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OUP CORRECTED PROOF – FINAL, 9/11/2016, SPi
published
1 Definite Descriptions
Paul Elbourne
2 Logic in Grammar
Polarity Free Choice, and Intervention
Gennaro Chierchia
3 Weak Island Semantics
Márta Abrusán
4 Reliability in Pragmatics
Eric McCready
5 Numerically Qualified Expressions
Christopher Cummins
6 Use-Conditional Meaning
Studies in Multidimensional Semantics
Daniel Gutzmann
7 Gradability in Natural Language
Logical and Grammatical Foundations
Heather Burnett
8 Subjectivity and Perspective in Truth-Theoretic Semantics
Peter Lasersohn
in preparation
Meaning over Time
The Foundations of Systematic Semantic Change
Ashwini Deo
Measurement and Modality
Daniel Lassiter
Plural Reference
Friederike Moltmann
The Semantics of Evidentials
Sarah E. Murray
A History of Formal Semantics
Barbara Partee
OUP CORRECTED PROOF – FINAL, 9/11/2016, SPi
Gradability in
Natural Language
Logical and Grammatical
Foundations
HEATHER BURNETT
1
OUP CORRECTED PROOF – FINAL, 9/11/2016, SPi
3
Great Clarendon Street, Oxford, ox2 6dp,
United Kingdom
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 in certain other countries
© Heather Burnett 2017
The moral rights of the author have been asserted
First Edition published in 2017
Impression: 1
All rights reserved. No part of this publication may be reproduced, stored in
a retrieval system, or transmitted, in any form or by any means, without the
prior permission in writing of Oxford University Press, or as expressly permitted
by law, by licence or under terms agreed with the appropriate reprographics
rights organization. Enquiries concerning reproduction outside the scope of the
above should be sent to the Rights Department, Oxford University Press, at the
address above
You must not circulate this work in any other form
and you must impose this same condition on any acquirer
Published in the United States of America by Oxford University press
198 Madison Avenue, New York, NY 10016, United States of America
British Library Cataloguing in Publication Data
Data available
Library of Congress Control Number: 2016936522
ISBN 978–0–19–872479–7 (hbk.)
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Printed in Great Britain by
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Links to third party websites are provided by Oxford in good faith and
for information only. Oxford disclaims any responsibility for the materials
contained in any third party website referenced in this work.
OUP CORRECTED PROOF – FINAL, 9/11/2016, SPi
To my parents
OUP CORRECTED PROOF – FINAL, 9/11/2016, SPi
OUP CORRECTED PROOF – FINAL, 9/11/2016, SPi
Contents
General preface x
Acknowledgments xi
List of abbreviations xiv
Introduction
.. Organization of the book
Vagueness and linguistic analysis
.. Introduction
.. Our Classical Semantic Theory
... Classical First order logic
... Extensions in linguistics
.. The phenomenon of vagueness
... Borderline cases
... Fuzzy boundaries
... The Sorites paradox
.. Tolerant, Classical, Strict
... Definition
... Account of the puzzling properties
.. Lasersohn’s () Pragmatic Halos
... Definition
... Comparison with Tolerant, Classical, Strict
.. Conclusion
Context sensitivity and vagueness patterns
.. Introduction
.. Adjectival context sensitivity patterns
.. Universal vs existential context sensitivity
.. Potential vagueness and adjectival vagueness patterns
... (A)symmetric vagueness
.. Conclusion
The Delineation Tolerant, Classical, Strict framework
.. Introduction
.. Language and classical semantics
OUP CORRECTED PROOF – FINAL, 9/11/2016, SPi
viii contents
contents ix
References
Index
OUP CORRECTED PROOF – FINAL, 9/11/2016, SPi
General preface
Acknowledgments
This work would never have been possible without all the help and
support that I have received from friends and colleagues during my
time as a doctoral student in the Department of Linguistics at UCLA
and as a Social Sciences and Humanities Research Council (SSHRC)
postdoctoral fellow in the Département de linguistique et de traduction
at l’Université de Montréal.
Although credit is due to many many people, I have to single out Paul
Égré for the enormous contributions that he has made to both the con-
tent of this monograph and my personal and professional development.
I thank him for so many things, including but not limited to: introducing
me to the exciting world of non-classical logics, showing me how to fill in
the CNRS application form, laughing at my jokes (even the ones making
fun of philosophers, Normaliens and French people), showing me how
to eat a hamburger French-style (i.e. with a knife and fork) and letting
me sleep in his kid’s room when I was homeless in New York.
More generally, I thank the members of the Institut Jean Nicod at the
Ecole normale supérieure in Paris (particularly members of the LIN-
GUAE group: Emmanuel Chemla, Vincent Homer, Philippe Schlenker,
Benjamin Spector, and Jérémy Zehr, as well as Claire Beyssade, Francis
Corblin, Alda Mari, David Nicolas, and François Récanati) for welcom-
ing me both as a student and as a visiting postdoctoral researcher. The
time that I have spent at Jean Nicod has been incredibly rewarding
(both academically and personally), and the influence of the ideas being
developed in this lab can be clearly seen in the major themes explored
in the book.
The first part of this book (chapters 2–5) is based on my 2012 disserta-
tion The Grammar of Tolerance: On vagueness, context-sensitivity and the
origin of scale structure, which was completed in the Linguistics depart-
ment at UCLA. My supervisors, Ed Keenan and Dominique Sportiche,
as well as Hilda Koopman, Jessica Rett, Yael Sharvit, and Ed Stabler
have all made innumerable vital contributions to this project, and I will
Credit for this last one is, of course, also due to Rachida, Amir, and, above all, to my
roommate Isaac.
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xii acknowledgments
acknowledgments xiii
List of abbreviations
list of abbreviations xv
SA scalar adjective
SC Strict Convexity
SP Shared Parts
SWO strict weak order
TA Total Axiom
TC Tolerant Convexity
TCS Tolerant, Classical, Strict
T-model Tolerant model
UD Upward Difference
VP verb phrase
wff well-formed formula
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1
Introduction
2 introduction
(2) For all x, y, if x is not tall and x and y’s heights differ by at most
1 cm, then y is also not tall.
Clearly the fact that both tall and not tall are tolerant creates a puzzle:
why do we not conclude that both the 1.9 m man and the 1.5 m man
are tall and not tall at the same time? Paradoxes of this type are known
as Sorites paradoxes, and they will be discussed in much greater detail
throughout the book.
Another adjective that shows a similar pattern is straight: In most
situations, adding a 1/10 mm bend to a stick is such an irrelevant change
that it will never be sufficient to make a stick that we call straight not
called straight. Thus, if we were to line up a set of sticks that differ by
1/10 mm bend from the perfectly straight ones to the really bendy ones,
then (3) seems true.
(3) For all x, y, if x is straight and x and y differ by a single 1/10 mm
bend, then y is also straight.
However, unlike tall, whose negation is also tolerant, even though
adding or subtracting a 1/10 mm bend is such a small change, the
corresponding statement with not straight is false: in particular (4) is
falsified by the case where we move from x that has a 1/10 mm bend (so
is not straight) to y that has absolutely no bends.
(4) False: For all x, y, if x is not straight and x and y differ by a single
1/10 mm bend, then y is also not straight.
In summary, on the one hand, adjectives like tall and straight are both
tolerant, but on the other, straight displays an asymmetry that tall
does not.
The second phenomenon that will be treated in this work is context
sensitivity. To be more specific, we will call a predicate P context sensitive
just in case, for some individual x, we can find a context in which P
applies to x, and we can find another context in which P does not apply
to x, without changing the properties of x and y. The adjectives tall and
straight both have this property: someone who can be considered tall
when we are considering jockeys might not be considered tall when we
The name of these puzzles comes from a puzzle attributed to Eubelides of Miletus known
introduction 3
are considering average men. Likewise, we saw above that an object with
a very small bend can be sometimes considered to be straight; however,
in a context in which very slight bends make a large difference to our
purposes, the very same object would not be considered straight.
This being said, tall and straight display a different pattern when it
comes to being context sensitive. For example, as discussed in Kennedy
(2007) and Syrett et al. (2010) (among others), adjectives like tall can
shift their criteria of application across contexts in a way that adjectives
like straight cannot. If I have two objects, one of which is (noticeably)
taller than the other, but neither are particularly tall, I can still use the
predicate tall to pick out the taller of the two.
(5) Pass me the tall one.
OK: even if neither/both is/are tall.
However, using straight in such a linguistic construction is only possible
if exactly one of the two is (very close) to perfectly straight.
(6) Pass me the straight one.
# if neither/both is/are straight.
The third phenomenon treated in this work is scalarity. Again, tall and
straight pattern alike on this dimension in that they can both appear in
the comparative and many other degree constructions (7).
(7) a. This stick is taller/straighter than that one.
b. This stick is very tall/straight.
However, once more, if we look at the full range of data concerning
gradability and scale structure, tall and straight show a different pattern:
for example, certain scalar modifiers like almost and completely are
natural with straight, but not with tall.
(8) a. ??John is almost/completely tall.
b. This stick is almost/completely straight.
The main goal of Chapters 2–5 is to develop an account of both the
similarities and differences between various subclasses of adjectives with
respect to each of these three phenomena (vagueness, context senstivity,
and scalarity). The principle subclasses that will be empirically distin-
guished are the following:
Consider, for example, the barrel of a rifle that must be perfectly straight for our shots
to be accurate.
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4 introduction
possible in some dialects of English (ex. This room is all empty ≈ This room is completely
OUP CORRECTED PROOF – FINAL, 7/11/2016, SPi
introduction 5
tou(te)s ‘all’, can combine with both adjectives like droit ‘straight’ and
definite plural DPs like les filles ‘the girls’ to create a parallel maximizing
interpretation.
(13) a. La rue est toute droite.
The road is all straight
‘The road is completely straight.’
b. Toutes les filles sont arrivées.
All the girls are arrived
‘All the girls arrived.’
Although examples such as (13) (and others to be discussed in Chapter
7) suggest that definite plural DPs and total adjectives have similar
scale structure properties, we will also see that context sensitivity/
vagueness/scale structure have slightly different manifestations with
DPs than with adjectives. In particular, I will argue that we see a different
typology of context sensitivity, vagueness, and scale structure patterns
in the DP domain than in the adjectival domain. Chapters 6 and 7 are
therefore devoted to capturing both the similarities (such as (13)) and
the proposed differences between adjectival and DP constituents within
a mereological extension (Simons, 1987; Hovda, 2008, among others) of
the DelTCS system (called M-DelTCS).
I will propose that the scales associated with DPs are derived from
statements about their context sensitivity and vagueness in the same
basic way as with adjectives. This is what creates the observed cross-
domain parallels. However, I will also propose that the different kinds
of ontological relations that characterize the domains into which DPs
and adjectives denote have important consequences for how the appli-
cation of these constituents can vary across comparison classes and how
they display the characteristic properties of vague language. In other
words, by virtue of the fact that DP constituents are interpreted into
domains that have mereological (i.e. part-structure) relations on them,
their context sensitivity and vagueness is constrained in a way that the
context sensitivity and vagueness of adjectival constituents is not. In
turn, by virtue of the logical structure of the M-DelTCS framework,
these differences in context sensitivity and vagueness will be translated
into differences in scale structure. Based on these results, I conclude that
empty). However, adjectival all is not fully productive in English in the way that its counter-
parts in the Romance languages (or even in German) are. Indeed, Moltmann () refers
to English all as ‘deficient’ with respect to its cognates in other Indo-European languages.
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6 introduction
sensitivity: RAs are universally context sensitive, both partial and AAT s
are existentially context sensitive, and NSs are not context sensitive.
This chapter also motivates an important empirical connection between
vagueness (i.e. the appearance of the properties described in Chapter 2)
and the scale structure classes in (9)–(12). In particular, I show that
the distribution of the puzzling properties of vague language is tied
to these lexical class distinctions, and, I propose, following authors
such as Kennedy and McNally (2005) and Kennedy (2007), that the
observed dependencies argue in favor of a closer relationship between
the phenomena of vagueness and scale structure than is often assumed
in the literature.
Chapter 4 (The Delineation Tolerant, Classical, Strict framework)
presents the DelTCS non-classical logical system for modelling the
relationship between context sensitivity, vagueness, and gradability in
the adjectival domain. I give an analysis of the context sensitivity/
vagueness patterns described in Chapter 3 within this framework, and
I discuss the empirical predictions that my analysis makes for a wide
range of semantic and pragmatic phenomena associated with adjectival
predicates.
Chapter 5 (Scale structure in Delineation semantics) presents both
new and previously discussed data associated with the scale structure
of members of the four principle classes of adjectives that are studied
in this work. Following much previous research, I argue that the adjec-
tives in each of the classes shown at the beginning of this chapter are
associated with scales that have different properties. In particular, as
we will see, there are empirical arguments for proposing that absolute
total adjectives are associated with scales that have maximal elements,
absolute partial adjectives are associated with scales that have minimal
elements, and relative adjectives are associated with scales that have
neither minimal nor maximal elements. I show in this chapter that
the association of an adjective with a scale with the correct properties
is already predicted by the analysis presented in Chapter 4 set within
the DelTCS architecture. In other words, I argue that, once we have an
(independently necessary) analysis of context sensitivity and vagueness
in the adjectival domain, we get an analysis of adjectival scale structure
“for free.”
Chapter 6 (Beyond Delineation semantics) studies a first class of
extensions of the framework developed in Chapters 2–5: theoreti-
cal/formal extensions. More precisely, this chapter explores to what
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8 introduction
2.1 Introduction
This chapter serves as an introduction to both one of the main empirical
phenomena to be analyzed in this monograph and the formal tools that
will be used in the analysis. As such, it has two main parts: in the first
part (Sections 2.2–2.3), I present the empirical phenomenon known as
vagueness in the linguistics and philosophical literatures, and I outline
why this phenomenon appears so threatening to our classical seman-
tic theories in logic and linguistics. Although the puzzles associated
with vague language have received an enormous amount of attention
in the field of philosophy, they have been much less studied from a
grammatical perspective. Therefore, in the first part of the chapter, I
describe the ways in which vague predicates challenge the currently
dominant approaches to natural language semantics. Thus, I argue that
the problem of accounting for vagueness is also a central problem for
the field of formal linguistics.
In the second part of the chapter (Sections 2.4–2.5), I present the basic
account of the puzzling properties of vague language that I will adopt
in this work: Cobreros et al.’s (2012b) Tolerant, Classical, Strict (TCS)
similarity-based non-classical logical framework. Unlike many other
works on this topic, I will not begin by reviewing all the many and varied
previous accounts of vague language, nor will I provide a comprehensive
comparison between the TCS approach and its competitors. There are
two reasons for this: firstly, excellent general introductions to the phe-
nomenon of vagueness and the wide variety of approaches on the market
already exist. Secondly, and more importantly, many of the debates in
the philosophical literature that have given rise to the wide range of
See, for example, Keefe (), Chapter of Smith (), the papers in Dietz and
Moruzzi () etc. See also van Rooij (), Cobreros et al. (b), and Cobreros et al.
(a) for comparisons between TCS and other frameworks.
A concrete example: As we will see in Section ., many speakers (the author included)
judge sentences of the form A is both P and not P as non-contradictory when A is a borderline
case of P. However, given that such FOL translations of such sentences are contradictions,
many philosophical theories maintain the contradictory nature of such statements. For
example, Keefe () says (p. ) (and see also similar sentiments in Fine () and
van Deemter ()):
Many philosophers would soon discount the paraconsistent option (almost) regardless of
how well it treats vagueness on the grounds of . . . the absurdity of p ∧ ¬p both being true
for many instances of p.
Thus, we can already see that theories of vagueness, which by design, have no way of dealing
with overt contradictions (either by allowing them, as in paraconsistent logics, or explaining
them away in a non-paraconsistent approach), are already inadequate semantic/pragmatic
theories for languages like English.
OUP CORRECTED PROOF – FINAL, 7/11/2016, SPi
Now we define the semantics for FOL. We first define models that consist
of a set of individuals D and a function m.
Definition 2.2.3 Model. A model is a tuple M = D, m where D is a
non-empty domain of individuals and m is a mapping on the non-logical
vocabulary satisfying,
1. for a constant a1 , m(a1 ) ∈ D
2. for a predicate P, m(P) ⊆ D.
In this chapter, for simplicity, I will limit the discussion to systems with unary predicate
because the n−ary predicate case is simply a straightforward generalization of the unary
predicate case.
OUP CORRECTED PROOF – FINAL, 7/11/2016, SPi
As is common, I will use a to refer to both the expression in the language and its
2
interpretation (I(a2 )), provided that it is clear from context which is meant.
Another random document with
no related content on Scribd:
its way. In Greenland, B. cordatus is very common in the dog, and probably also
in man, though few cases have been recorded. B. mansoni Cobb. (= B.
liguloides Leuck.) was, till recently, known only in the larval state from China and
Japan. Iijima, however, has found older specimens in the latter country. B.
cristatus Dav. is a species founded somewhat doubtfully on two fragments
found, one in a child, the other in a man, in France.
In sheep the most noteworthy and dangerous parasite is Coenurus cerebralis (or
the cystic stage of the dog-taenia, T. coenurus), which gives rise to the disease
known as "gid" or "staggers." It is found in various parts of the brain or spinal
cord, and the symptoms differ according to the position of the parasite. If this
presses upon one hemisphere the sheep describes circles and finally falls: if on
the optic lobes, the eyes are affected: if the pressure affects the cerebellum the
movements of the sheep are uncertain and incoordinated. Four or six weeks
after the appearance of the symptoms, death results from cerebral paralysis, or
from general debility, and the loss of sheep incurred by this disease (happily less
frequent in England than formerly) has been calculated by Youatt at a million for
France annually; at 35 per cent of the flocks for England in bad seasons; and
about 2 per cent for Germany. Besides sheep, which are most subject to "gid"
during their first year, various ruminants—Goat, Ox, Moufflon, Chamois, Roe,
Antelope, Reindeer, Dromedary—are attacked in the same way. A similar form,
Coenurus serialis Baill., is common in the wild rabbit in this country, and in
Australia in the hare and squirrel. It forms large swellings in the connective
tissue of various parts of the body, but usually does not affect the health of the
host. It is not known in what carnivore Taenia serialis Baill. normally occurs.
Experiments have, however, shown that it develops rapidly in dogs.
The preventive measures which are steadily diminishing the prevalence of the
Cestode parasites in man in some parts of Western Europe cannot be dealt with
here, but it may be noticed that the Jewish observance with regard to swine is
the surest preventive measure against taeniasis and trichinosis. Careful
inspection of meat and general cleanliness, are the leading measures that in
these hygienic matters secure the greatest immunity from disease.
The reproductive organs, unlike the preceding systems, are discontinuous from
one proglottis to the next. The male and female organs and their mutual
connexions, especially in the unsegmented Cestodes, may be compared in
detail with those of Trematodes, but the difference between the arrangement of
the generative organs of various Cestodes is very great.[107] The penis (Fig. 41,
cs) is evaginated through the male pore (Fig. 41, ♂ ), and inserted far into the
vagina (♀, vag) of the same or another segment of the tape-worm.
From this fact and the anatomical relations of the vagina, it is becoming
increasingly probable that the so-called uterus of Trematodes is an organ
corresponding to the vagina of Cestodes, and not to the uterus of Cestodes. The
latter opens to the exterior in Schistocephalus, Bothriocephalus, and some other
Cestodes of fishes by a special pore (Fig. 41, uto). Through this, some of the
eggs (which in these genera give rise to ciliated larvae) are enabled to escape,
and need not wait for the detachment of the proglottis, as must happen in the
Taeniidae, where the uterus is closed. This uterus, a true physiological one, is
probably the homologue of the "canal of Laurer" ("Laurer-Stieda canal," or
"vagina") of Trematoda. The fertilised ovum and yolk are brought together into
the "ootype," where the shell-gland forms the egg-shell around them (Fig. 41,
sh.gl) and the egg is then passed into the uterus. The ovum segments to form a
minute six-hooked larva, which may (Bothriidae, Fig. 42) or may not (Taeniidae)
be ciliated. Thus in Taenia serrata the proglottides are shed with the faeces of
the host (dog), and they protect the young from the desiccating influence of the
surroundings. If inadvertently eaten by a rabbit along with herbs, the proglottis
and larval envelope are digested, and by its six hooks the tiny larva bores
through the gastric wall into the portal vein, and so into the liver. Here the hooks
are thrown off, and the solid mass of cells becomes vacuolated.
Table for the Discrimination of the more usual Cestodes of Man and
Domestic Animals.[108]
I. Scolex in most cases with hooks; uterus with a median and lateral
branches; yolk-glands simple, median; genital pore single; dorsal
excretory vessel narrower than the ventral, without a circular
commissural trunk; eggs without pyriform apparatus (processes of the
ovarian membrane)
Gen. Taenia L. (s. str.)
A. Genital ducts pass on the ventral side of the
nerve and of the two longitudinal excretory
vessels T. crassicollis Rud.
B. Genital ducts pass between the dorsal and ventral longitudinal vessels.
a. Nerve present on dorsal side of genital ducts.
α. Head armed T. solium Rud.
β. Head unarmed T. saginata Goeze.
b. Nerve on ventral side of genital ducts.
Dog-Taeniae[109]
Head armed; genital pore marginal and
— Single
Many proglottides; strobila several centimetres long; small hooks
with guard.
Bifid hooks, which are
— 230µ-260µ long[110]; genital pore very
distinct T. serrata Goeze.
— 136µ-157µ long; genital pore not very
salient T. serialis Ball.
Entire large hooks, which are
— 180µ-220µ long; length of mature
segments double their width T. marginata Batsch.
— 150µ-170µ long; length of mature
segments treble their width T. coenurus Küch.
3-4 segments; a few mm. long T. echinococcus
v. Sieb.
— Double and bilateral Dipylidium caninum
L.
Head unarmed; two genital pores on ventral Mesocestoides
surface lineatus Goeze.
II. Scolex without hooks; one or two transverse uteri present; one or two
genital pores and yolk-glands, the latter never median; genital ducts
pass on the dorsal side of the nerve; eggs with pyriform apparatus.
A. One transverse uterus present.
a. Uterus with bullate egg-sacs; pyriform apparatus without horns; genital
ducts between dorsal and ventral vessels
Thysanosoma Dies.
α. Head large (1.5 mm.); square lobed testes in median field; posterior
margin of segments fimbriated; genital pore double
T. fimbriata Dies.
β. Head small; no fimbriae; pore rarely double T. giardii Riv.
b. Uterus without saccular dilatations; segments Anoplocephala E.
short, thick, and slightly imbricate Blanch.
Horse-Taeniae.
α. Head very large A. plicata
— No posterior lobes Zed.
— Four posterior lobes A. perfoliata
Goeze.
β. Head small, without posterior lobes A. mamillana Mehl.
B. Two uteri and two genital pores present; horns of pyriform apparatus
well developed; genital ducts pass on the dorsal side of the
longitudinal vessels
Moniezia R. Bl.
a. Interproglottidal glands [111] arranged in linear series (planissima
group)
M. planissima S. and H. M. benedeni Mz. M. neumani Mz.
b. Interproglottidal glands saccular (expansa group)
M. expansa Rud. M. oblongiceps S. and H. M. trigonophora S. and H.
c. Interproglottidal glands absent (denticulata M. denticulata Rud.
group) M. alba Perr.
C. Uterus single or double, without spore-like egg-sacs; eggs with a single
shell; genital pores irregularly alternate; strobila narrow; testes absent
from median part of the field
Stilesia Raill.
a. A transverse uterus in middle part of median S. centripunctata
field; head 2 mm. diameter Riv.
b. Two lateral uteri in each segment; head less
than 1 mm. in diameter S. globipunctata Riv.
III. Scolex almost invariably provided with hooks; genital pores on left border
of segment; eggs with three shells but no cornua. Segments broader
than long; posterior angles salient.
Hymenolepis Weinl.
a. Scolex with a single series of 24-30 hooks, each 14-18µ long
H. nana v. Sieb. H. murina Duj.
b. Scolex very small, unarmed H. diminuta Rud.
IV. Scolex provided with two elongated muscular pits. Body segmented; three
genital apertures in middle of ventral surface
Bothriocephalus Rud.
Body 2-20 metres in length
B. latus Brems. B. cristatus Dav. (doubtful species). B. cordatus Leuck. B.
mansoni Cobb. (= B. liguloides Leuck.)
CHAPTER IV
MESOZOA
DICYEMIDAE—STRUCTURE—REPRODUCTION—OCCURRENCE: ORTHONECTIDAE—
OCCURRENCE—STRUCTURE: TRICHOPLAX: SALINELLA.
The Mesozoa are an obscure group, the position of which in the animal kingdom
is still doubtful. The name Mesozoa was given to the group by its discoverer, E.
van Beneden,[112] as he concluded that they were intermediate between the
Protozoa and the higher Invertebrates. Recent authors, however, have called
attention to the resemblance existing between them and the "sporocysts" of
Trematodes, and though we still are ignorant of certain important points in their
life-histories, the Mesozoa are most conveniently (and probably rightly)
considered as an appendix to the Platyhelminthes.
The animals composing this group are minute and parasitic, and are composed
of a small number of cells. They may be divided into two families: the
Dicyemidae, which occur exclusively in the kidneys of certain Cephalopods
(cuttle-fish); and the Orthonectidae, which live in the brittle-star Amphiura
squamata, the Nemertine Nemertes lacteus, or the Polyclad Leptoplana
tremellaris. In addition to the undoubted Mesozoa, certain anomalous forms—
Trichoplax adhaerens and Salinella salve—may be referred to this group.
The occurrence of the known species of Dicyemids (a group which has not been
investigated on our coasts) is as follows:—
Species. Host.
Dicyema typus van Ben. Octopus vulgaris.
D. clausianum van Ben. O. macropus.
D. microcephalum Whit. O. de Filippi.
D. moschatum Whit. Eledone moschata.
D. macrocephalum van Ben. Sepiola rondeletii.
D. truncatum Whit. Rossia macrosoma, Sepia elegans, S.
officinalis.
D. schultzianum van Ben. S. biseralis, Octopus vulgaris.
Dicyemennea eledones Wag. Eledone moschata, E. aldrovandi.
D. mülleri Clap. E. cirrosa.
D. gracile Wag. Sepia officinalis.
Conocyema polymorphum S. officinalis, Octopus vulgaris.
van Ben.
Rhopalura giardii is of distinct sexes. Either males or females are found in one
Amphiura. Two kinds of females, flattened unsegmented, and cylindrical
segmented forms, are known. They consist of a ciliated ectodermal layer
enclosing an endodermal mass of eggs, between which is a fibrillar layer usually
considered to be of a muscular nature. The cylindrical female gives rise to eggs
which develop, probably exclusively, into males. The flattened female produces
eggs from which females alone arise, though the origin of the two forms of this
sex is not well ascertained. The males contain spermatozoa which fertilise the
eggs of the cylindrical female, whereas the ova of the flat form probably develop
parthenogenetically.
BY
LILIAN SHELDON
Staff Lecturer in Natural Science, Newnham College, Cambridge.
CHAPTER V
NEMERTINEA
INTRODUCTORY—EXTERNAL CHARACTERS—ANATOMY—CLASSIFICATION—
DEVELOPMENT—HABITS—REGENERATION—BREEDING—GEOGRAPHICAL
DISTRIBUTION—LAND, FRESH-WATER, AND PARASITIC FORMS—AFFINITIES
The Nemertinea form a compact group, the affinities of which have not been at
present clearly determined. Several species were mentioned and described in
the works of various naturalists during the latter half of the eighteenth century,
though their anatomy was not understood until considerably later. The first
mention of any member of the group was made by the Rev. W. Borlase in his
Natural History of Cornwall, published in 1758. He gives a short description and
a rough figure of Lineus marinus. From that time the increase in the knowledge
of the group was very gradual. New species were from time to time described,
but few of the descriptions could boast of much completeness, and many
erroneous views were held until comparatively recent years. The group was very
variously classified, but the general arrangement in early times seems to have
been to unite it with the Planarians. Valuable contributions to the history of the
development were made in 1848 and the few subsequent years by Desor,[118]
Gegenbaur,[119] Krohn,[120] and Leuckart and Pagenstecher[121]; and more
recently by Metschnikoff[122] and Salensky.[123]
Nemertines for the most part closely resemble one another in all essential
points, though they differ considerably in size, colour, and external details. They
vary in length from less than an inch to thirty yards, this extreme size being
attained by Lineus marinus.
Fig. 48.—Lineus marinus Mont., from the living specimen in the coiled condition.
Plymouth. × 1. a, Anterior end; b, posterior end.
Fig. 49.—L. marinus, from the same specimen as Fig. 48, in the expanded
condition. a, Anterior end; b, posterior end.
Nemertines are common on the British coasts; about forty species have been
recorded from this area. On turning over a stone on a sandy or muddy shore in a
pool left by the receding tide, there may often be seen a coiled mass, having the
appearance of a uniform slimy string twisted into a complicated knot. If it be
carefully removed, the ends can generally be made out, one bluntly rounded and
the other slightly tapering (Fig. 48, a and b). Occasionally there may be seen
attached to the blunter end a fine thread, which moves about freely. This thread
may, by an instantaneous movement, be drawn into the body, no trace of its
existence being left except at the tip of the head, where a small pore is visible;
this is the orifice through which it was withdrawn. Shortly afterwards the thread
may be again shot out, the process being instantaneous and often accomplished
with great force. This thread (Fig. 50, p) is the proboscis, a very important and
characteristic organ in Nemertines.
Most Nemertines are marine; they are mostly indifferent to climate and to the
nature of the soil on which they live.
Fig. 50.—Side view of head of Cerebratulus (Micrura) tristis Hubr., showing the
everted proboscis. Naples. × 2. Drawn from a spirit specimen. c.s, Cephalic
slit; m, mouth; p, proboscis.
In the ordinary forms the posterior end of the body is pointed either bluntly or
sharply. The head is somewhat broader than the rest of the body, and often
assumes a spatulate form. Eyes (Fig. 51, e) are usually present either in one or
several pairs, or in symmetrically-arranged groups on each side of the head. The
mouth (Fig. 58, m) is situated near the front end of the body on the ventral
surface, and is usually rendered conspicuous by being surrounded by thick
tumid lips. It varies in form from being slit-like to elliptical. At the anterior end of
the body a small terminal pore occurs; this is the external opening of the
proboscis (Fig. 51, p.p).
Nemertines are often very diversely and brilliantly coloured, the hues most
commonly found being white, yellow, green, deep purple, and various shades of
red and pink. The ventral surface is usually paler in colour than the dorsal, and
the latter is often marked by longitudinal and transverse stripes (Fig. 59) in
contrasting colours.
1. An external epidermic layer (ep), consisting of ciliated cells, among which are
placed numerous unicellular glands. These glands probably secrete the mucus
in which the Nemertine is usually enveloped; their contents when in the body are
very highly refracting. The epidermis rests on a basement membrane (b.m).
2. The two or three muscular layers, arranged as either an external circular and
an internal longitudinal, or an inner and an outer circular separated by a
longitudinal layer, or, as in the figure (c.m and l.m), two longitudinal separated by
a circular layer.
Fig. 51.—Amphiporus lactifloreus Johnst., drawn from the living specimen, from
the dorsal surface. Plymouth. × 2. e, Eyes; g, generative organs; n.g, nerve
ganglion; p.p, proboscis pore; p, proboscis.
3. A fairly thick connective-tissue layer often found between the epidermis and
the muscles, into which latter it gradually merges (s.t).
The Digestive System.—The mouth is placed on the ventral surface near the
anterior end of the body (Figs. 53, 58, m). It leads into a straight oesophagus
(Fig. 53, oes), whence passes off the intestine (int), which is continued as a
straight non-convoluted tube to the anus (a), situated terminally at the posterior
end of the body. The intestine is thrown out throughout the greater part of its
course into paired lateral pouches.
The eversion is effected by a turning inside out of the anterior part of the
proboscis (Fig. 54). The process whereby the proboscis is retracted has been
very aptly compared to the effect which would be produced by the inversion of
the finger of a glove, accomplished by pulling a string attached to its tip on the
inside, the anterior muscular part being comparable to the finger and the
glandular part to the string. It is thus obvious that in the everted condition the