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1
Oxford University Press is a department of the University of Oxford. It furthers
the University’s objective of excellence in research, scholarship, and education
by publishing worldwide. Oxford is a registered trade mark of Oxford University
Press in the UK and certain other countries.
This publication is made possible in part by the Dragan Plamenac Endowment of the American
Musicological Society, funded in part by the National Endowment for the Humanities and the
Andrew W. Mellon Foundation.
9 8 7 6 5 4 3 2 1
Printed by Sheridan Books, Inc., United States of America
v
For Eric
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vi
Contents
Acknowledgments ix
2 Reputational Entrepreneurs 29
vii
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ix
Acknowledgments
This book would never have been written without the generous assistance
I received from numerous institutions and individuals. My research was made
possible through various departments at the University of Texas at Austin. Two
sabbaticals from the Faculty Development Program proved invaluable, the first
for providing the initial impetus to embark on this project and the second for the
much-needed time to complete the manuscript. In addition, support from a uni-
versity research grant, two creative research grants from the College of Fine Arts,
and a Sarah and Ernest Butler Faculty Development Award from the Butler School
of Music funded research trips to libraries and archives, without which this type
of project would have been impossible. Throughout my travels, I benefited greatly
from the expertise of librarians and archivists, who helped me access the materi-
als I needed and shared their knowledge of the collections. I thank all those who
assisted me at the Archives de Paris, the Médiathèque Musicale Mahler, the music
department of the Bibliothèque nationale de France, and the Fonds ancien of the
Bibliothèque municipale de Lyon. Closer to home, I received invaluable assistance
from the Harry Ransom Center at the University of Texas at Austin, the New York
Public Library, and the Frick Art Reference Library at the Frick Collection, where
my dear friend and archivist, Julie Ludwig, helped me with my request.
My project has benefited immensely from friends and colleagues who offered
crucial feedback at the various stages of this book’s preparation, from grant
ix
x
x Acknowledgments
proposals and conference presentations to article and chapter drafts. My sincere
thanks go to Denis Herlin, Roy Howat, Barbara Kelly, Tamara Levitz, Christopher
Moore, Jann Pasler, Marie Rolf, Philip Rupprecht, and John Turci-Escobar for
their time, support, and astute commentary. Once again, I am extremely grate-
ful to my editor, Suzanne Ryan, for her expertise and enthusiasm as she ushered
my book through the publication process—I am delighted to be working with her
again. Diane Gervais, director of the Alliance Française d’Austin, was indispen-
sable for her expertise in matters of translation and for helping me to decipher
illegible handwriting on the various documents I brought to her. Finally, I thank
Ryan Beavers, Scott Schumann, and Christopher Prosser for their meticulous work
in preparing the music examples.
My colleagues at the Butler School of Music provided me with moral support
and humorous respites throughout the entire process of writing this book. In par-
ticular, I thank John Turci-Escobar and the stalwarts of happy hour, Jim Buhler,
Charles Carson, Eric Drott, and Hannah Lewis. Finally, to my husband and col-
league, Eric Drott, once again I am indebted for more than I can say. It was much
more fun researching and writing this book with him in my life. His support, en-
couragement, and advice have helped to shape my project from the initial idea to
the final manuscript. Debussy’s Legacy is dedicated to him.
xi
1
The Construction of Reputation and the Case of Debussy
On 17 June 1932, a monument to Debussy was unveiled near his home in the
sixteenth arrondissement on the outskirts of Paris. The creation of the sculptors
Jan and Joël Martel, this tribute to the composer comprised a large stone triptych
with two statues framing a central panel, all of which was mirrored in a reflecting
pool that extended to the front of the monument (see Figure 5.6 in Chapter 5).
Commentators were immediately drawn to the intricate design of the central
panel, entitled La grande tapisserie de pierre (the large stone tapestry). Louis Laloy,
for example, noted how its “bas-reliefs can be deciphered like hieroglyphics” and
he went on to identify “Saint Sébastien in his armor, Mélisande’s hair, the Faun
with his slanting flute, the waves of the sea engulfing the cathedral” (Figure 1.1).1
In his description, Gabriel Astruc provided even more detail, adding Pelléas be-
neath the evocation of Mélisande and her flowing hair, an enamored nymph below
the Faun, and angels next to the figure of Saint Sébastien, who was tied to a laurel
tree, an arrow piercing his heart. Astruc also noted the waves of La Mer, a mer-
maid representing “Sirènes” (Nocturnes), the pagoda from Estampes, the fish of
“Poissons d’or” (Images), and the toy figures of La Boîte à joujoux.2
In addition to describing the details on the face of the monument, several writers
commented on the numerous features that adorned the back of the central panel.
There was a quotation from the composer, writing as Monsieur Croche: “Search for
discipline within freedom. Listen to no-one’s advice except that of the wind that
1
2
Figure 1.1. Jan and Joël Martel, “The large stone tapestry,” Monument à Debussy (1932). By
permission of the Harry Ransom Center, University of Texas at Austin.
3
Figure 1.2. Jan and Joël Martel, “The symbolic concert,” Monument à Debussy (1932).
Photograph by Debretagne. By permission of the Harry Ransom Center, University of Texas at
Austin.
5
Once an artist dies, his or her reputation comes to rest irrevocably in other
hands. Only a small minority will have achieved the renown and gained a
following sufficient to carry them through the ages. The remembrance of
most . . . is highly dependent on survivors with an emotional and/or financial
stake in the perpetuation of their reputation.8
In Debussy’s case, it could be argued that he possessed both the renown and a
following that should have guaranteed a smooth transition from a living to a post-
humous reputation. But as documented throughout this book, these were not suf-
ficient in themselves. Since most studies of Debussy understandably end with his
death, few scholars take note of the vitriolic backlash against the composer in the
years after the First World War and the decade-long campaign that fought to keep
his musical aesthetic relevant in an era when it was characterized as outmoded,
decadent, and even harmful for the future of French music. To overturn such per-
ceptions entailed the actions of numerous individuals who devoted the time and
effort necessary to combat the downturn in the composer’s posthumous fortunes.
Howard S. Becker elaborates further on this collective process, stating that
“lasting [is] a crucial criterion of artistic greatness, that is to say, of reputation”
and that “artistic work lasts when it has an organizational basis that preserves
and protects it.”9 Becker emphasizes the need for a “network of cooperating
6
6 Debussy’s Legacy
people” whose activities are essential not only during the artist’s lifetime, but
also, even more so, after the artist’s death.10 Identifying Debussy’s network is a
straightforward matter, since in the latter part of his life it was well established
and efficient. It included Jacques Durand, who had an exclusive contract to pub-
lish and distribute the composer’s works from 1906 on.11 When pressed for time
and facing a fixed deadline, Debussy or Durand looked to colleagues for support in
the final stages of composition, recruiting assistance for tasks such as correcting
proofs, creating piano reductions, and completing orchestrations. The compos-
ers André Caplet and, on one occasion, Charles Koechlin performed such tasks.12
Once completed, the scores made their way to the impresarios, conductors, and
performers responsible for rehearsing and directing the premiere. Through their
professional standing, Debussy gained access to orchestras, ensembles, venues,
concert subscription series, and publicity, all of which ensured that tickets for the
event would be sold. At this point, critics sympathetic to the composer entered
the network, writing both to promote the work in advance of its performance
and to shape its reception afterward. Thus, Debussy worked at the “center of a
network of cooperating people” that aided him in the completion, distribution,
performance, and reception of his compositions, “all of whose work [was] essen-
tial to the final outcome.”13
After the composer’s death in 1918, this same network continued to operate,
the members of his cooperative circle simply retaining the roles and functions
they had performed during his lifetime. Both their sustained activity and their
ability to marshal a wide range of professional resources ensured that Debussy’s
music remained a permanent fixture on concert programs, facilitating high-profile
events featuring his music, restaging his theatrical works, and introducing nu-
merous unpublished compositions to concert audiences (discussed further below).
An important condition of this productivity, Bruno Latour asserts, is “the multi-
plicity of objects any course of action mobilizes along its trail.” For Latour, “objects
too have agency” and “any thing that does modify a state of affairs by making a dif-
ference is an actor.”14 From this perspective, the work of the members of Debussy’s
cooperative network depended not only on their capacity to mobilize the varied
resources at their disposal, but also on the generation of new entities such efforts
left in their wake. Whether it was the orchestrations, scores, concert programs,
and articles outlined previously or the plaques, monuments, and festivals dis-
cussed in Chapter 5, these artifacts made a difference, the steady accumulation of
objects large and small serving to anchor Debussy’s memory in the postwar years.
In this way, a concept as illusive and capricious as reputation—which resides solely
in the perceptions and opinions of others—was rendered visible, accrued a mate-
rial presence, and left a trace. Taken together, the ventures of this cooperative
7
8 Debussy’s Legacy
shaped how Debussy’s music was received after his death and how its aesthetic
worth was consolidated.
Chapter 1 considers some of the general mechanisms by which artistic figures
are consecrated and weighs their relative contribution to the construction of
Debussy’s reputation. Drawing on Gladys Engel Lang and Kurt Lang’s analysis of
the survival of reputation in the fine arts, four areas emerge that would seem to be
particularly relevant to the case of Debussy: (1) the initiatives undertaken by the
composer to establish his own legacy; (2) the posthumous reception of the corpus
of works left behind; (3) the actions of heirs and family members on behalf of the
deceased; and (4) the efforts of the composer’s close friends and collaborators.
Examining each of these categories in turn, however, suggests that the first two
prove to be the most unreliable for securing the composer’s early legacy. As Tia
DeNora observes, “a reputation that rests on what an individual says and does to
maintain it is perpetually vulnerable, however culturally powerful that individual
may be.”15 Such was the case for Debussy and, consequently, the two remaining
groups—heirs and family members on the one hand, friends and colleagues on the
other—played a significant and often antagonistic role in shaping the composer’s
posthumous fortunes.
There is no better evidence of Debussy’s preoccupation with his legacy than the
works that emerged during the last years of his life. Diagnosed with cancer in
1909, he was increasingly aware that his final years were approaching and this,
in combination with the heightened nationalism of the First World War, forced a
new level of introspection. In the last four years of his life, he engaged with mu-
sical traditions he had previously shunned and self-consciously worked to define
himself as a “musicien français” by connecting his works to key figures in France’s
artistic patrimony. The compositions that emerged in these years exhibited cer-
tain characteristics that suggested that Debussy was acutely aware of the issues
involved in reputation building. First, his choice of genre was significant in that
he decided to enter into and leave his imprint on some of the more densely popu-
lated areas of the musical canon. In 1915, the instrumental sonata and piano etude
were new to Debussy and both represented a volte-face from his previous disdain
for the continued use of traditional forms in contemporary composition. Second,
their prefaces and dedications (as well as the composer’s correspondence) evinced
Debussy’s concern for the historical affiliations of his works and the specific gene-
alogy of the genre with which he wished to be associated. By linking the sonatas
9
All that, it will be now “the things before the war,” that is to say things will be
separated from us by a wall, a true wall… . It will be necessary to begin again
to live, on a new conception of life, which does not mean that all that was
made before the war will be forgotten, but that all things made after must be
otherwise.19
Roussel’s letter acknowledged the huge schism brought about by the First World
War and how these four years would henceforth be bracketed from consideration,
with musical life leading up to 1914—“the things before the war”—separated from
01
10 Debussy’s Legacy
1919 and “all things made after.” Darius Milhaud elaborates on this view with a
narrative that similarly bypassed the war years by jumping directly from 1913 to
1919, but in this instance incorporating the figure of Erik Satie as a link between
the two dates: “After the shock produced by the premiere of Le sacre du printemps,
Satie reaches his full potential and brings music back to simplicity, thus opening
the way for the young musicians who formed the postwar French school.”20 On
the one hand, Milhaud’s account is admittedly self-serving, as he characterizes
Le sacre as the musical forerunner to the postwar French school, of which he is a
member. On the other hand, there is a delicate dance to be performed with regard
to Satie, who provides the only connection to the recent French musical past. In
his account, Milhaud takes care to emphasize Satie’s activities prior to the war
and his leadership role after the war. With a single sentence, he touches lightly on
the wartime premiere of Parade (1917) before swiftly moving on to devote the re-
mainder and majority of the paragraph to the premiere of Socrate (1919).21
Debussy’s death in the final months of the war served only to consolidate this
bifurcated view of recent French music history. Alongside the historical watershed
marked by the end of the war, the death of the leader of the prewar musical avant-
garde came to symbolize the end of an era, punctuating the close of one period and
the commencement of another. Georges Auric commented on this symbolism when
he wrote, “The historical importance of Debussy is such that he marks, I believe,
both the end of a style and the beginning of a new one.”22 But acknowledging the
divide between prewar and postwar musical life was only one part of the agenda.
As evident from Roussel’s letter cited above, the other was to magnify the differ-
ences between them. Debussy’s death at the end of the war, therefore, not only
signaled the end of an era, but also provided a convenient foil for a new generation
of composers seeking to find their place in the Parisian musical milieu. As Auric put
it, “Debussy remains a valued musician despite the upheaval that appears to me
to disrupt and destroy his aesthetic.”23 Looking back at the immediate postwar pe-
riod from the vantage point of 1926, Roussel commented on “this natural reaction
against existing trends,” noting how “the recent cataclysm that we witnessed” [ . . .]
has “not been negligible in influencing the current orientation of the arts.”24 Ravel,
who had been a frequent target in the backlash against the prewar avant-garde,
described this musical reaction in much stronger language than Roussel, employing
terms of aggression, violence, destruction, and demolition:
There are the postwar “young people,” in other words, the restless, rash,
and somewhat aggressive adolescents, who had to begin afresh the work of
musical civilization on a completely topsy-turvy planet. Their task was dif-
ficult and ungrateful. They felt an instinctive need to break violently with
1
In emphasizing the rift and antagonisms between prewar and postwar musical
life, such narratives simply omitted the war years. As the above citations demon-
strate, Roussel, Milhaud, and Ravel all chose to contrast either 1913 or 1914 with
1919 and, in the process, consigned the intervening years to oblivion.
Surprisingly, the debussyists were equally complicit in bypassing the war years
in their accounts of the composer’s life and works. Rather than challenge the stark
pre-and postwar dichotomy of recent music history, the debussyists reinforced it
by performing a similar maneuver of their own. But whereas the next generation
of composers disregarded the war years to move forward, the composer’s support-
ers disregarded these same years to return to the past. Thus, in remembering the
composer, they chose to forget: the past they elevated was a highly selective and
edited portion of the composer’s entire career. Vuillermoz’s outline of the land-
marks of Debussy’s oeuvre is revealing in this respect, as he traversed from “the
Quartet to the Prélude à l’Après-Midi d’un Faune, from the Faune to the Nocturnes,
from the Nocturnes to Pelléas, from Pelléas to La Mer, and from La Mer to Le Martyre
de Saint Sébastien!”26 According to Vuillermoz’s summary, the masterworks worthy
of mention come to a halt in 1911, with Le Martyre de Saint Sébastien considered
the last example of the composer in his prime. Vuillermoz was not alone in his ex-
alted view of this work. In the same year, Laloy wrote, “The truth is that the music
of Le Martyre, which is more accomplished than that of Pelléas, is no less animated,
and radiates a celestial clarity which was foreign to the entirely human and earthly
Pelléas.”27 For Laloy, Le Martyre stood out not only in Debussy’s oeuvre, but also
in the history of theatrical works in general. Alongside Stravinsky’s Rossignol, he
asserted that “only the revelations of Parsifal, Boris Godunov, Pelléas et Mélisande
and Saint Sébastien [were] comparable.”28
If Debussy’s detractors tended to treat his career as having all but ended by 1913
or 1914, his supporters selected an even earlier date, choosing the 1911 premiere of
Le Martyre as the culmination of his musical oeuvre. Le Martyre was fashioned into
a swansong, its elevation occurring at the expense of the remaining seven years
of Debussy’s compositional career.29 But there were other reasons why Le Martyre
assumed such a prominent position in the minds of many debussyists. Like Pelléas
before, it was a theatrical work in which they played a decisive role. Just as their
attendance and activism were largely responsible for keeping Pelléas et Mélisande
21
12 Debussy’s Legacy
in the theater after its premiere, their practical contributions to Le Martyre were
largely responsible for delivering it to the theater in the first place. Accounts of
the rehearsal period recall how the composer’s “cooperative network” shifted into
high gear to prepare the work for its first performance: Astruc was the impre-
sario responsible for managing the enterprise and promoter of La grande Saison
de Paris in which the work was to appear; Debussy, working on a tight schedule of
four months, delivered the manuscript in fragments to Durand, who engraved and
published the parts as soon as they appeared; Caplet helped Debussy to orches-
trate the score, rehearsed the orchestra, and conducted the premiere; Vuillermoz
and Inghelbrecht directed the chorus during rehearsals, and Inghelbrecht con-
ducted the chorus for the premiere.30 Given this level of participation, the debussy-
ists’ championing of Le Martyre went beyond aesthetic preferences to elevate a
work in which their contributions were essential. Thus, in remembering the com-
poser, they were remembering themselves: their truncated view of Debussy’s ca-
reer emphasized precisely those works in which they were the most active and
consequently their memories were the most vivid. Astruc’s reminiscences confirm
this, with his overview of Debussy’s career prioritizing less the works themselves
and more the controversies associated with them: “After the skirmishes of the pre-
mieres at Lamoureux and Colonne, after the great battle of Pelléas and the unique
combat of the Faune, came the affair of Le Martyre de Saint Sébastien.”31 Despite its
chronological precedence—the “unique combat of the Faune” referring to the 1912
Ballets Russes production of L’Après-midi d’un faune—Astruc placed Le Martyre at
the end of his account, its premiere representing the final point of combat and, by
implication, the final highpoint of the composer’s career.
For better and for worse, Debussy’s early legacy was shaped in response to this
“collective amnesia,” the selective forgetting and remembering generally empha-
sizing the years 1902 to 1911—that is, from the premiere of Pelléas et Mélisande to
the premiere of Le Martyre.32 The ramifications of this foreshortening of historical
perspective were twofold. First, it had the unfortunate consequence of emphasiz-
ing precisely those works by Debussy that were in the process of being devalued
by the next generation of composers and critics: lengthy theatrical works with
symbolist or mystical libretti and large orchestral compositions with picturesque
titles and programs. Indeed, by continually promoting Le Martyre, the debussyists
exhibited a certain lack of musical acuity, demonstrating how tone-deaf they were
to the concerns of postwar musical aesthetics. Second, the truncation of Debussy’s
creative career omitted from consideration the last years of the composer’s life,
the years that embodied a total reorientation of his creative endeavors and mani-
fested his self-conscious attempts to define his musical legacy. If the debussyists
had a habit of highlighting their own interactions with the composer, then his
31
14 Debussy’s Legacy
those works that were published during his lifetime. In reality, the boundaries of
his oeuvre were just as variable as was opinion on his aesthetic. As we have seen,
the number of his compositions was continually subject to contraction, with the
various curtailments of his creative career. At the same time, however, his oeuvre
was also subject to expansion, with the discovery, distribution, and performance
of previously unpublished manuscripts. Table 1.1 lists the posthumous premieres
in the first decade after Debussy’s death, demonstrating how his corpus of works
was far from stable, with ten “new” compositions appearing in as many years.34
Table 1.1 also illustrates the efficacy of Debussy’s organizational network,
which continued to produce works despite the absence of the composer. Debussy’s
former friends and colleagues maintained their roles as orchestrators, conductors,
and soloists to bring previously unpublished compositions to the concert stage.
But not all of these posthumous premieres were viewed positively. The members
of Debussy’s cooperative circle had reputations too, and the individual reputations
of those involved at each step in the process contributed to the overall acceptance
or rejection of the work. At one end of the spectrum, the posthumous premieres
of La Boîte à joujoux and Khamma caused little consternation, both because of the
state of the compositions and because of the accepted authority of the personnel
who made the first performance possible. The piano versions of La Boîte à joujoux
and Khamma had been completed by the composer and published by Durand dur-
ing his lifetime. Moreover, Debussy had started the orchestration of both works,
and their completion by Caplet and Koechlin, respectively, aroused no cause for
concern. It was widely known that Caplet had orchestrated for Debussy before
and, although many were unaware that Koechlin had performed this service for
Khamma, it was announced in the press that its orchestration was completed
during the composer’s lifetime, under his supervision, and with his complete ap-
proval.35 The authority of Durand to publish these works was never questioned;
neither was the appropriateness of Caplet and Gabriel Pierné to conduct their
first performances, both of whom had worked numerous times with the composer
before. With each composition, the reputations of publisher, orchestrator, and
conductor came together to ensure the legitimacy and propriety of the posthu-
mous premiere. At the other end of the spectrum, however, were the first perfor-
mances of 1928, which included three works from Debussy’s student years and
his final composition, the Ode à la France, incomplete at the time of his death.
The decision to publish and perform these compositions, the questionable state of
the manuscripts, and the choice of publisher, in addition to the qualifications of
the composer selected to “finish” these works and conduct the premiere, all com-
bined to cast doubt on the concert. The names of Choudens and Marius-François
51
Table 1.1.
16 Debussy’s Legacy
Gaillard were new to this enterprise and went well beyond the cooperative net-
work sanctioned by the composer.
The posthumous premieres of the 1920s represented a double-edged sword
with regard to Debussy’s legacy. On the one hand, they had a negative effect
in that they reminded the younger generation of composers and critics exactly
what it was they were trying to renounce, a state of affairs made ever more
apparent with less successful compositions from Debussy’s oeuvre. André
Schaeffner enunciated this view: “The first performance of unpublished works
by Debussy (Le Roi Lear, Lindaraja) offered only the employment of procedures
that Debussy had been the first to abuse.”36 On the other hand, the excitement
generated by a newly discovered composition was positive, keeping Debussy’s
name in newspapers, in music periodicals, and on concert programs. The
rounds of publicity that attended each “new” work were, in this respect, bene-
ficial for the perpetuation of his legacy. Furthermore, it was a clear indicator of
renown that any composition that carried his name was automatically consid-
ered worthy of interest. In his review of the 1928 posthumous premieres, for
example, Henry Prunières asserted, “nothing that Debussy composed should
leave us indifferent.” But he immediately followed this statement with an-
other: “I admit that the student works that were revealed to us contribute
little to his glory.”37
Prunières’s second statement was a common refrain among reviews of the post-
humous premieres and perhaps represented the most neutral comment a critic
could make. Vuillermoz uttered practically the same phrase concerning Le Roi Lear
and Lindaraja: “these exhumations, as well as the Fantaisie pour piano, [ . . .] will
add nothing to the glory of the author.” Maurice Boucher, reviewing the same com-
positions, presented a slight variation on this theme: “they add absolutely noth-
ing to the admiration that we have for Debussy.”38 Although seemingly innocuous
(and evidently part of the critical lexicon), this phrase tacitly acknowledged what
Becker refers to as the works “counted up in the balance of [a]reputation.” Becker
describes how “artists’ reputations are a sum of the values we assign to the works
they have produced. Each work that can definitely be attributed to [Debussy] adds
to or subtracts from the total on the basis of which we decide how great an art-
ist [Debussy] was.”39 Thus, the statements of Prunières, Vuillermoz, and Boucher
were perhaps more than merely neutral: that is, they refused to allow these works
to count in assessments of the composer’s reputation. In some instances, the post-
humous premieres were even judged to “subtract” from the balance of Debussy’s
reputation. By 1928, Vuillermoz feared that these endless “exhumations” had the
potential to damage the composer’s legacy: “The friends of Debussy are begin-
ning to get alarmed by this unexpected swelling of posthumous works, which one
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CHAPTER III
CRUSTACEA (CONTINUED): COPEPODA
BRANCH I. EUCOPEPODA.
Sub-Order 1. Gymnoplea.
The division between the front and hind part of the body falls
immediately in front of the genital openings and behind the fifth thoracic
feet. The latter in the male are modified into an asymmetrical copulatory
organ.
TRIBE I. AMPHASCANDRIA.
TRIBE II.
HETERARTHRANDRIA.
The first antennae of the male are asymmetrical, one, usually the
right, being used as a clasping organ.
The males of the Centropagidae, Candacidae and Pontellidae,
besides possessing the asymmetrically modified thoracic limbs of the
fifth pair also exhibit a modification of one of the first antennae,
which is generally thickened in the middle, and has a peculiar joint in
it, or geniculation, which enables it to be flexed and so used as a
clasping organ for holding the female.
Fam. 1.—Centropagidae.—These Copepods are very common
in the pelagic plankton, and some of the species vie with the
Calanidae in plumed ornaments, e.g. Augaptilus filigerus, figured by
Giesbrecht in his monograph. The use of these ornaments, which are
possessed by so many pelagic Copepods, is entirely obscure.[40]
Certain of the Centropagidae live in fresh water. Thus Diaptomus is
an exclusively fresh-water genus, and forms a most important
constituent of lake-plankton; various species of Heterocope occur in
the great continental lakes, and certain Eurytemora go up the
estuaries of rivers into brackish water.
An excellent work on the fresh-water Copepods of Germany has
been written by Schmeil,[41] who gives analytical tables for
distinguishing various genera and species. The three fresh-water
families are the Centropagidae, Cyclopidae, and Harpacticidae (see
p. 62). The Centropagidae may be sharply distinguished from the
other fresh-water families by the following characters:—The
cephalothorax is distinctly separated from the abdomen; the first
antennae are long and composed of 24–25 segments, in the male
only a single antenna (generally the right) being geniculated and
used as a clasping organ. The fifth pair of limbs are not rudimentary;
a heart is present, and only one egg-sac is found in the female. The
second antennae are distinctly biramous.
Diaptomus.—The furcal processes are short, at most three times as
long as broad; endopodite of the first swimming appendage 2–
jointed, endopodites of succeeding legs 3–jointed.
Heterocope.—The furcal processes are short, at most twice as long
as broad; endopodites of all swimming legs 1–jointed.
Eurytemora.—The furcal processes are long, at least three and a
half times as long as broad; the endopodite of the first pair of legs
1–jointed, those of the other pairs 2–jointed.
It has been known for a long time that some of the marine
Copepods are phosphorescent, and, indeed, owing to their numbers
in the plankton, contribute very largely to bring about that liquid
illumination which will always excite the admiration of seafarers. In
northern seas the chief phosphorescent Copepods belong to
Metridia, a genus of the Centropagidae; but in the Bay of Naples
Giesbrecht[42] states that the phosphorescent species are the
following Centropagids: Pleuromma abdominale and P. gracile,
Leuckartia flavicornis and Heterochaeta papilligera; Oncaea
conifera is also phosphorescent. It is often stated that Sapphirina (p.
69) is phosphorescent, but its wonderful iridescent blue colour is
purely due to interference
colours, and has nothing to do
with phosphorescence.
Giesbrecht has observed that the
phosphorescence is due to a
substance secreted in special
skin-glands, which is jerked into
the water, and on coming into
contact with it emits a
phosphorescent glow. This
substance can be dried up
completely in a desiccated
specimen and yet preserve its
phosphorescent properties, the
essential condition for the actual
emission of light being contact
with water. Similarly, specimens
preserved in glycerine for a long
period will phosphoresce when
compressed in distilled water.
From this last experiment
Giesbrecht concludes that the
phosphorescence can hardly be
due to an oxidation process, but
the nature of the chemical
reaction remains obscure.
Fam. 2. Candacidae.—This
family comprises the single genus
Candace, with numerous species
distributed in the plankton of all
seas. Some species, e.g. C.
pectinata, Brady, have a
practically world-wide
distribution, this species being
recorded from the Shetlands and
from the Philippines.
Fam. 3. Pontellidae.—This is
a larger family also comprising
widely distributed species found
Fig. 27.—Dorsal view of Anomalocera
pattersoni, ♂, × 20. (After Sars.)in the marine plankton.
Anomalocera pattersoni (Fig. 27)
is one of the commonest elements in the plankton of the North Sea.
Sub-Order 2. Podoplea.
The boundary between the fore and hind part of the body falls in
front of the fifth thoracic segment. The appendages of the fifth
thoracic pair in the male are never modified as copulatory organs.
TRIBE I. AMPHARTHRANDRIA.
The first antennae in the male differ greatly from those in the
female, being often geniculated and acting as prehensile organs.
Fams. 1–2. Cyclopidae and Harpacticidae, and other allied
families, are purely free-living forms; they are not usually pelagic in
habit, but prefer creeping among algae in the littoral zone or on the
sea-bottom, or especially in tidal pools. Some genera are,
nevertheless, pelagic; e.g. Oithona among Cyclopidae; Setella,
Clytemnestra, and Aegisthus among Harpacticidae.
The sketch (Fig. 28) of Euterpe acutifrons ♀ , a species widely
distributed in the Mediterranean and northern seas, exhibits the
structure of a typical Harpacticid, while Fig. 29 shows the form of the
first antenna in the male.
Several fresh-water representatives of these free-living families
occur. The genus Cyclops (Cyclopidae) is exclusively fresh-water,
while many Harpacticidae go up into brackish waters: for example
on the Norfolk Broads, Mr. Robert Gurney has taken Tachidius
brevicornis, Müller, and T. littoralis, Poppe; Ophiocamptus
brevipes, Sars; Mesochra lilljeborgi, Boeck; Laophonte littorale, T.
and A. Scott; L. mohammed, Blanchard and Richard; and
Dactylopus tisboides, Claus.
Schmeil[43] gives the following scheme for identifying the fresh-
water Cyclopidae and Harpacticidae (see diagnosis of Centropagidae
on p. 59):—
Fam. 1. Cyclopidae.—The
cephalothorax is clearly separated
from the abdomen. The first
antennae of the female when bent
back do not stretch beyond the
cephalothorax; in the male both
of them are clasping organs. The
second antennae are without an
exopodite. The fifth pair of limbs
are rudimentary, there is no
heart, and the female carries two
egg-sacs.
Cyclops.—Numerous species, split
up according to segmentation of
rudimentary fifth pair of legs,
number of joints in antennae, etc.
Fam. 2. Harpacticidae.—
The cephalothorax is not clearly
separated from the abdomen. The
first antennae are short in both
sexes, both being clasping organs
in the male. The second antennae
have a rudimentary exopodite.
The fifth pair of limbs are
rudimentary and plate-shaped; a
heart is absent, and the egg-sacs
Fig. 28.—Euterpe acutifrons, ♀ , × 70. of the female may be one or two
Abd.1, 1st abdominal segment; Th.5, in number.
5th thoracic segment. (After
Giesbrecht.) 1. Ophiocamptus (Moraria).—Body
worm-shaped; first antennae of
female 7–jointed, rostrum
forming a broad plate.
2. Body not worm-shaped; first antennae of female 8–jointed,
rostrum short and sharp.
(a) Endopodites of all thoracic limbs 3–jointed. The first
antennae in female distinctly bent after the second joint.
Nitocra.
(b) Endopodite of at least
the fourth limb 2–jointed;
first antennae in female
not bent. Canthocamptus.
3. Ectinosoma.—Body as in
2, but first antennae are very
short, and the maxillipede
does not carry a terminal
hooked seta as in 1 and 2.
Fam. 3. Peltiidae.[44]—This is
an interesting family, allied to the
Harpacticidae, and includes
species with flattened bodies
somewhat resembling Isopods,
and a similar habit of rolling
themselves up into balls. No
parasitic forms are known,
though Sunaristes paguri on the
French and Scottish coasts is said
to live commensally with hermit-
crabs.
The first antennae are short, similar in the two sexes, and are
never used by the male as clasping organs. This function may be
subserved by the second maxillae.
Fams. Oncaeidae, Corycaeidae, Lichomolgidae, Ergasilidae,
Bomolochidae, Chondracanthidae, Philichthyidae,
Nereicolidae, Hersiliidae, Caligidae, Lernaeidae,
Lernaeopodidae, Choniostomatidae.
The families Oncaeidae and Corycaeidae contain pelagic forms of
flattened shape and great swimming powers, but the structure of the
mouth-parts in the Corycaeidae points to a semi-parasitic habit.
Fam. 1. Oncaeidae.—This family, including the genera Oncaea,
Pachysoma, etc., does not possess the elaborate eyes of the next
family, nor is the sexual dimorphism so marked.
Fam. 2. Corycaeidae.—These are distinguished from the
Oncaeidae, not only by their greater beauty, but also by the
possession of very elaborate eyes, which are furnished with two
lenses, one at each end of a fairly long tube. The females of
Sapphirina are occasionally found in the branchial cavity of Salps,
and their alimentary canal never contains solid particles, but is filled
with a fluid substance perhaps derived by suction from their prey. S.
opalina may occur in large shoals, when the wonderful iridescent
blue colour of the males makes the water sparkle as it were with a
sort of diurnal phosphorescence. The animal, however, despite the
opinion of the older observers, is not truly phosphorescent. It may be
that the ornamental nature of some of the males is correlated with
the presence of the curious visual organs, which are on the whole
better developed in the females than in the males. As in so many
pelagic Copepods, the body and limbs may bear plumed setae of
great elaboration and beautiful colour, e.g. Copilia vitrea (Fig. 37).
We now pass on to the rest of the parasitic Copepods,[51] which
probably belong to the tribe Isokerandria, and we meet with the
same variety of degrees of parasitism as in the Ampharthrandria,
often leading to very similar results.
Fig. 37.—Copilia vitrea (Corycaeidae), ♀, × 20. (After Giesbrecht.)