Professional Documents
Culture Documents
Contemporary Irish
Popular Culture
Transnationalism, Regionality, and Diaspora
Anthony P. McIntyre
Film
University College Dublin
Dublin, Ireland
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For Maria and Annie
Acknowledgements
I’d like to thank all those who helped in producing this book. First, thank
you to Camille Davies, Jack Heeney and Imogen Higgins at Palgrave
Macmillan for all their support. Two anonymous reviewers provided excel-
lent suggestions that have improved the final work considerably. Valuable
feedback was also provided by audiences at several academic conferences
and seminars where I presented papers that would eventually find their
way in modified form into this book. I’d like to thank audiences and
speakers at the L’Irlande en series conference at Université Paris Ouest,
Nanterre in 2014; the Post Celtic-Tiger Irishness Symposium at Trinity
College Dublin in 2016; the Global Irish Diaspora Congress at University
College Dublin in 2017; the Sports, Media and the Cultural Industries in
Ireland Symposium at Dublin City University in 2018; and the European
Popular Culture Association Conference 2019 at the University of
Limerick.
I am sincerely grateful to Diane Negra who, throughout my time at
UCD has been an exemplary colleague and friend and who provided gen-
erous feedback throughout the development of this book. My research
with Diane and Eleanor O’Leary on aspects of Irish contemporary culture
has taken place alongside my work on this book and has informed it con-
siderably. David McKinney, Marcus Free and Colin Coulter read work in
progress at different points and offered valuable advice. Any errors within
the book are, of course, my own.
Portions of the present work have appeared in earlier publications.
Several sections in Chap. 2 develop ideas that appeared in a version origi-
nally published as McIntyre, Anthony P., “Moone Boy and the Elision of
vii
viii ACKNOWLEDGEMENTS
ix
x CONTENTS
Index251
List of Figures
xi
xii LIST OF FIGURES
Fig. 6.1 The Fricker Irish mammy memes utilise Brenda Fricker’s role
as Mrs. Brown in My Left Foot, and in particular her stern
expression, connoting both the domesticity associated with
previous eras and the unglamorous ‘common sense’
of the figure 211
Fig. 6.2 Posters for the cinematic release of Philomena conflate the
Irish mammy with mobility through a white doodle on a
plain yellow background that summarises the journey taken
in the movie 229
Fig. 7.1 In “Strangers on a (Dublin) Train” emerging interpersonal
protocols of the pandemic era are recast as novel and romantic 242
CHAPTER 1
Introduction—“Fractured Movement”:
Transnationalism, Regionality, and Diaspora
in Contemporary Irish Popular Culture
at the center of social life” (3). It is this influence across multiple spheres,
as well as the adaptability of this variant of capitalism that is key to its per-
sistence, despite the notable social and financial catastrophes with which it
has come to be associated, not least the “Great Recession” of 2007–2009
and the austerity policies that swiftly followed in its wake. Jamie Peck and
Nik Theodore (2019) suggest the necessity of “confront[ing] neoliberal-
ism as an emergent mode of regulation, one that has become cumulatively
embedded across multiple sites and spaces such that it increasingly defines
the rules of the game and the terrain of struggle, even if never acting alone
or monopolizing that terrain.” (Peck and Theodore 2019, 246). The pur-
pose of this book is to contribute to a growing body of Irish cultural stud-
ies scholarship that seeks to interrogate the role of popular culture in
installing neoliberal values as common sense (see, for instance, Kiersey
2014; Brick and Davidson 2017; Free and Scully 2018; Negra and
McIntyre 2020; McIntyre 2021).
For scholars such as Peck and Theodore, neoliberalism is never a mono-
lithic construct, but one that is variegated. This plasticity, in their account,
should act as “an invitation to conjunctural analysis, sensitive to variable
(local) projects, formations, struggles, and contestations, and at the same
time recognizing the openness of emergent pathways and future hori-
zons” (246). This approach aligns with scholarship on Irish manifestations
of this economic model. Geographers Rob Kitchin et al. (2012) in an
influential article tracing the economic and spatial impacts of neoliberal-
ism suggest that Ireland differed from the UK and the US, where neolib-
eralism was “an ideologically informed project” instigated under the
premierships of Margaret Thatcher and Ronald Reagan. For Kitchin et al.,
“Irish neoliberalism was produced through a set of short-term (intermit-
tently reformed) deals brokered by the state with various companies, indi-
viduals, and representative bodies, which cumulatively restructured Ireland
in unsustainable and geographically ‘uneven’ ways” (1306). Nevertheless,
while neoliberalism may not have been imposed upon the Irish in the top-
down manner of the US and the UK, the role of media figures and the
culture industries more broadly was also crucial to the shifting ideological
consensus that helped broker neoliberalism’s acceptance in a newly secula-
rised Ireland and a post-conflict Northern Ireland.
Irish studies scholar Joe Cleary (2018) suggests Ireland’s reputation for
“creativity”—a notable neoliberal buzzword, and one which, as I detail in
Chap. 5 has been seized upon with gusto by the Irish state—combined
with several other factors (a mobile, educated workforce; lack of
1 INTRODUCTION—“FRACTURED MOVEMENT”: TRANSNATIONALISM… 7
in cultural studies, media studies and Irish studies. The conjunctures that
this book intersects with, and which are associated in some ways with the
populist currents that Volcic and Andrejevic (2016) contend are precipi-
tated by globalist and neoliberal tendencies are Trumpism in the US,
Brexit in the UK, (both of which I consider in the remainder of this sec-
tion) and a post-Celtic Tiger Ireland characterised for many years by aus-
terity and economic migration. The global COVID-19 pandemic that
swept the world in 2020 is dwelt upon in the coda of this book.
The Brexit referendum of 2016 and Britain’s subsequent departure
from the European Union demonstrates the complex interplay between
nation states. The sundering of the UK from the EU has reignited ten-
sions in Northern Ireland given the Irish border’s sudden reconfiguration
as a boundary not only between Britain and Ireland, but between Britain
and the EU. This momentous political recalibration has undermined the
1998 Good Friday Agreement that marked the putative end of The
Troubles in Northern Ireland and the beginning of a power-sharing sys-
tem underwritten by British and Irish state support. The fact that the UK
has historically been the destination to which a majority of Irish emigrants
depart contributes also to the fraught relations between the two nations.
Certain figures and texts analysed herein (footballer McClean, Derry Girls
[2018–]; Mrs Brown’s Boys [2011–] in Chap. 6; sitcoms focusing on the
Irish in Britain such as Catastrophe [2015–2019] and This Way Up [2019]
in Chap. 5) all register in complex ways the seismic shifts under way in the
second decade of the 2000s.
In particular, my examination of McClean’s refusal to wear the Earl
Haig poppy year on year, while playing in the English football leagues
demonstrates how a personal protest informed by the player’s experience
of British militarism on the streets of Derry became a cultural lightning
rod. In many ways McClean’s protest (begun in 2012) prefigured the rise
of a virulent nationalism in the UK that was tied in complex ways to
notions of remembrance and the valorisation of sacrifices made in conflict,
in particular the First World War, and which manifested in its most potent
expression in the Brexit vote to take the UK out of the European Union.
The vote has compounded uncertainties in terms of national belonging
that have emerged for Irish and other national subjects in these years,
uncertainties often rooted in shared and troubled histories of colonialism
and migration.
It is important to clarify at this point that the transnational shifts I track
in this book are for the most part related to anglophone countries, and
1 INTRODUCTION—“FRACTURED MOVEMENT”: TRANSNATIONALISM… 11
primarily the interplay between Ireland, the US and the UK. In many ways
this is due to the triangulation of diasporic movement between these
nations, as well as the dominance of both the UK and the US in terms of
cultural production pertaining to Ireland. Sociologist Mary J. Hickman
(2002) emphasizes this connection in her consideration of the definition
and impact of the Irish diaspora, characterising it as:
trans denotes both moving across space or across lines, as well as changing
the nature of something. Besides suggesting new relations between nation
states and capital, transnationality also alludes to the transversal, the transac-
tional, the translational, and the transgressive aspects of contemporary
behaviour and imagination that are incited, enabled, and regulated by the
changing logics of states and capitalism. (4)
1 INTRODUCTION—“FRACTURED MOVEMENT”: TRANSNATIONALISM… 13
culture” that I deploy in the title to this book is also not without its diffi-
culties, as Stuart Hall (1981) cautions. For my purposes, I take the term
to designate the realm of cultural activity that while reaching a broad audi-
ence, still tends to be conceptualised as “low culture,” far from the rarefied
realm of Art, Literature or even cinematic auteurism. While the discipline
of Irish studies has begun, in fits and starts, to see such texts as advertising,
sitcoms, non-traditional sports broadcasts, reality television, celebrity cul-
ture and memes as within its purview, the present study contends such an
approach is essential to any account of contemporary Irishness and the
shifts it has undergone in the twenty-first century.
While the archive I assemble is somewhat variegated, certain genres or
modes find greater representation. Comedy, whether in television sitcom
form or associated hybrid genres of comedy-drama or dramedy, is a key
feature of this book. This, in part, is related to comedy’s capacity, similar
indeed to that of sports, to generate feelings of togetherness. As Alenka
Zupančič (2020) has suggested, “Laughter is not only or simply an expres-
sion of individual relief and pleasure, it is decidedly a collective-forming
affect, more so perhaps than any other” (2020, 281). Similarly, for Andy
Medhurst (2007), “Above all else, comedy is an invitation to belong”
(2007, 19). That comedy can foster a sense of collectivity and belonging,
also registers in an opposing formation. As some feel included or part of a
collective, others will also feel excluded. Mrs Brown’s Boys, a sitcom exam-
ined in Chap. 6, is perhaps particularly exemplary of this facet of comedy,
given the strong polarisation the sitcom provokes, which skews along gen-
erational and regional lines.
My focus on the popular aligns with the multiple scalar levels with
which this study is concerned. As geographer Jason Dittmer (2005) argues
“Popular culture is one of the ways in which people come to understand
their position both within a larger collective identity, and within an even
larger geopolitical narrative, or script” (626). This aspect of popular cul-
ture is essential for this book’s purposes, given its concern with Irishness
on a regional, national, and transnational level. Part of my argument in
this book centres on the expanded reach of contemporary communica-
tions channels. So, for instance, the sitcom Derry Girls, originally broad-
cast on Channel 4 in the UK, which I examine in Chap. 3 has an expanded
reach since it has been made available internationally on streaming video
on demand (SVOD) provider Netflix. This implications of this are that the
vernacular language and specific accents that index the sitcom’s regionality
are foregrounded, a feature of the sitcom that can both elicit positive and
1 INTRODUCTION—“FRACTURED MOVEMENT”: TRANSNATIONALISM… 19
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CHAPTER 2
This chapter, and several of the others to follow, suggests that an analysis
of contemporary iterations of stardom and celebrity provides a useful
prism through which to view shifts in wider Irish society in the post-Celtic
Tiger era. The heightened individualism of stardom/celebrity allows for
an analysis that can span multiple textual formations, from journalistic
portrayals of the individual in question and social media postings on the
part of the celebrity, to media content (films and television programmes,
for instance) featuring the figure, and the critical and popular reception of
these. This form of cultural studies approach, when used in tandem with
interdisciplinary scholarship drawn from academic fields including
Sociology, History and Politics enables the generation of insights derived
from multiple axes of analysis. To that end, the present chapter considers
the career of Irish actor Chris O’Dowd as a site of embodied social knowl-
edge regarding Ireland and its diasporic populations in the twenty-first
century.
O’Dowd—born in Sligo, raised in neighbouring Roscommon and now
living in Los Angeles, having worked in London for a substantial portion
of his professional career—is an example of the diasporic and transnational
tendencies that are often central to a successful career in the performing
arts. Although Ireland in many ways has an arts scene that belies the coun-
try’s comparatively small size, cosmopolitan centres of cultural production
such as London and Los Angeles have long exerted a powerful draw upon
those trying to make their way in the creative industries (Barton and
Murphy 2020). The flow of aspirants seeking a place in these industries
mirrors a wider transnational movement among the Irish population with
well-established roots. It has been argued that a recourse to emigration is
almost instinctive in the Irish, especially in times of economic duress, with
the exodus of prior generations having established a symbolic and material
infrastructure for Irish people to leave their homeland for professional,
economic, or other reasons (O’Toole 2016). The present chapter, through
an examination of O’Dowd’s professional career limns some of the con-
tours of contemporary Irish diasporic life patterns and the ideological
shaping of this constitutive element of Irishness, particularly as they mani-
fest in popular culture.
An established body of scholarship contends that the mediated con-
struction of stars and celebrities, figures who hold a prominent position
within an ever-expanding mediascape, undergird notions of selfhood and
subjectivity. The significant decline in the influence of traditional reli-
gion—Catholicism predominantly in the case of the Republic of Ireland—
has seen stardom and celebrity partially fill the void left through the
provision of models of behaviour, however problematic in their own ways
these new social configurations may be. For sociologist Chris Rojek
(2001), in this context, celebrity functions as one of several “replacement
strategies that produce new orders of meaning and solidarity,” with celeb-
rity culture, in the main, constituting “a significant institution in the nor-
mative achievement of social integration” (99). In a similar vein, P. David
Marshall (1997) claims that “celebrities represent subject positions that
audiences can adopt or adapt in their formation of social identities” (65),
a process he sees as providing “an embodiment of collective configurations
within individual representations” (51). Taking the nation as such a col-
lective configuration, we can conceive that in a country such as Ireland,
with its long histories of migration and substantial diasporic communities,
the embodiment of a transnational identity that unites a sense of belong-
ing within the home nation with the possibilities afforded by relocation is
one of the key functions of contemporary Irish celebrity.1
As outlined in the introductory chapter, Ireland has for some time pur-
sued economic policies predicated on securing outside investment and
developing an open economy that positions the nation as the gateway to
Europe for US corporations. This, and the dispersal of a populace that is
also shaped by such economic strategies—predicated as they are on the
transnational flow of people, material goods and capital—is reflected in
2 STAR LEVERAGE, LOCAL MATTERS, AND TRANSNATIONAL MEDIA: CHRIS… 25
Film scholar Ruth Barton (2006) has noted the centrality of male star
discourse to contemporary notions of Irishness. While one might assume
the emergence of Oscar-nominated stars such as Saoirse Ronan and Ruth
Negga suggests a redress of the gender balance in terms of international
stardom is currently under way (see Barrett 2015), male stars still outnum-
ber their female compatriots by a considerable measure. O’Dowd is, along
with Domhnall Gleeson, Andrew Scott, and Jamie Dornan, part of a gen-
erational cohort of prominent male Irish stars born in the late seventies
and early eighties who move fluidly between television and cinema in the
twenty-first century. O’Dowd has not achieved the consistent headline
billing attained by more high-profile stars like Michael Fassbender and
Colin Farrell, yet arguably this distance from the apex of the star system
renders him a more approachable and demotic form of star, in sync with
an era in which the longstanding hierarchies that obtain within the film
industry are under increasing duress.
After working as a jobbing actor in the UK for several years, O’Dowd
first came to prominence, in the UK and Ireland, playing the hapless soft-
ware support worker Roy in fellow Irishman Graham Linehan’s sitcom
The IT Crowd (2006–2013). The role would prove pivotal in the actor’s
career not only for giving him his first critical success, but also for facilitat-
ing his big break in the US. Bridesmaids’ director Paul Feig was an IT
Crowd fan and asked O’Dowd to try the audition for the film in his own
accent after an initial try in an American one failed to impress (Solomons
2011). The incident demonstrates the growing importance of transna-
tional creative networks predicated on specific genre knowledge and
appreciation—a phenomenon I deal with in more detail in considering the
2 STAR LEVERAGE, LOCAL MATTERS, AND TRANSNATIONAL MEDIA: CHRIS… 27
from elitist notions of stardom and Hollywood glamour, which are increas-
ingly associated with punishing regimes of bodily regulation.
This aspect of the O’Dowd’s appeal is often foregrounded in media
profiles of the actor. An April 2014 issue of GQ magazine, for instance,
features O’Dowd on the cover alongside a beautiful glamour model with
the tagline “Frankly, we can’t believe Chris O’Dowd’s luck either!”
(Fig. 2.1) The cover, and tagline, suggests that part of the star’s appeal is
his evocation of an ordinariness that secures empathy with audiences, with
the “we” of the headline interpellating an audience who might, by similar
quirk of fate, end up in O’Dowd’s “lucky” position. This narrative of an
everyman finding himself incongruously situated among the elite of the
entertainment industry is further emphasized through the self-deprecating
humour O’Dowd employs in media interviews.
Such self-deprecation is evident in a 2012 interview with Conan
O’Brien on his TBS talk show Conan (2010–2021), in which the actor is
promoting the comedy This is 40. O’Dowd discusses at length his tall phy-
sique and how unusual his body looks naked, joking that it has led to vari-
ous nude scenes being omitted from films he has acted in, and drawing a
contrast between his own physique and the petite frame of Megan Fox,
the glamorous co-star with whom he shared a poolside scene in the movie.
The interview also aligns O’Dowd with the similarly built physique of the
host, and O’Brien’s own rhetorical mainstay of an ethnicized bodily self-
deprecation (usually regarding his red hair or fair skin). A later interview
on the same show in May 2020 has O’Brien clearly savouring and imitat-
ing O’Dowd’s pronunciation of his dog Potato’s name. Both the ethni-
cally inflected corporeal comparisons and accent appreciation and mimicry
on display in these interviews foreground the connections between Irish
and Irish-American performative modes and serve as a public attestation
of good will between Irish diasporic subjects of different generational
removes from the home nation.2
O’Dowd’s demotic positioning undergirds his emergence as a post-
Celtic Tiger figure, distanced from the perceived hedonistic excesses of
Irish actors such as Colin Farrell and Jonathan Rhys Meyers, physically
attractive performers who were among the most prominent exemplars of
Irish male masculinity on screen at the outset of the twenty-first century.
The well-documented issues with alcohol and drug misuse that have
tainted the careers of Farrell and Rhys Meyers, leading to public disavow-
als of past behaviours and habits on the part of both actors, are not a fea-
ture of O’Dowd’s star discourse. In part this is due to a multi-mediated
30 A. P. MCINTYRE
and coherent screen presence on the part of O’Dowd. While O’Dowd has
had some prominent examples of public drunkenness, particularly on an
episode of British chat show The Last Leg (2012–) in October 2018,
2 STAR LEVERAGE, LOCAL MATTERS, AND TRANSNATIONAL MEDIA: CHRIS… 31
public opinion about the appearance was generally favourable. The actor
had been noticeably inebriated, but in a very good-humoured manner,
telling a rambling joke to the evident amusement of the hosts and fellow
guests on the show. O’Dowd in this role personifies Irish conviviality and
humour and the darker associations between alcohol and Irishness are
attenuated. The seeming shift in popularity from stars imbued with a
hedonistic quality to O’Dowd’s more everyman characteristics suggest the
complex ways in which stardom works as a barometer of social processes.
It also attests to the labile quality of Irish performativity, a somewhat capa-
cious mode of expression that nevertheless tends to operate within a fixed
set of coordinates. The shift from hedonism to the demotic qualities with
which O’Dowd is associated roughly aligns with a post-Celtic Tiger
national sense conveyed in national discourse, evident across a range of
media, that the Irish had “lost the run of ourselves” in the preceding
boom period (Free and Scully 2018).
The incident on The Last Leg also demonstrates O’Dowd’s facility for
correcting public missteps and the latitude afforded the performer on
account of his gregarious good humour. The actor is a regular user of
social media platform Twitter, and he posted the day after the production
expressing both contrition and bemusement at the incident. Responding
to a Twitter user who, unlike many of the responders to his tweet, criti-
cised the actor’s performance as “embarrassing” and “distasteful,”
O’Dowd posted, “I am a little embarrassed to be honest. But to keep
things in perspective, it’s a tipsy comedian on a late-night chat show on
Channel 4. I wasn’t doing meth on bbc breakfast. But thanks for your
concern, genuinely” (Starkey 2018). Similarly, O’Dowd used humour
during a podcast interview with broadcaster Louis Theroux to walk back
his role in the recording of John Lennon’s “Imagine” organized by Israeli
actress Gal Gadot when many parts of the world went into lockdown due
to the COVID-19 pandemic. The viral video, featuring a number of celeb-
rities singing along to the classic song, was publicly reviled as “tone deaf”
in conception, given its seeming equation between the lives of Hollywood
stars holed up in their mansions with the plight of others in more strait-
ened circumstances. After laughing through the host’s straight-faced
description of the song as “proving divisive on the internet,” and his sub-
sequent jibes, O’Dowd characterises the video as part of “that first wave of
creative diarrhoea that seemed to encase the world [at an early point in the
pandemic],” and further concedes that “any backlash was fairly justified”
(“Grounded with Louis Theroux 10. Chris O’Dowd” 2020). This mixture
32 A. P. MCINTYRE
we got the script done, then improvised a bit and then went back to the
script and shot that again when we were all kind of loosened up and threw
in bits of improvisation with bits of script. […] We did this improvisation
when we were actually filming. (Hagerty 2011)
Families
Agnostidae (p. 244).
Shumardiidae (p. 245).
Trinucleidae (p. 245).
Harpedidae (p. 245).
Paradoxidae (p. 246).
Conocephalidae = Conocoryphidae (p. 247).
Olenidae (p. 247).
Calymenidae (p. 247).
Asaphidae (p. 249).
Bronteidae (p. 249).
Phacopidae (p. 249).
Cheiruridae (p. 250).
Proëtidae (p. 251).
Encrinuridae (p. 251).
Acidaspidae (p. 251).
Lichadidae (p. 252).
ARACHNIDA (p. 255).
Orders.
Families.
Decolopodidae (p. 531).
Colossendeidae = Pasithoidae (p. 532).
Eurycididae = Ascorhynchidae (p. 533).
Ammotheidae (p. 534).
Rhynchothoracidae (p. 535)
Nymphonidae (p. 536).
Pallenidae (p. 537).
Phoxichilidiidae (p. 538).
Phoxichilidae (p. 539).
Pycnogonidae (p. 539).
CRUSTACEA
CHAPTER II
BY
The Crustacea are almost exclusively aquatic animals, and they play a
part in the waters of the world closely parallel to that which insects play
on land. The majority are free-living, and gain their sustenance either as
vegetable-feeders or by preying upon other animals, but a great number
are scavengers, picking clean the carcasses and refuse that litter the
ocean, just as maggots and other insects rid the land of its dead cumber.
Similar to insects also is the great abundance of individuals which
represent many of the species, especially in the colder seas, and the
naturalist in the Arctic or Antarctic oceans has learnt to hang the
carcasses of bears and seals over the side of the boat for a few days in
order to have them picked absolutely clean by shoals of small Amphipods.
It is said that these creatures, when crowded sufficiently, will even attack
living fishes, and by sheer press of numbers impede their escape and
devour them alive. Equally surprising are the shoals of minute Copepods
which may discolour the ocean for many miles, an appearance well known
to fishermen, who take profitable toll of the fishes that follow in their
wake. Despite this massing together we look in vain for any elaborate
social economy, or for the development of complex instincts among
Crustacea, such as excite our admiration in many insects, and though
many a crab or lobster is sufficiently uncanny in appearance to suggest
unearthly wisdom, he keeps his intelligence rigidly to himself, encased in
the impenetrable reserve of his armour and vindicated by the most
powerful of pincers. It is chiefly in the variety of structure and in the
multifarious phases of life-history that the interest of the Crustacea lies.
Before entering into an examination of these matters, it will be well to
take a general survey of Crustacean organisation, to consider the plan on
which these animals are built, and the probable relation of this plan to
others met with in the animal kingdom.
The Crustacea, to begin with, are a Class of the enormous Phylum
Arthropoda, animals with metamerically segmented bodies and usually
with externally jointed limbs. Their bodies are thus composed of a series
of repeated segments, which are on the whole similar to one another,
though particular segments may be differentiated in various respects for
the performance of different functions. This segmentation is apparent
externally, the surface of a Crustacean being divided typically into a
number of hard chitinous rings, some of which may be fused rigidly
together, as in the carapace of the crabs, or else articulated loosely.
Each segment bears typically a pair of jointed limbs, and though they
vary greatly in accordance with the special functions for which they are
employed, and may even be absent from certain segments, they may yet
be reduced to a common plan and were, no doubt, originally present on
all the segments.
Passing from the exterior to the interior of the body we find, generally
speaking, that the chief system of organs which exhibits a similar
repetition, or metameric segmentation, is the nervous system. This
system is composed ideally of a nervous ganglion situated in each
segment and giving off peripheral nerves, the several ganglia being
connected together by a longitudinal cord. This ideal arrangement,
though apparent during the embryonic development, becomes obscured
to some extent in the adult owing to the concentration or fusion of ganglia
in various parts of the body. The other internal organs do not show any
clear signs of segmentation, either in the embryo or in the adult; the
alimentary canal and its various diverticula lie in an unsegmented body-
cavity, and are bathed in the blood which courses through a system of
narrow canals and irregular spaces which surround all the organs of the
body. A single pair, or at most two pairs of kidneys are present.
The type of segmentation exhibited by the Crustacea is thus of a limited
character, concerning merely the external skin with its appendages, and
the nervous system, and not touching any of the other internal organs.[1]
In this respect the Crustacea agree with all the other Arthropods, in the
adults of which the segmentation is confined to the exterior and to the
nervous system, and does not extend to the body-cavity and its contained
organs; and for the same reason they differ essentially from all other
metamerically segmented animals, e.g. Annelids, in which the
segmentation not only affects the exterior and the nervous system, but
especially applies to the body-cavity, the musculature, the renal, and often
the generative organs. The Crustacea also resemble the other Arthropoda
in the fact that the body-cavity contains blood, and is therefore a
“haemocoel,” while in the Annelids and Vertebrates the segmented body-
cavity is distinct from the vascular system, and constitutes a true
“coelom.” To this important distinction, and to its especial application to
the Crustacea, we will return, but first we may consider more narrowly
the segmentation of the Crustacea and its main types of variation
within the group. In order to determine the number of segments which
compose any particular Crustacean we have clearly two criteria: first, the
rings or somites of which the body is composed, and to each of which a
pair of limbs must be originally ascribed; and, second, the nervous
ganglia.
Around and behind the region of the mouth there is very little difficulty
in determining the segments of the body, if we allow embryology to assist
anatomy, but in front of the mouth the matter is not so easy.
In the Crustacea the moot point is whether we consider the paired eyes
and first pair of antennae as true appendages belonging to two true
segments, or whether they are structures sui generis, not homologous to
the other limbs. With regard to the first antennae we are probably safe in
assigning them to a true body-segment, since in some of the
Entomostraca, e.g. Apus, the nerves which supply them spring, not from
the brain as in more highly specialised forms, but from the commissures
which pass round the oesophagus to connect the dorsally lying brain to
the ventral nerve-cord. The paired eyes are always innervated from the
brain, but the brain, or at least part of it, is very probably formed of
paired trunk-ganglia which have fused into a common cerebral mass; and
the fact that under certain circumstances the stalked eye of Decapods
when excised with its peripheral ganglion[2] can regenerate in the form of
an antenna, is perhaps evidence that the lateral eyes are borne on what
were once a pair of true appendages.
Now, with regard to the segmentation of the body, the Crustacea fall
into three categories: the Entomostraca, in which the number of segments
is indefinite; the Malacostraca, in which we may count nineteen
segments, exclusive of the terminal piece or telson and omitting the
lateral eyes; and the Leptostraca, including the single recent genus
Nebalia, in which the segmentation of head and thorax agrees exactly
with that of the Malacostraca, but in the abdomen there are two
additional segments.
It has been usually held that the indefinite number of segments
characteristic of the Entomostraca, and especially the indefinitely large
number of segments characteristic of such Phyllopods as Apus, preserves
the ancestral condition from which the definite number found in the
Malacostraca has been derived; but recently it has been clearly pointed
out by Professor Carpenter[3] that the number of segments found in the
Malacostraca and Leptostraca corresponds with extraordinary exactitude
to the number determined as typical in all the other orders of Arthropoda.
This remarkable correspondence (it can hardly be coincidence) seems to
point to a common Arthropodan plan of segmentation, lying at the very
root of the phyletic tree; and if this is so, we are forced to the conclusion
that the Malacostraca have retained the primitive type of segmentation in
far greater perfection than the Entomostraca, in some of which many
segments have been added, e.g. Phyllopoda, while in others segments
have been suppressed, e.g. Cladocera, Ostracoda. It may be objected to
this view of the primitive condition of segmentation in the Crustacea that
the Trilobites, which for various reasons are regarded as related to the
ancestral Crustaceans, exhibit an indefinite and often very high number
of segments; but, as Professor Carpenter has pointed out, the oldest and
most primitive of Trilobites, such as Olenellus, possessed few segments
which increase as we pass from Cambrian to Carboniferous genera.
The following table shows the segmentation of the body in the
Malacostraca, as compared with that of Limulus (cf. p. 263), Insecta, the
primitive Myriapod Scolopendrella, and Peripatus. It will be seen that the
correspondence, though not exact, is very close, especially in the first four
columns, the number of segments in Peripatus being very variable in the
different species.
Table showing the Segmentation of various Arthropods
Malacostraca. Limulus. Insecta. Myriapoda. Peripatus.
(Scolopendrella).
1 Eyes Median Eyes
eyes
2 1st antennae Rostrum Antennae Feelers Feelers
3 2nd antennae Chelicerae Intercalary
segment
4 Mandibles Pedipalpi Mandibles Mandibles Mandibles
5 1st maxillae 1st walking Maxillulae Maxillulae 1st jaw-claw
legs
6 2nd maxillae 2nd „ „ 1st 1st maxillae 2nd jaw-
maxillae claw
7 1st maxillipede 3rd „ „ 2nd 2nd maxillae 1st leg
maxillae
8 2nd maxillipede 4th „ „ 1st leg 1st leg 2nd „
9 3rd maxillipede Chilaria 2nd „ 2nd „ 3rd „
10 1st ambulatory Genital 3rd „ 3rd „ 4th „
operculum
11 2nd „ 1st gill- 1st 4th „ 5th „
book abdominal
12 3rd „ 2nd „ 2nd „ 6th „ 6th „
13 4th „ 3rd „ 3rd „ 6th „ 7th „
14 5th „ 4th „ 4th „ 7th „ 8th „
15 1st abdominal 5th „ 5th „ 8th „ 9th „
16 2nd „ No 6th „ 9th „ 10th „
appendages
17 3rd „ „ 7th „ 10th „ 11th „
18 4th „ „ 8th „ 11th „ 12th „
19 5th „ „ 9th „ 12th „ 13th „
20 6th „ „ 10th „ Reduced limbs 14th „
21 [4] „ Cercopods [5]
The essential fact that the two types of limb are built on the same plan
may be considered as established; but it may be urged that the biramous
type represents this common plan more nearly than the foliaceous. It is,
at any rate, certain that in the maxillipedes of the Decapoda we witness
the conversion of the biramous type into the foliaceous by the expansion
of the basal joints concomitantly with the assumption by the maxillipedes
of masticatory functions. Thus in the Decapoda the first maxillipede is
decidedly foliaceous owing to the expanded “gnathobases” (Fig. 1, A, bp,
cxp), and the second maxillipedes are flattened, with their basal joints
somewhat expanded and furnished with biting hairs; but in the
“Schizopoda” (e.g. Mysis) the first maxillipede is a typical biramous limb,
though the expanded gnathobases in some forms are beginning to project
(Fig. 1, E), while the limb following, which corresponds to the second
maxillipede of Decapods, is simply a biramous swimming leg. Besides this
obvious conversion of a biramous into a foliaceous limb, further evidence
of the fundamental character of the biramous type is found, first, in its
invariable occurrence in the Nauplius stage, which does not necessarily
mean that the ancestors of the Crustacea possessed this type of limb in
the adult, but which does imply that this type of limb was possessed at
some period of life by the common ancestral Crustacean; and, second, the
limbs of the Trilobita, a group which probably stands near the origin of
the Crustacea, have been shown by Beecher to conform to the biramous
type (Fig. 1, H). Furthermore, the thoracic limbs of Nebalia, an animal
which combines many of the characteristics of Entomostraca and
Malacostraca, and is therefore considered as a primitive type, despite
their flattened character, are really built upon a biramous plan (Fig. 1, G).
In conclusion, we may point out that this view of the Crustacean limb,
as essentially a biramous structure, agrees with the conclusion derived
from our consideration of the segmentation of the body, and points less to
the Branchiopoda as primitive Crustacea and more to some generalised
Malacostracan type.
So far we have shortly dealt with those systems of organs which are
clearly affected by the metameric segmentation of the body; we must now
expose the condition of the body-cavity to a similar scrutiny. If we
remove the external integument of a Crustacean, we find that the internal
organs do not lie in a spacious and discrete body-cavity, as is the case in
the Annelids and Vertebrates, but that they are packed together in an
irregular system of spaces (“haemocoel”) in communication with the
vascular system and containing blood. In the Entomostraca and smaller
forms generally, a definite vascular system hardly exists, though a central
heart and artery may serve to propel the blood through the irregular
lacunae of the body-cavity; but in the larger Malacostraca a complicated
system of arteries may be present which pour the blood into fairly
definitely arranged spaces surrounding the chief organs. These spaces
return the blood to the pericardium, and so to the heart again through the
apertures or ostia which pierce its walls.
This condition of the body-cavity or haemocoel is reproduced in the
adults of all Arthropods, but in some of them by following the
development we can trace the steps by which the true coelom is replaced
by the haemocoel. In the embryos of all Arthropods except the Crustacea,
a true closed metamerically segmented coelom is formed as a split in the
mesodermal embryonic layer of cells, distinct from the vascular system.
During the course of development the segmented coelomic spaces and
their walls give rise to the reproductive organs and to certain renal organs
in Peripatus, Myriapoda, and Arachnida (nephridia and coxal glands),
but the general body-cavity is formed as an extension of the vascular
system, which is laid down outside the coelom by a canaliculisation of the
extra-coelomic mesoderm. In the embryos of the Crustacea, however,
there is never at any time a closed segmented coelom, and in this respect
the Crustacea differ from all other Arthropods. The only clear instance in
which metamerically repeated mesodermal cavities have been seen in the
embryo Crustacean is that of Astacus; here Reichenbach[7] states that in
the abdomen segmental cavities are formed which subsequently break
down; but even in this instance no connexion has been shown to subsist
between these embryonic cavities and the reproductive and excretory
organs of the adult.
Since the connexion between the coelom and the excretory organs is
always a very close one throughout the animal kingdom, interest naturally
centres upon the renal organs in Crustacea, and it has been suggested
that these organs in Crustacea represent the sole remains, with the
possible exception of the gonads, of the coelom. Since, at any rate, a part
of the kidneys appears to be developed as a closed sac in the mesoderm,
and since they possess a possible segmental value, this suggestion is
plausible; but, on the other hand, since there are never more than two
pairs of kidneys, and since they are totally unconnected with the gonads
or with any other indication of a segmented coelom, the suggestion
remains purely hypothetical.
The renal organs of the Crustacea, excluding the Malpighian tubes
present in some Amphipods which open into the alimentary canal, and
resemble the Malpighian tubes of Insects, consist of two pairs—the
antennary gland, opening at the base of the second antenna, and the
maxillary gland, opening on the second maxilla. These two pairs of glands
rarely subsist together in the adult condition, though this is said to be the
case in Nebalia and possibly Mysis; the antennary glands are
characteristic of adult Malacostraca[8] and the larvae of the Entomostraca,
while the maxillary glands (“shell-glands”) are present in adult
Entomostraca and larval Malacostraca, that is to say, the one pair
replaces the other in the two great subdivisions of the Crustacea. The
shell-gland of the Entomostraca is a simple structure consisting of a
coiled tube opening to the exterior on the external branch of the second
maxilla, and ending blindly in a dilated vesicle, the end-sac. The
antennary gland of the Malacostraca is usually more complicated: these
complications have been studied especially by Weldon,[9] Allen, and
Marchal[10] in the Decapoda. In a number of forms we have a tube opening
to the exterior at the base of the second antenna, and expanding within to
form a spacious bladder into which the coiled tubular part of the kidney
opens, while at the extremity of this coiled portion is the vesicle called the
end-sac. This arrangement may be modified; thus in Palaemon Weldon
described the two glands as fusing together above and below the
oesophagus, the dorsal commissure expanding into a huge sac stretching
dorsally down the length of the body. This closed sac with excretory
functions thus comes to resemble a coelomic cavity, and the view that it is
really coelomic has indeed been upheld.
A modified form of this view is that of Vejdovský, who describes a
funnel-apparatus leading from the coiled tube into the end-sac of the
antennary gland of Amphipods; he regards the end-sac alone as
representing the coelom, while the funnel and coiled tube represent the
kidney opening into it.
Not very much is known of the development of these various structures.
Some authors have considered that both antennary and maxillary glands
are developed in the embryo from ectodermal inpushings, but the more
recent observations of Waite[11] on Homarus americanus indicate that the
antennary gland at any rate is a composite structure, formed by an
ectodermal ingrowth which meets a mesodermal strand, and from the
latter are produced the end-sac and perhaps the tubular excretory
portions of the gland with their derivatives.
With regard to the possible metameric repetition of the renal organs, it
is of interest to note that by feeding Mysis and Nebalia on carmine,
excretory glands of a simple character were observed by Metschnikoff
situated at the bases of the thoracic limbs.
The alimentary canal of the Crustacea is a straight tube composed of
three parts—a mid-gut derived from the endoderm of the embryo, and a
fore- and hind-gut formed by ectodermal invaginations in the embryo
which push into and fuse with the endodermal canal. The regions of the
fore- and hind-gut can be recognised in the adult by the fact of their being
lined with the chitinous investment which is continued over the external
surface of the body forming the hard exoskeleton, while the mid-gut is
naked. The chitinous lining of fore- and hind-gut is shed whenever the
animal moults. In the Malacostraca, in which a complicated “gastric mill”
may be present, the chitinous lining of this part of the gut is thrown into
ridges bearing teeth, and this stomach in the crabs and lobsters reaches a
high degree of complication and materially assists the mastication of the
food. The gut is furnished with a number of secretory and metabolic
glands; the so-called liver, which is probably a hepatopancreas, opening
into the anterior end of the mid-gut, is directed forwards in most
Entomostraca and backwards in the Malacostraca, in the Decapoda
developing into a complicated branching organ which fills a large part of
the thorax. In the Decapoda peculiar vermiform caeca of doubtful
function are present, a pair of which open into the gut anteriorly where
fore-passes into mid-gut, and a single asymmetrically placed caecum
opens posteriorly into the alimentary tract where mid- passes into hind-
gut.
The disposition of these caeca, marking as they do the morphological
position of fore-, mid-, and hind-gut, is of peculiar interest owing to the
variations exhibited. From some unpublished drawings of Mr. E. H.
Schuster, which he kindly lent me, it appears that in certain Decapods,
e.g. Callianassa subterranea, the length of the mid-gut between the
anterior and posterior caeca is very long; in Carcinus maenas it is
considerable; in Maia squinado it is greatly reduced, the caeca being
closely approximated; while in Galathea strigosa the caeca are greatly
reduced, and the mid-gut as a separate entity has almost disappeared.
The relation of these variations to the habits of the different crabs and to
their modes of development is unknown.
The reproductive organs usually make their appearance as a small
paired group of mesodermal cells in the thorax comparatively late in life;
and neither in their early development nor in the adult condition do they
show any clear signs of segmentation or any connexion with a coelomic
cavity. The sexes are usually separate, but hermaphroditism occurs
sporadically in many forms, and as a normal condition in some parasitic
groups (see pp. 105–107). The adult gonads are generally simple paired
tubes, from the walls of which the germ-cells are produced, and as these
grow and come to maturity they fill up the cavities of the tubes; special
nutrient cells are rarely differentiated, though in some cases (e.g.
Cladocera) a few ova nourish themselves by devouring their sister-cells
(see p. 44). The oviducts and vasa deferentia are formed as simple
outgrowths from the gonadial tubes, which acquire an opening to the
exterior; they are usually poorly supplied with accessory glands, the
epithelium of the canals often supplying albuminous secretions for
cementing the eggs together, while the lining of the vasa deferentia may
be instrumental in the formation of spermatophores for transferring large
packets of spermatozoa to the female. In the vast majority of Crustacea
copulation takes place, the male passing spermatophores or free
spermatozoa into special receptacles (spermathecae), or into the oviducts
of the female. The spermatophores are hollow chitinous structures in