategy research. The ecosystem construct reaffirms the importance
of considering the aggregate level of analysis in assessing firms’ competitive advantage: If
firms gain from others participating in an ecosystem, but cannot fully control them, what does
that imply for how they attain advantage? Frameworks such as the RBV mostly concern
themselves with owned resources. How should this perspective change when the resources
exist not at the level of the firm, but at the level of the ecosystem? And, linking the RBV,
dynamic capabilities, and ecosystems, what sort of resources and capabilities could be
valuable for firms in this dynamic context (see Helfat and Campo-Rembado, 2015)? Helfat
and Raubitschek (2018) have recently argued that innovation capabilities, environmental
scanning and sensing capabilities, and, in particular, integrative capabilities are critical for
ecosystem orchestration and platform leadership. The question remains, how does the value of
resources and capabilities differ depending on the role firms take within the ecosystem (hub
vs. participants)?
Our objective in this paper was to advance our understanding of ecosystems, and to propose
elements of a positive theory of ecosystems—the role of modularity, and the impact of
different types of complementarity (and the resulting fungibility) as they tie ecosystem
members together in a web of interdependent yet autonomous activities. We hope that the
directions offered will enrich both research on ecosystems, and research in mainstream
strategy, as firms become increasingly engaged in, and respond to ecosystem growth.

27
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Figure 1. Different Types of Value Systems

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Figure 2. Types of Complementarities and Ecosystems

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APPENDIX: ECOSYSTEMS’ RELATION TO EXTANT RESEARCH
The real litmus test of a theory must be its ability to add value: to extend existing research, not
duplicate it. In this section, we ask, “What does the ecosystem lens show us that we could not
otherwise see?”
Starting with industry analysis, the focus on ecosystems appears to be a clear complement.
Traditional analysis of sectors and their evolution lacks the vocabulary to consider groupings
such as ecosystems or examine their dynamics. On the rare occasions that ecosystems relate to
a single sector, their function is likely to affect the success of that sector, and the patterns of
value distribution within it, in idiosyncratic ways that are worthy of further study. In looking
at these patterns, ecosystem research would benefit if it employed specific methodologies that
have been developed to study joint value creation and distribution—as opposed to merely
alluding to them. So, there is gain to be had here—phenomenologically and theoretically.
Moving to recent work on Industry Architecture (Jacobides et al., 2016), there are clear and
strong connections. Ecosystems appear to be one of many ways that a sector or set of sectors
can be structured; that is, they seem to represent a specific type of industry architecture. The
nature of this architecture will affect the potential patterns of value creation and distribution
both between different, potentially competing ecosystems, and between the participants in
each ecosystem. There is, already, research that considers how ecosystems develop in the
context of particular IAs (Tee & Gawer, 2009) and how ecosystem structure affects value
distribution (Dedrick et al., 2010). The concept of bottlenecks (Baldwin, 2014; Jacobides &
Tae, 2015), central to IA, is clearly relevant to ecosystems too (Hannah & Eisenhardt, 2016).
However, IA does not yet have a developed repertoire that examines how these specific,
particular types of groupings (i.e. ecosystems) come about or are governed, or how they shape
value distribution.

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Research on modularity (Baldwin, 2014) could be usefully employed to help us understand
ecosystems, and vice-versa. As we posited, we see modularity as a precondition for the
emergence of ecosystems. But, is it the case that modular structures emerge as a result of
conscious design, e.g. by hub firms? If so, what determines what gets “opened up” and what
does not? When do we see modular systems emerge, and what is the role of existing
ecosystems in either enhancing or interfering with modularity? What are the results of
different strategic choices of hubs and of other ecosystem participants in this regard?32
The more established literature on alliances also stands to benefit from the construct of
ecosystems. Phenomenologically, ecosystems overlap only partly with alliances (Gulati et al.,
2012), and clearly represent a very specific subset of them. Sometimes, ecosystem
relationships do not require a formal alliance, and do not bind firms, since participation in an
ecosystem might merely be a function of adhering to certain specifications. However, we
could consider ecosystem participation as a particular type of loose alliance, where the link
between firms expresses the co-dependence brought about by their mutual co-specialization
(e.g., Alexy et al., 2013; Kapoor & Lee, 2013). As we stressed in the text, a defining feature
of ecosystems is the provision of standardized rules in terms of the alliances offered, which
are specific to the roles that are acknowledge in an ecosystem. These types of standardized
alliances, which need to imply some non-generic, non-fungible investment to qualify as being
the base for an ecosystem could be the basis of future research.
32 Note that the ecosystem analysis helps provide a strategic angle to the analysis of industry evolution. Consider,
for instance, Langlois (1992), whose thesis is that transaction costs are a transient phenomenon, and that as
contracting parties figure out a way to coordinate, hierarchy will give way to the market. Our analysis here
suggests that coordination (and modularity) is designed, and while there may be technical abilities of shifting to a
market through the use of standards, the strategic dynamics may not let that happen. Apple could surely create
open standards and migrate away from a closed ecosystem to an open market-based structure, but this would not
serve its strategic interests, and as such it will not do it. The exploration of these dynamics and of the strategic
use of ecosystem as a means of organizing, and potentially dominating, through a careful design of
complementarities, modularity, and of the governance of ecosystems remains a fascinating area for future
research.

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The nature of standardized and more bespoke alliances and (more rarely) joint ventures could
become a promising area for the understanding of ecosystem governance and management.
How do firms that co-sponsor ecosystems manage them? When do they use alliances to do so?
Also, alliances can be used not only as tools for ecosystem participants, but at a higher level,
to set the ground for building an ecosystem. How do these alliances differ from those that
simply pertain to ecosystem participation? For instance, how did the alliance of Microsoft and
Intel, by shaping Wintel and the nature of this platform ecosystem, affect both the outcome
and the nature of the looser alliances between Wintel and PC manufacturers? Furthermore, the
analysis of ecosystems and their associated alliances could provide fresh research questions
for alliance research, since the focus would be neither on the individual alliance nor on the
portfolio of alliances of a single firm (Wassmer, 2010), but rather on the alliances pertaining
to an ecosystem (Hannah & Eisenhardt, 2016).33
Research on networks (Powerll, 2003) could also benefit ecosystem analysis, and vice versa.
For all the solid work on networks, they have largely focused on the dynamics of one industry
(e.g. Uzzi, 1997). An ecosystem (which, by definition, encompasses firms with non-generic
group-level complementarities) could be mapped as a network, yet it is distinct, both because
it can have a cross-sector nature, and because of the existence of a set of distinct and
asymmetrical links tied at the group level by specific complementarity. There is significant
value in documenting the network structure of ecosystems (Iyer et al.; 2006; Venkatraman &
Lee, 2004), but there is greater promise in studying how network analysis metrics (such as
centrality, closeness etc.) apply to such segment-spanning networks, theoretically and
empirically.
33 To be fair to ecosystem research, the alliance literature does focus almost exclusively on dyads (see Dyer &
Singh, 1998); even the triad as a focus of analysis is a novelty (Davis, 2016). As such, analysis of a structured set
of alliances based on ecosystems goes beyond the “alliance portfolio” research (see Wassmer, 2010).

37
To reiterate, ecosystems are defined by non-generic complementarities at the group level,
which means that while there is competition to attract profits within the ecosystem, there is
alignment in how all members benefit from the success of the collective enterprise (i.e., the
ecosystem) (Adner, 2017) and thus gain advantage over another collective enterprise (i.e.,
another ecosystem) or a set of unrelated firms. For instance, firms participating in the Android
ecosystem clearly have issues on how to divide the total spoils (between and within different
parts of the value-adding process), but they also share a desire to beat the iPhone.34 It is this
particular incentive and organizational structure, distinct from ecosystems as we define them,
that yields the mix of cooperation and competition (sometimes efficient, sometimes less so)
that the literature has picked up. Thus, we have a different canonical problem from the one
institutional economics focuses on—one that provides opportunities to leverage and extend
existing research, much as the canonical buyer-supplier analysis led to the boom in TCE
(Williamson, 1985).
Some recent research is pointing in this direction. West and Wood (2013) document how the
structure of relationships and governance mechanisms between key actors in the Symbian
ecosystem created conflicting incentives that constrained the ecosystem’s capacity to evolve
34 Note that the extent of such interest is a direct function of the extent of complementarity that developers have.
At the margin, if an app can work seamlessly in both ecosystems, then there is little that binds these
organizations together, since the success of one ecosystem is irrelevant to their prospects—they can inhabit
another at zero cost. A further subtlety here is that if we take one organization that belongs to different
ecosystems (such as multi-homing app developers, who work with all major ecosystems), at the corporate level
there may be little vesting to particular ecosystems, even though at the business level, each unit tied to an
ecosystem does have co-dependencies. This observation opens up another set of interesting questions. First, for
firms that can participate in multiple competing ecosystems, what is the right strategy? Bresnahan et al. (2014),
for instance, show that the two platforms in our example target different segments of consumers, thus capturing
different parts of the market, which not only explains why they can coexist, but also why app developers find it
beneficial to participate in both, without having a vested interest in one ecosystem withering. Second, some
complementors may act strategically, and decide to multi-home to prevent a single platform ecosystem winning
(eg., Cennamo & Santaló, 2013). In the videogame industry, developers such as Electronic Arts have used this
multi-homing strategy to increase their value capture ability vis-à-vis any single platform. As such, over and
above the strategies of “the focal firm,” we can develop rich typologies and prescriptions for strategies of firms
that consider their participation in multiple ecosystems.

38
and create a thriving market for applications such as those later created by Apple and then
Google. With a clearer understanding of the underlying dynamics, we can progress further.35
Ansari et al. (2015) document how TiVo, a start-up firm that pioneered the digital video
recorder in the U.S. television industry, elicited greater acceptance and support for its
disruptive technology (even from incumbents) by influencing positioning and relationships
among members of the evolving TV industry ecosystem. Kapoor and Furr (2015) show how,
in the solar photovoltaic industry, complementarities at both the firm and ecosystem level can
explain the diversifying technology choices of firms participating in an ecosystem, while
Hannah and Eisenhardt (2016) look at how different firms within one ecosystem followed
diverse strategies in the attempt to strengthen their positions and become the bottleneck.36
35 There is an increase in studies that consider the ways in which organizations shape their ecosystems—and the
implications of these choices. Gatignon and Capron (2017) consider Natura, a Brazilian eco-friendly cosmetics
firm, and show how it built an ecosystem, linking with underprivileged women across Brazil and tribes in the
Amazon and other biomes to ensure the distribution and supply of its products. They point out that Natura chose
not to become the sole hub, but rather to create a broader ecosystem involving entire communities as well as
public, private, and non-profit partners collaborating together, given that the benefits from this multilateral
approach outweighed its expected benefit from being the sole hub.
36 Note that, per the narrow definition of ecosystems we advocate, while bottlenecks are integral parts of
ecosystems, bottlenecks can also emerge outside ecosystems. In a related set of sectors, where there is a unique
but generic set of complementarities (e.g., the requirement of rare earths for the production of mobile telephony
devices) the relatively more scarce component even in a sector with no ecosystem-like arrangements may lead to
a bottleneck in the spirit of Jacobides and Tae (2015), Baldwin (2014), or Ethiraj and Posen (2013). If there are
some non-generic complementarities, however, then we will have both ecosystem dynamics and bottlenecks—
and the design of the ecosystem may lead to bottlenecks.