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Venture Capital Redefined The Economic Political and Social Impact of Covid On The VC Ecosystem 1St Edition Darek Klonowski All Chapter
Venture Capital Redefined The Economic Political and Social Impact of Covid On The VC Ecosystem 1St Edition Darek Klonowski All Chapter
Venture Capital Redefined The Economic Political and Social Impact of Covid On The VC Ecosystem 1St Edition Darek Klonowski All Chapter
Venture Capital
Redefined
The Economic, Political,
and Social Impact of
COVID on the VC
Ecosystem
Venture Capital Redefined
Darek Klonowski
Venture Capital
Redefined
The Economic, Political, and Social Impact
of COVID on the VC Ecosystem
Darek Klonowski
Brandon University
Brandon, MB, Canada
© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer
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Thanks be to God
Preface
vii
viii PREFACE
Of this book’s many objectives, the primary focus has been its use as a
tool to understand the severity of the novel coronavirus’s impact upon the
venture capital industry. Analysis performed for this book project suggests
that the industry has undergone profound changes since the early onset
of government restrictions designed to halt the spread of COVID-19. A
second area of emphasis within this book is an assessment of the potential
long-term impact of the economic, political, and social restrictions on the
venture capital ecosystem post COVID-response. Thirdly, the perspec-
tives of various stakeholders involved in the venture capital ecosystem,
including general partners (GPs), limited partners (LPs), entrepreneurs,
and other important stakeholders (including the state) are considered,
particularly in the context of the post COVID-response period. Lastly,
the ultimate aim of this book is to answer the question of whether current
changes to the venture capital industry are likely to renew and promote
the industry’s overhaul, or simply perpetuate its decline.
in the industry are likely to emerge, although these trends have beset the
industry for some time and were visible prior to COVID-response.
The third chapter also outlines how COVID-response has redefined
personal relations in the venture capital ecosystem. Since venture capital
is a people-centric business, restrictions have affected virtually every
aspect of GPs’ interactions with entrepreneurs and LPs, in addition to
their in-house operations and relationships with other key stakeholders.
Considering recent disturbances to the already precarious venture capital
industry, the second part of the chapter extrapolates what the future
of venture capital may look like, and describes in detail the stages the
industry may evolve through; these include the global consolidation of
GP partners (i.e., growing emergence of mega funds), global pooling
of LPs (i.e., emergence of global LPs), and the development of more
futuristic structures based on the complete digitization of the investment
process and the wide-ranging dis-intermediation of GPs.
The fourth chapter broadly discusses the major external undercur-
rents that have emerged post COVID-response and are likely to impact
venture capital in terms of both existing portfolio firms and new invest-
ment opportunities. The six major trends detailed include changes to
industrial structures, alterations to consumer behavior, mass digitiza-
tion, the increased role of home as the center of human gravity, public
health apprehensions, and the abolition of the middle class in devel-
oped countries. Available investment opportunities in specific segments
of the economy are also examined as they pertain to each of these key
trends. Lastly, the chapter concludes by illuminating the likely sources
of venture capital transactions in the COVID era, which may include
the privatization of state firms, disposals from major multinationals, and
founders.
The venture capital industry has been under pressure post COVID-
response, which is a trend that is likely to continue in the future; it
is therefore valuable to explore changes to the venture capital invest-
ment process, which are analyzed in chapter five. Normal patterns of
venture capital’s daily operations, deal processing routines, regular habits
of communication, and other tacit behaviors that were previously estab-
lished in the pre-COVID era have been disrupted. Furthermore, state-led
COVID restrictions have affected the venture capital community differ-
ently from country to country, although the venture capital industry
generally went into a retreat-and-hide mode during March and April
of 2020. The situation improved in most countries by June, although
PREFACE xi
it was evident by the fall of 2020 that the venture capital community
would not be able to engage in its normal processes for the foresee-
able future because of the strict COVID-response measures that once
again emerged. This chapter discusses the many dislocations to the normal
patterns of processing venture capital deals, as well as the challenges expe-
rienced within each stage of the investment process (i.e., deal generation,
evaluation and screening, financial contracting, monitoring, and exiting).
Moreover, the chapter highlights one of the most profoundly affected
phases of the investment process, namely due diligence and monitoring,
and discusses problems related to portfolio firm underperformance and
valuation.
The sixth chapter of this book identifies the future of venture capital
performance (i.e., financial returns) by further analyzing the LP-GP rela-
tionship. The chapter begins with a discussion of fund formation and the
provision of capital by LPs to the venture capital ecosystem. It is stressed
that LPs can contribute to the deterioration of financial performance by
providing capital to suboptimal GPs, performing poor due diligence on
funds, tolerating limited access to information, and so on. The chapter
subsequently discusses the cost of “carried interest” and the provision
of venture capital services to LPs, which are two of the most expen-
sive components within the LP-GP arrangement. Moreover, the chapter
examines value chain analysis, provides another historical look at venture
capital performance, assesses the industry’s return prospects in the context
of a longitudinal study, and outlines the profiles of superior and inferior
GPs.
One of the most important components of the venture capital
ecosystem is the SME sector, which, as noted above, was one of the
hardest hit segments of the economy post COVID-response. Many
entrepreneurial firms were left financially devastated because their profits,
savings, and “sweat equity” disappeared within a matter of weeks or
months following the introduction of restrictive measures mandated by
public officials. Thus, the third major section is called “Entrepreneur-
ship Redefined Post COVID-Response” and consists of two chapters that
focus on the critical issues impacting the SME sector, including its ability
to access entrepreneurial finance.
The first chapter within the third section, or the seventh chapter
overall, concentrates on the entrepreneurial crisis created by COVID-
response; evidence suggests that up to 80 percent of firms were affected,
with about 40 to 50% of firms impacted so severely that they were unsure
xii PREFACE
of business survival. In the U.S., for example, the number of active busi-
nesses declined from 15.0 million to 11.7 million, which represents the
largest decrease in the number of operating firms in the U.S.’ modern
economic history. Furthermore, the harm to the SME sector was indi-
rectly related to the firm size (i.e., larger firms were less impacted by
COVID-response). Although firms operating within the SME sector are
typically flexible and agile in their ability to pivot and generate revenue
through different streams, their capacity to respond was thwarted by the
speed and strictness of public restrictions. The negative impact of public
restrictions has also affected different market segments to varying degrees,
as firms operating in retail, hospitality, restaurant and food services, arts
and entertainment, transportation, fitness, and leisure sectors have been
disproportionately impacted.
The changing landscape for entrepreneurs, particularly in terms of
their ability to procure entrepreneurial finance, is discussed in the eighth
chapter of this book. Research indicates that access to finance is one
of the most critical challenges for entrepreneurial firms, although there
are numerous other issues that have acutely impacted areas of the SME
sector post COVID-response. Primarily, many firms have faced a severe
decline, or a complete loss, of revenue. SME firms have also experi-
enced significant difficulties covering their fixed financial obligations. The
combination of revenue decline or loss with high fixed costs of busi-
ness operations has directly impacted the profitability, cash flow, and
liquidity of SMEs. Secondly, challenges to the businesses’ financial param-
eters have negatively influenced the value of entrepreneurial firms. A third
issue that has beset entrepreneurial firms post COVID-response has been
their ability to maintain personnel; while many firms initially aimed to
sustain their employees, they ultimately resorted to temporary or perma-
nent layoffs due to the firms’ abysmal financial position. The full impact
of these staffing reductions on unemployment will likely be manifested in
the next two or three years.
Additionally, it is noted in chapter eight that many firms from the SME
sector were forced to close while large “big box” stores remained open,
which created an unequal playing field for smaller firms. There is also
preliminary evidence to suggest that the SME sector’s ability to secure
access to entrepreneurial finance is further challenged post COVID-
response, as, for example, data from venture capital firms indicates that
deal volume is on the decline. Business angels have similarly reduced their
financing activities to instead focus on existing investments, while financial
PREFACE xiii
Future Research
Venture capital has been a topic of academic inquiry for a long period
of time. Debate on this subject has been both useful and productive; it
has provided many comprehensive perspectives on various aspects of the
venture capital investment process, including screening and evaluation,
financial contracting, and investee firm-GP relationships. Furthermore,
academic inquiry has not omitted some of the most difficult topics within
the industry, such as the divergence of interest between GPs and LPs,
declining venture capital performance, and the extent of GPs’ effective-
ness in their assistance of investee firms. However, due to current changes
to the venture capital industry as a result of COVID-response, academic
inquiry into venture capitalism is in many ways at a point of partial restart.
There are at least five critical areas of academic investigation that
should be considered moving forward, with the first and most important
area of research undoubtedly related to the redefinition of the relation-
ship between GPs and investee firms. Venture capital prides itself on the
provision of assistance to entrepreneurial firms, in spite of the often imper-
fect nature of this aid. As outlined in chapters three and five, this critical
relationship in the venture capital ecosystem has very evidently been rede-
fined post COVID-response. Thus, the key questions that should now be
asked are as follows: How effective are new modes of online interaction
between GPs and investee firms? Are GPs capable of providing effective
PREFACE xv
assistance to investee firms post COVID-response, and if so, what are the
crucial areas of assistance that investee firms require? Are there new areas
of investee firm assistance that have been previously overlooked by GPs?
A second fruitful area of new research should explore the future
of venture capital financial performance. In the past, there have been
multiple studies that provide a historical perspective on venture capital
performance, many of which reach differing conclusions as to whether or
not GPs are able to deliver returns in excess of those available in public
equities markets. If GPs are unable to generate repeatable returns that
exceed returns from public equities markets, the entire venture capital
ecosystem would continually be upset and thus subject to LPs satis-
fying their automatic “bucket filling” allocations in the asset class without
much reflection. The vital questions regarding venture capital perfor-
mance should be as follows: What are the key determinants of returns
across various types of private capital investing (i.e., buyouts, expansion
capital, seed and early-stage capital, etc.)? In what ways could the venture
capital change to generate better and more consistent returns? What are
the returns post COVID-response, and how do they differ from those
obtained during other economic crises? Has the divergence between top
quartile performance and the remaining GPs decreased or widened?
The third facet of study that has arisen post COVID-response specifi-
cally relates to the SME sector, which has disproportionately experienced
operational challenges, market uncertainties, financial difficulties, and
problems with financial management, which, in turn, have elevated the
sector’s difficulties in accessing finance. Important questions pertaining
to entrepreneurial finance could include: What are the key bootstrapping
strategies entrepreneurial firms can still rely on, and how has bootstrap-
ping changed for the SME sector? Can fintech fill the intermediate gap
by providing finance to entrepreneurial firms? Are there any new forms of
entrepreneurial finance that are likely to emerge post COVID-response?
What is the new role of the state in assisting the SME sector with regard
to the provision of finance? Are government loan guarantee programs
sustainable alternatives to bank lending?
A fourth area of possible research relates to actual GP operations. As
noted in chapters three and five, in-person interaction has been the bread-
and-butter of the venture capital industry; this form of interaction has
now been entirely redefined. Significant thought should be given to the
effectiveness of GPs’ interactions with their portfolio firms, LPs, and other
xvi PREFACE
Acknowledgments
I wrote this book during the period of subsequent COVID-response
restrictions (between January and May of 2021), after having read about
70 or so newspaper, magazine, and Internet-based articles on the topic
over the Christmas break; this was a useful starting point in my inquiry,
as it allowed me to dive further into the study of this new and ever-
changing subject. The daily grind of working on this project proved to
be interesting, challenging, and thought provoking. The old Benedictine
monks’ axiom of ora et labora has been valuable in keeping this project
moving forward.
While preparing the initial manuscript, I benefited from informal
(email) discussions with Thomas Meyer (SimCorp) and Mike Casey
(Portico). Discussion with Stephen Richmond (Abris Capital), who has
provided many favors over the years, has also been extremely beneficial; he
has been a go-to person whenever I needed feedback and insight from real
practitioners within the venture capital industry. I have relied heavily on
my previous research, contemplation, and writing while completing this
book, namely Strategic Entrepreneurial Finance and The Venture Capital
Deformation.
I would like to express gratitude to senior administrators at Brandon
University, and especially Steve Robinson, who have supported this book
project through a research appointment. I would also like to thank
PREFACE xvii
xix
xx CONTENTS
Index 277
List of Figures
xxi
List of Tables
xxiii
PART I
There are other reasons why venture capitalists may not be such
vigorous contributors to innovation, the first of which is because venture
capital’s profit maximization, short-term determinism, and exit orienta-
tion often result in less investment in long-term R&D, innovation, and
commercialization within entrepreneurial firms. In fact, venture capitalists
often insist on reducing expenses and capital expenditures associated with
innovation in order to maximize profits and cash flow, especially ahead
of a desired exit. Secondly, venture capitalists focus on an increasingly
narrow range of industries and therefore make a limited impact on the
broader economy. Venture capitalists frequently direct their capital toward
deals where the innovation cycles (i.e., innovation-to-commercialization)
are relatively short; thus, venture capital’s desired time frame to exit may
simply be too short to develop any long-lasting innovation. Additionally,
the cyclical nature of the venture capital industry itself may make any true
focus on innovation uneven, unsteady, unstable, and irregular.
Only a limited number of venture capitalists understand that true inno-
vation may come from advances in organizational structures, systems, and
approaches rather than technologically driven applications and services.
Of course, these organizational innovations cannot be patented and show-
cased. Instead, “breakthrough” products and services, which venture
capitalists often finance, are favored in spite of the fact that they may not
grant long-lasting leadership and competitive positions in the marketplace
because they can be quickly mimicked by a more efficient service provider
or manufacturer. Another claim regularly perpetuated by venture capital
is its ability to increase patents in investee firms, which is supposed to be
indicative of its contribution to innovation. Of course, the act of encour-
aging investee firms to register patents for unique products or services that
existed prior to venture capital participation should not be confused with
the creation of innovation that is frequently allocated to venture capital.
An increase in patents following an investment by venture capital may
be reflective of the fact that GPs are simply more effective and vigorous
in patenting activities. However, patenting may be unnecessary for the
majority of firms.
Finally, it is also important to comment on the general contribution
of venture capital to the local economy. Despite its frequent claims,
venture capital’s contribution to employment is questionable, mixed,
and controversial. Although venture capital firms frequently report the
number of employees employed at their investee firms, in addition to
1 VENTURE CAPITAL PRIOR TO THE AGE OF COVID 11
their firm’s revenue and profitability, they often fail to disclose the incre-
mental number of employees hired during their tenure. Research suggests
that venture capital’s contribution to incremental employment growth is
not homogenous and depends upon unique circumstances. For example,
management buyouts are likely to create jobs while management buy-ins
are prone to destroy them. Of course, as stated previously, venture capital
seeks to perpetually increase its short-term profits in a rapid manner, and
this near-sighted orientation is frequently in conflict with both increasing
costs (i.e., hiring more people) and investing in R&D.
a
2,500
Global venture capital 25%
20%
10%
1,500
$ billion
5%
1,000
0%
-5%
500
-10%
0 -15%
2000 2002 2004 2006 2008 2010 2012 2014 2016 2018 2020
Global illiquidity premium Global fundraising Global investing Cumulative dry powder
190
$ billion
140
90
40
-10
2003 2005 2007 2009 2011 2013 2015 2017 2019
Fig. 1.1 Key statistics in global venture capital (a) Global private equity
fundraising, investing, dry powder accumulation, and the illiquidity premium
(b) Key data in venture capital in emerging markets (Source Various sources,
including Prequin, Bain, Cambridge Associates, EMPEA. Updated from
Klonowski [2018])
achieved from public equities markets or alternative asset classes. The U.S.
S&P 500 index was selected as the benchmark for approximating these
returns from public equities markets.
Figure 1.1(a) illustrates a strong acceleration in global fundraising and
investing activities in the middle 2000s, followed by a rapid decline during
the 2008 financial crisis. In the last twenty years, between 2000 and 2020,
global venture capital firms raised $8,711.0 billion and invested $6,771.1
billion. While the average growth rates in fundraising and investing were
equal to 12.9% and 14.9%, respectively, over this twenty-year period, the
growth rates in recent years have been irregular, volatile, and inconsis-
tent. For example, between 2000 and 2010, growth rates in fundraising
accelerated by 18.0%, while in the following period (2011 to 2020) it was
equal to 6.1%, which represents a significant slowdown. The most signifi-
cant declines in fundraising occurred during and after the “dotcom” crisis,
resulting in a 34.5% decline in 2001, and after the 2008 financial crisis,
which amounted to a 53.3% decline. Other significant periods of decline
in fundraising occurred in 2003 (−19.8%), 2016 (−21.4%), and 2020
(−11.3%).
There have also been periods of rapid increase in fundraising, especially
following the dotcom correction (2004–122.6%; 2005–68.6%; 2006–
54.2%), as well as in the years after the 2008 financial crisis (2011–17.7%;
2012–15.1%; 2013–35.1%). Meanwhile, the average annual fundraising
activities have been equal to $414.8 billion. Investing activities show
similar growth and decline patterns, with the most significant decline in
investing occurring in 2009 by almost two-thirds from $121.0 billion to
$35.0 billion. However, investing activity bounced back to about $254.0
billion the following year in 2010. Additionally, it is important to note
that fundraising for younger entrepreneurial firms is one of the fastest
growing categories of broadly defined private equity. LPs and GPs gener-
ally question whether the industry has reached its peak, and if that is the
case, whether this peak is cyclical or more permanent.
Figure 1.1(a) confirms that in the period between the dotcom era
and the 2008 financial crisis, average fundraising and investing activ-
ities have been well matched. The capital deployment efficiency ratio
(CDER), which may be defined as a longitudinal ratio of cumulative
investing to fundraising, was equal to 91.5%. The above graph also clearly
demonstrates that since the 2008 financial crisis, fundraising has outpaced
investing; the CDER ratio post-2008 has been equal to 73.4 (although
the CDER ratio over the entire 20-year period has been equal to 77.7%).
18 D. KLONOWSKI
from, and when the returns were calculated. Some studies, which predom-
inantly focused on investments made in the 1980s and 1990s, illustrate
strong above-average returns; the period encompassing the early to mid-
1980s appears to be the most profitable era in the development of the
venture capital industry in the U.S., the U.K., and other countries.
In any case, it is important to note that the average GP has not been
able to generate the excess returns equal to between 30 and 50% that have
often been promoted to the public. For example, one study published by
the Kauffman Foundation, which provides an analysis of the foundation’s
investments into its GPs, reports that only about 25% of 100 GPs in the
foundation’s portfolio have been able to beat the returns from public
equities markets by at least three percentage points; nearly 50% of GPs
analyzed were not able to provide any returns on invested capital. Thus,
it is not surprising that further evidence shows a significant divergence in
the return performance between the top quartile and the rest of the GPs.
In other words, there is a significant drop-off in performance between
the top and remaining quartiles, and there is no doubt that these top
performers unduly influence the industry averages. To illustrate this, for
example, the top quartile GPs in 2013 in Europe were able to secure
net returns equal to 20.3%, while the second quartile GPs were only
able to achieve returns of 5.9%; this represents a difference of 14.4% in
performance between the first and second quartile performance.
Additional academic studies further dispute venture capital’s ability to
generate above-average returns. Figure 1.2 demonstrates a comparison of
academic studies on venture capital returns in comparison with returns
achieved from public equities markets. The studies are presented in the
following five columns: venture capital returns worse than public equi-
ties markets; equal to public equities markets; better than public equities
markets; better than public equities markets by more than three percent;
better than public equities markets by five percent. These studies indicate
that venture capital funds have not been able to exceed LPs’ minimum
requirements in any consistent manner. It is also prudent to note that the
right-hand side of the figure is quite empty.
Research furthermore confirms a discrepancy between returns gener-
ated by LPs and GPs, indicating that GPs may be able to generate positive
returns for themselves, but substantially lower returns for LPs. This is
inevitably due in part to GPs’ excessive fees and carried interest payouts,
which cut into LPs’ profits or net returns. Additionally, as noted above,
VC returns < public VC returns > public VC returns > public VC returns > public
24
VC returns ≈ public
returns returns returns, but by less returns, but by more returns, but by more
than 3% than 3% than 5%
(2012, JF)
Harris, Jenkinson & Kaplan
Sorensen, Wang & (2014, JF)
Yang
Kaplan & Schoar Phalippou (2014, RF)
(2014, RFS)
(2005, JF)
Cochrane Brophy & Guthner
Ivashina & Lerner (1988, JBV)
(2005, JFE)
(2020, JFE)
Other publications
(2012, report) Ljundgqvist &
Phalippou (2009, JEP) Richardson
(2003, unpublished)
Manigart, Joos &
Vos (1994, JSBF)
Fig. 1.2 A summary of academic studies on venture capital returns Abbreviations: AER American Economic Review,
FAJ Financial Analyst Journal, JAI Journal of Alternative Investments, JBV Journal of Business Venturing, JEP Journal
of Economic Perspectives, JIM Journal of Investment Management, JF Journal of Finance, JFE Journal of Financial
Economics, JFQA Journal of Finance and Quantitative Analysis, JPM Journal of Portfolio Management, JPE Journal
of Private Equity, JSBF Journal of Small Business Finance, RF Review of Finance, RFS Review of Financial Studies.
Updated from Klonowski (2018)
1 VENTURE CAPITAL PRIOR TO THE AGE OF COVID 25
100%
Percentage returns
80%
60%
40%
20%
0%
1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 2008 2010 2012 2014 2016 2018
Time
-20%
VC illiquidity premium VC returns S&P 500 PME returns LP's required 3% premium
30
25
Percentage returns
20
15
10
0
1986 1988 1990 1992 1994 1996 1998 2000 2002 2004 2006 2008 2010 2012 2014 2016 2018
-5 Time
-10
Fig. 1.3 Historical perspective on venture capital and private equity returns in
the U.S. (a) Returns from venture capital between 1986 and 2018 in the U.S.
(b) Returns from private equity between 1986 and 2018 in the U.S. (Source
Cambridge Associates. Updated from Klonowski [2018])
Besides the above, all of which are littoral or more or less shallow-water species,
we have another series of forms, or, to speak more correctly, we have two other
series of forms, from the deep Atlantic waters within the British area. In the cold
area of the Faeroe Channel we have Boreonymphon robustum, Bell; Nymphon
elegans, Hansen; N. sluiteri, Hoek; N. stenocheir, Norman; Colossendeis
proboscidea, Sabine; C. angusta, Sars. In the warm waters south and west of the
Wyville-Thomson ridge we have Chaetonymphon spinosissimum, Norman;
Nymphon gracilipes, Heller (non Fabr.); N. hirtipes, Bell; N. longitarse, Kr.; N.
macrum, Wilson; Pallenopsis tritonis, Hoek (= P. holti, Carpenter); Anoplodactylus
oculatus, Carpenter, and A. typhlops, G. O. Sars; and to the list under this section
Canon Norman has lately made the very interesting addition of Paranymphon
spinosum, Caullery, from the Porcupine Station XVII., S.S.E. of Rockall, in 1230
fathoms. Lastly, and less clearly related to temperature, we have Chaetonymphon
tenellum, Sars; N. gracilipes, Fabr.; N. leptocheles, Sars; N. macronyx, Sars; N.
serratum, Sars; and Cordylochele malleolata, Sars.
Of the species recorded in the above list as a whole, Anoplodactylus virescens,
Nymphon gracile, and Pallene spectrum reach their northern limit in the southern
parts of our own area; Ammothea echinata, Anoplodactylus petiolatus, Pallene
brevirostris, and Phoxichilus spinosus (or very closely related forms) range from the
Mediterranean to Norway, the last three also to the other side of the Atlantic;
Nymphon brevirostre and N. rubrum range from Britain, where they are in the
main East Coast species, to Norway. Of the Atlantic species, other than the Arctic
ones, the majority are known to extend to the New England coast.
INDEX
Abalius, 312
Abdomen, of Malacostraca, 110;
of Acantholithus, 178;
of Birgus, 176;
of Cenobita, 176;
of Dermaturus, 178;
of Hapalogaster, 178;
of Lithodes, 178;
of Pylopagurus, 178;
of Trilobites, 235;
of Scorpions, 297;
of Pedipalpi, 309;
of Spiders, 317;
of Palpigradi, 422;
of Solifugae, 426;
of Pseudoscorpions, 431;
of Podogona, 440;
of Phalangidea, 440, 443;
of Acarina, 457;
of Pentastomida, 489;
of Pycnogonida, 502
Abdominal glands, of Chernetidea, 432
Abyssal region (marine), 204;
(lacustrine), 209
Acantheis, 418
Acanthephyra, 163
Acanthephyridae, 163
Acanthoctenus, 415
Acanthodon, 388
Acanthogammarus, 138
Acantholeberis, 53
Acantholithus, 181;
A. hystrix, 178
Acanthophrynus, 313
Acari, 454 (= Acarina, q.v.)
Acaridea, 454 (= Acarina, q.v.)
Acarina, 258, 454 f.;
parasitic, 455;
external structure, 457;
spinning organs, 457;
internal structure, 459;
metamorphosis, 462;
classification, 464
Acaste, 249
Accola, 390
Acerocare, 247
Achelata, 529
Achelia, 534;
A. longipes, 506
Achtheres, 75;
A. percarum, 75
Acidaspidae, 251
Acidaspis, 226, 227, 230, 231, 235, 241, 251;
A. dufrenoyi, 250;
A. tuberculata, larva, 240;
A. verneuili, 231;
A. vesiculosa, 231
Aciniform glands, 335, 349
Acoloides saitidis, 367
Acroperus, 53;
A. leucocephalus, 52
Acrosoma, 410
Acrothoracica, 92
Actaea, 191;
habitat, 198
Actinopodinae, 387
Actinopus, 387
Aculeus, of scorpion, 303
Admetus, 313
Aegidae, 126
Aegisthus, 61
Aeglea laevis, 169;
distribution, 212
Aegleidae, 169
Aeglina, 227, 249;
Ae. prisca, 248
Agelena, 416;
A. brunnea, 367;
A. labyrinthica, 352, 353, 378, 380, 381, 416;
A. naevia, 339
Agelenidae, 325, 352, 353, 415
Ageleninae, 416
Aggregate glands, 335, 349
Aglaspis, 279
Agnathaner, 66
Agnathonia, 529
Agnostidae, 244
Agnostini, 243
Agnostus, 222, 223, 225, 231, 234, 245;
A. integer, 245
Agraulos, 247
Agroeca, 397;
A. brunnea, cocoon, 358
Albunea, 171;
respiration, 170;
distribution, 201
Albuneidae, 171
Alcippe, 92;
A. lampas, 92, 93
Alcock, on Oxyrhyncha, 192;
on phosphorescence, 151
Alepas, 89
Alima, larva of Squilla, 143
Alimentary canal, of Crustacea, 14;
of Phyllopoda, 28;
of Cladocera, 42;
of Squilla, 142;
of Malacostraca, 110;
of
Trilobites, 222;
of Arachnida, 256;
of Limulus, 268;
of Scorpions, 304;
of Pedipalpi, 310;
of Spiders, 329;
of Solifugae, 427;
of Pseudoscorpions, 134;
of Phalangidea, 444;
of Acarina, 459;
of Tardigrada, 480;
of Pentastomida, 491;
of Pycnogous, 513
Alitropus (Aegidae), habitat, 211
Allman, on larvae of Pycnogons, 523
Alloptes, 466
Alona (including Leydigia, Alona, Harporhynchus, Graptoleberis),
53
Alonopsis, 53
Alpheidae, 163;
habitat, 198
Alpheus, 163;
reversal of regeneration, 156
Alveolus, of palpal organ of Spiders, 322
Amaurobius, 399;
A. fenestralis, 399;
A. ferox, 399;
A. similis, 399;
spinnerets, 326
Amblyocarenum, 388
Amblyomma, 470;
A. hebraeum, 456, 470
Amblypygi, 312
Ammothea, 505, 534;
A. achelioides, 534;
A. brevipes, 541;
A. echinata, 505, 509, 510, 534, 541, 542;
A. fibulifera, 522, 534, 541;
A. franciscana, 541;
A. grandis, 534;
A. hispida, 534, 535, 541;
A. laevis, 541;
A. longicollis, 533;
A. longipes, 506, 534, 541;
A. magnirostris, 534, 541;
A. typhlops, 542;
A. uniunguiculata, 534
Ammotheidae, 534
Amopaum, 452
Ampharthrandria, 61
Amphascandria, 57
Amphion, 251
Amphipoda, 136 f.;
pelagic, 202;
fresh water, 211
Ampullaceal glands, 335, 349
Ampycini, 243
Ampyx, 231, 245;
A. roualti, 230
Anabiosis, in Tardigrada, 484
Analges, 455, 466
Analgesinae, 466
Ananteris, 306
Anaphia, 539
Anaspidacea, 115;
distribution, 211, 217
Anaspidae, 89
Anaspides, 115, 117;
relation to Schizopoda, 112;
distribution, 211;
A. tasmaniae, 115, 116;
habitat, 211
Anaspididae, 115
Anelasma squalicola, 89
Anelasmocephalus, 452
Angelina, 247
Anisaspis bacillifera, 387
Anisopoda, 122
Anomalocera pattersoni, 60;
distribution, 202, 203
Anomopoda, 51
Anomorhynchus, 532
Anomura, 167;
relation to Thalassinidea, 167
Anoplodactylus, 511, 538;
A. lentus, 524;
A. neglectus, 539;
A. oculatus, 542;
A. petiolatus, 508, 510, 539, 541, 542;
A. virescens, 540, 542
Anopolenus, 247
Antarctic zone (marine), 200
Antarctica, evidence on, 200, 217
Antennae, of Crustacea, 5, 8;
of Phyllopoda, 24;
of Cladocera, 37;
of Copepoda, 55;
of Cirripedia, 81 f.;
of Ostracoda, 107;
of Malacostraca, 110;
of Anomura, 168;
of Corystes cassivelaunus, 170, 183, 189;
used in respiration, 170;
of Trilobites, 237
Antennary gland, 13 (= green gland, q.v.)
Anthrobia, 406;
A. mammouthia, 334, 366
Anthura, 124
Anthuridae, 124
Ants and spiders, 370
Anyphaena accentuata, 397
Aphantochilinae, 414
Aphantochilus, 414
Apoda, 94
Apodidae, 19, 21, 22, 23, 27, 28, 29, 31, 36, 241
Aponomma, 470
Appendages (incl. legs, limbs), of Crustacea, 7;
of Entomostraca, 18;
of Phyllopoda, 24;
of Cladocera, 40;
of Copepoda, 55;
of Cirripedia, 80 f.;
of Ostracoda, 107;
of Malacostraca, 110;
of Nebalia, 111;
of Eumalacostraca, 113;
of Anaspides, 115;
of Mysidacea, 118 f.;
of Cumacea, 120;
of Isopoda, 121 f.;
of Amphipoda, 136 f.;
of Stomatopoda, 142;
of Euphausiacea, 144 f.;
of Decapoda, 152;
of Macrura, 153;
of their larvae, 159;
of Anomura, 167 f.;
of Birgus, 175;
of Brachyura, 181 f.;
alterations caused by parasites, 100 f.;
by hermaphroditism, 102 f.;
of Trilobita, 236, 237;
of Arachnida, 255 f.;
of Limulus, 262, 263;
of Eurypterus, 285 f.;
of Scorpions, 301, 303;
of Pedipalpi, 309;
of Spiders, 319;
of Palpigradi, 422;
of Solifugae, 426;
of Pseudoscorpions, 432;
of Podogona, 440;
of Phalangidea, 443;
of Acarina, 458;
of Tardigrada, 479;
of Pentastomida, 493;
of Pycnogons, 503 f.
Apseudes spinosus, 123
Apseudidae, 122
Apstein, 335
Apus, 21, 23, 25, 28, 30, 32, 34, 36, 221, 242, 243;
segmentation, 6;
A. australiensis, 36;
A. cancriformis, 36;
habitat, 34
Arachnida, introduction to, 255;
segmentation of body, 255–6;
primitive, 256–7;
coxal glands, 257;
endosternite, 257;
sense-organs, 257;
classification, 258
Araneae, 258, 314 f.
Araneida, 314
Araneina, 314
Araneus, 408 n.
Aratus pisonii, 195
Arbanitis, 388
Archaeolepas, 84;
A. redtenbacheri, 84
Archea, 411;
A. paradoxa, 383;
A. workmani, 411
Archeidae, 321, 411
Archisometrus, 306
Arctic zone, 199
Arcturidae, 127
Arcturus, 127
Arcyinae, 410
Arcys, 410
Arethusina, 223, 230, 251;
A. konincki, 250
Argas, 457, 469;
A. persicus, 469;
A. reflexus, 469
Argasidae, 469
Arges, 252
Argiope, 408;
A. aurelia, 340, 379;
A. bruennichi, 408;
A. cophinaria, 349, 365;
A. trifasciata, 408
Argiopidae, 406 n.
Argiopinae, 408
Argulidae, 76
Argulus foliaceus, 77
Argyrodes, 402;
A. piraticum, 367;
A. trigonum, 367
Argyrodinae, 402
Argyroneta, 336, 415;
A. aquatica, 357, 415
Ariadna, 395
Ariamnes, 402;
A. flagellum, 318
Arionellus, 247
Aristaeus, 162;
A. crassipes, 159;
A. coruscans, phosphorescence, 151
Armadillidium, 129
Artema, 401
Artemia, 23, 24, 35;
A. fertilis, anal region, 23;
head, 26;
limb, 27;
A. salina, 23, 33, 36;
A. urmiana, 23
Arthrolycosa antiqua, 383
Arthropoda, 4;
segmentation, 7;
a natural group, 17
Arthrostraca, 121
Asagena, 404
Asaphellus, 249
Asaphidae, 249
Asaphini, 243
Asaphus, 222, 225, 227, 229, 235, 236, 249;
A. cornigerus, 227;
A. fallax, eye, 228;
A. kowalewskii, 227;
A. megistos, 236;
A. platycephalus, 236
Ascidicola rosea, 66
Ascidicolidae, 66
Asconiscidae, 130
Ascorhynchus, 505, 533;
A. abyssi, 506, 509, 519;
A. cryptopygius, 513 n.;
A. minutus, 517;
A. ramipes, 513 n.
Ascothoracica, 93
Asellidae, 128
Asellota, 127
Asellus, 127;
habitat, 209, 211;
A. aquaticus, 127, 209;
A. cavaticus, 209, 210;
A. forelii, 209
Aspidoecia, 76
Astacidae, 157;
distribution, 213, 216
Astacoides, 157;
distribution, 213
Astacopsis, 157;
distribution, 213;
A. franklinii, 214
Astacus, 104, 157;
appendages, 10;
distribution, 213;
hermaphroditism, 104
Astacus gammarus (= Homarus vulgaris), 154
Asterocheres violaceus, 67
Asterocheridae, 67
Asterope oblonga, 108
Astia, 421;
A. vittata, 381
Astigmata, 465
Astridium, 540
Atax, 462, 472;
A. alticola, 472;
A. bonzi, 472
Atelecyclidae, 190
Atelecyclus, 191;
respiration, 189
Atops, 247
Attidae, 376, 381, 419
Attus, 421;
A. pubescens, 372, 421;
A. saltator, 372, 421
Atya, 163
Atyephyra, 163;
habitat, 210
Atyidae, 159, 163;
distribution, 212
Atypidae, 390
Atypoides, 391
Atypus, 391;
A. abboti, 356;
A. affinis, 356, 391;
A. beckii, 391
Auditory organ, of Anaspides, 116;
of Decapoda, 153;
of Mysidae, 119
Augaptilus filigerus, 59
Austrodecus glacialis, 535
Austroraptus polaris, 535
Autotomy, 155
Avicularia, 389
Aviculariidae, 316, 327, 386;
bite of, 365;
poisonous hairs of, 365
Aviculariinae, 389
Axial furrows, 223
Baglivi, 361
Baikal, Lake, Crustacea of, 212
Balanus, 91;
B. porcatus, shell, 90;
B. tintinnabulum, 91;
anatomy, 90
Ballus variegatus, 420
Barana, 506, 513, 533;
B. arenicola, 512, 513, 533;
B. castelli, 512, 513 n., 533
Barnacles, origin of term, 79
Barrande, J., on development of Trilobites, 238;
on their classification, 243
Barrandia, 249
Barrois, 435 n.
Barrus, 429
Barychelinae, 389
Basse, on Tardigrada, 481
Baster, Job, 503
Bates, 373
Bathynomus giganteus, 126;
habitat, 205
Bathynotus, 247
Bathyphantes, 406
Bdella lignicola, 471
Bdellidae, 458, 471
Beecher, C. E., on facial sutures of Agnostus and Olenellus, 225;
on development of Trilobites, 238;
on their classification, 243
Beetle-mites, 467
Beetle-parasites, 470
Belinurus, 275, 279;
B. reginae, 278
Belisarius, 308
Belt, 368, 371
Beltina, 283 n.
Bernard, 311, 424, 426, 433 n., 434 n.
Bertkau, 323, 365, 395 n.
Beyrich, E., on facial suture of Trinucleus, 226
Billings, E., on appendages of Trilobites, 236
Bipolarity, 200
Birds and Spiders, 370
Birds’ feather Mites, 466
Birgus, 181;
B. latro, habits, 174;
structure, 175, 176
Black Corals, Cirripedia parasitic on, 93, 94
Blackwall, 348, 359 n., 365, 368, 385
Blindness, in Crustacea, 149, 209, 210;
in Spiders, 334
Blood, haemoglobin supposed in, 30, 68
Boas, on classification of Malacostraca, 113
Boeckella, distribution, 216
Boeckia, 138
Böhmia, 535
Bolocera, Pycnogonum with, 524
Bolyphantes, 406
Bomolochidae, 71
Bomolochus, 71, 72
Bon, 360
Bont-tick, 456
Boophilus, 456, 469;
B. australis, capitulum of, 468
Bopyridae, 130, 133
Bopyrina, 129, 130, 132
Bopyrus fougerouxi, 133;
male, 133;
adult female, 134
Bopyrus larva, of Bopyrina, 129, 133
Boreomysis, 120;
B. scyphops, distribution, 201
Boreonymphon, 536;
B. robustum, 506, 507, 511, 512, 542
Bosmina, 52, 53;
occurrence in Southern hemisphere, 216;
B. longirostris, habitat, 206
Bosminidae, 53;
appendages, 41;
alimentary canal, 42
Bothriuridae, 306, 308
Bothriurus, 308
Bouvier, 528 n.
Boys, 348, 360, 376
Brachybothrium, 391
Brachymetopus, 251
Brachythele, 390
Brachyura, 181;
eyes, 150
Branchiae (= gills) of Crustacea, 16;
of Decapoda, 152;
of Limulus, 269;
of Eurypterids, 288
Branchinecta, 25, 35;
B. paludosa, 35;
range, 34
Branchiopoda, 18 f.
Branchiopodopsis, 35;
B. hodgsoni, 35
Branchiostegite, 152
Branchipodidae, 19, 22, 35, 241