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ASSESSMENT IN SECOND LANGUAGE
PRONUNCIATION

Assessment in Second Language Pronunciation highlights the importance of pronunciation

in the assessment of second language speaking proficiency. Leading researchers from
around the world cover practical issues as well as theoretical principles, enabling the
understanding and application of the theory involved in assessment in pronunciation.
Key features of this book include:

•â•¢ Examination of key criteria in pronunciation assessment, including intelligibility,

comprehensibility and accentedness;
•â•¢ Exploration of the impact of World Englishes and English as a Lingua Franca
on pronunciation assessment;
•â•¢ Evaluation of the validity and reliability of testing, including analysis of scoring
methodologies;
•â•¢ Discussion of current and future practice in assessing pronunciation via speech
recognition technology.

Assessment in Second Language Pronunciation is vital reading for students studying

modules on pronunciation and language testing and assessment.

Okim Kang is Associate Professor of Applied Linguistics/TESL at Northern

Arizona University, USA.

April Ginther is Associate Professor of Second Language Studies at Purdue

University, USA.
ASSESSMENT IN
SECOND LANGUAGE
PRONUNCIATION

Edited by Okim Kang and April Ginther

First published 2018
by Routledge
2 Park Square, Milton Park, Abingdon, Oxon OX14 4RN
and by Routledge
711 Third Avenue, New York, NY 10017
Routledge is an imprint of the Taylor & Francis Group, an informa business
© 2018 selection and editorial matter, Okim Kang and April
Ginther; individual chapters, the contributors
The right of the editors to be identified as the authors of the
editorial material, and of the authors for their individual chapters,
has been asserted in accordance with sections 77 and 78 of the
Copyright, Designs and Patents Act 1988.
All rights reserved. No part of this book may be reprinted or
reproduced or utilised in any form or by any electronic, mechanical,
or other means, now known or hereafter invented, including
photocopying and recording, or in any information storage or
retrieval system, without permission in writing from the publishers.
Trademark notice: Product or corporate names may be trademarks
or registered trademarks, and are used only for identification and
explanation without intent to infringe.
British Library Cataloguing-in-Publication Data
A catalogue record for this book is available from the British Library
Library of Congress Cataloging-in-Publication Data
A catalog record for this book has been requested

ISBN: 978-1-138-85686-8 hbk

ISBN: 978-1-138-85687-5 pbk
ISBN: 978-1-315-17075-6 ebk

Typeset in Bembo
by Swales & Willis Ltd, Exeter, Devon, UK
CONTENTS

List of figures vii

List of tables viii
Acknowledgements ix
Notes on contributors x

Introduction 1
Okim Kang and April Ginther

PART I
Current issues in pronunciation assessment 9

1 Measurement of accentedness, intelligibility,

and comprehensibility 11
Ron Thomson

2 Validity in pronunciation assessment 30

Luke Harding

3 Pronunciation assessment in the context of World Englishes 49

Slobodanka Dimova

4 Listeners and raters: similarities and differences in

evaluation of accented speech 67
Xun Yan and April Ginther
viâ•…Contents

5 Assessing pronunciation for research purposes with

listener-based numerical scales 89
Daniel R. Isbell

PART II
Technology and pronunciation assessment 113

6 Pronunciation features in rating criteria 115

Romy Ghanem and Okim Kang

7 Using speech processing technology in assessing

pronunciation 137
Alistair Van Moere and Masanori Suzuki

8 Automated assessment of pronunciation in spontaneous

speech 153
Anastassia Loukina, Larry Davis, and Xiaoming Xi

Index 172
FIGURES

2.1 Inferential links in an interpretive argument 33

5.1 Histograms of averaged and individual comprehensibility scores 96
5.2 Histograms of comprehensibility scores awarded by each rater 98
5.3 Category probability curves for comprehensibility scores 100
5.4 Item characteristic curve for the comprehensibility scale 101
5.5 Histograms of averaged and individual accentedness scores 102
5.6 Histograms of accentedness scores awarded by each rater 104
5.7 Category probability curves for accentedness scores 105
5.8 Item characteristic curve for the accentedness scale 106
6.1 Long voicing lead of consonant /k/ as produced by an L1 speaker 123
6.2 Short voicing lead of consonant /b/ as produced by an L1 speaker 123
6.3 First formant spectrograms of monophthong /i/ using PRAAT 124
6.4 Vowel measurement of the tense vowel /i/ using PRAAT 124
6.5 Representation of a silent pause using PRAAT 125
6.6 Stressed syllable in children as produced by an
advanced Chinese speaker of English 126
6.7 The prominent syllable of a tone unit as produced
by an advanced Chinese speaker 126
7.1 Speech contours of the word “strategy” 147
TABLES

1.1 Sample of studies measuring strength of foreign accent

and their operationalization 16
1.2 Sample of studies measuring intelligibility and their
operationalization 18
1.3 Sample of studies measuring comprehensibility and their
operationalization 20
2.1 Some key validity questions for pronunciation assessment 35
2.2 Common research methods in pronunciation assessment
validation research 42
5.1 Descriptive statistics for comprehensibility scores 96
5.2 Reliability indices for comprehensibility ratings 97
5.3 Summary statistics of speaker and listener measures
for comprehensibility 97
5.4 Descriptive statistics for accentedness scores 102
5.5 Reliability indices for accentedness ratings 103
5.6 Summary statistics of speaker and listener measures
for accentedness 103
ACKNOWLEDGEMENTS

We would like to begin by acknowledging the influence and importance of the

annual conference on Pronunciation in Second Language Teaching and Learning
(PSLLT) to the genesis of this volume. The conference theme, Setting the Course
for Pronunciation Teaching and Assessment, PSLLT, 2012, Vancouver, BC, Canada,
and the many inspiring presentations and discussions generated in response, set the
course for the development of Assessment in Second Language Pronunciation.
There are many people who we thank for their support during the preparation
of this volume. First and foremost, we thank the authors for contributing their
work. Their perseverance and persistence in the preparation of the volume are
much appreciated. Helen Tredget at Routledge has been of enormous help and
support in getting all of the pieces in order and of working with us in the process of
finalizing the product. We also thank the anonymous reviewers for their valuable
comments and suggestions. Nadia Seemungal-Owen, Senior Editor at Routledge,
was instrumental for her initial support and in carrying this book from an idea to
its current form. Our thanks go to Dawn Burns as well for her valuable help with
copy editing and preparation of the final manuscript.
CONTRIBUTORS

Larry Davis is a Managing Research Scientist at Educational Testing Service,

Princeton, NJ, USA. His primary research focus is speaking assessment, including
speaking constructs, task design, scoring rubrics, rater expertise and behavior, and
applications of automated scoring technology. His publications include articles
on rater expertise, partner effect in paired oral tests, and the use of rubrics in
language teaching.

Slobodanka Dimova is an Associate Professor at the University of Copenhagen,

Denmark. Her research interests include language testing and measurement, oral
language production, and English medium instruction in higher education. She is
the Test Coordinator of the Test of Oral English Proficiency for Academic Staff
(TOEPAS) and serves as a Book Editor of the Language Testing journal.

Romy Ghanem is a doctoral candidate at Northern Arizona University, USA.

Her primary research interests include speech production and perception as well as
language structure. She has conducted research in linguistic and reverse linguistic
stereotyping and used corpus methods to examine the formation of genitives. Her
current research involves the effect of first language on the speech alignment of
different linguistic features.

April Ginther is an Associate Professor of Second Language Studies and Linguistics

at Purdue University, USA, where she teaches courses in language testing and
quantitative research. Her research interests include the development and use of
English language proficiency tests, the integration of testing and instruction, and
the measurement of language proficiency over time. She is currently the Co-Editor
of the journal Language Testing.
 Contributorsâ•… xi

Luke Harding is a Senior Lecturer of Linguistics and English Language at Lancaster

University, UK. His research interests are mainly in the area of language assessment,
particularly listening, speaking, assessor decision-making, language assessment lit-
eracy, and the challenges of World Englishes and English as a Lingua Franca for
language assessment.

Daniel R. Isbell is a PhD student in Second Language Studies at Michigan State

University, USA. His research interests include language assessment, instructed
SLA, and L2 pronunciation. Daniel has conducted research on pronunciation instruc-
tion and the role of pronunciation in task-based interaction. He has presented his
work at international conferences and has published in the journal Language Testing.

Okim Kang is an Associate Professor of Applied Linguistics at Northern Arizona

University, USA. Her research specialties are L2 pronunciation, oral proficiency
assessment, language attitudes, speech perception and production, and automated
scoring systems. She serves on Editorial Boards of several journals (e.g., TESOL
Quarterly, Language Testing, Journal of Second Language Pronunciation) and is Co-Editor
of an online journal, TESL-EJ Book Reviews.

Anastassia Loukina is Research Scientist in Natural Language Processing and

Speech group at Educational Testing Service, Princeton, NJ, USA. She is a phoneti-
cian by training and her research focuses on automated speech scoring and especially
the automated evaluation of pronunciation and intelligibility. She published articles
on various aspects of automated speech scoring as well as other applications of natu-
ral language processing technologies and machine learning in assessment.

Masanori Suzuki is Director of Test Development at Pearson’s automated scor-

ing division where he oversees development projects and validation research of
automated language assessments. He holds an MA in Teaching English to Speakers
of Other Languages (TESOL) from San Francisco State University, USA. His
research interests are language testing, SLA, and psycholinguistics.

Ron Thomson is a Professor of Applied Linguistics at Brock University in the

Niagara Region, Canada. His research focuses on the development of L2 oral flu-
ency and pronunciation. He is also the creator of www.englishaccentcoach.com, a
free web-based application which helps English learners improve their perception
of English vowels and consonants.

Alistair Van Moere is head of a division in Pearson that researches, develops, and
delivers various educational assessments, including automatically scored language
tests. He has worked in language training and assessment for over 20 years and
has published 20 research articles in peer-reviewed journals on the subjects of oral
language assessment and automated scoring.
xiiâ•…Contributors

Xiaoming Xi is Executive Director of Global Education and Workforce at

Educational Testing Service, Princeton, NJ, USA. Her research spans broad areas
of theory and practice, including validity and fairness issues, test validation methods,
approaches to defining test constructs, validity frameworks for automated scoring,
automated scoring of speech, the role of technology in language assessment and
learning, and test design, rater, and scoring issues. She edited a special issue on
automated scoring and feedback systems in Language Testing and has been awarded
five patents in this area.

Xun Yan is Assistant Professor of Linguistics and SLATE (Second Language

Acquisition and Teacher Education) at University of Illinois at Urbana-Champaign,
USA. His research interests include post-admission language assessments, language
assessment literacy, pronunciation and speech intelligibility, and test score use in
educational settings. His work has been published in Language Testing, Assessing
Writing and Journal of Second Language Writing.
INTRODUCTION
Okim Kang and April Ginther

The assessment of second language (L2) speaking proficiency has been of central
interest to researchers in Applied Linguistics since the first discussions of commu-
nicative competence (Hymes, 1972; Canale & Swain, 1980); however, research on
pronunciation, once marginalized in part due to its association with discrete aspects
of oral production (Lado, 1961, 1964), is now emerging as a revitalized field of
inquiry with its own important implications and concerns. Part of this resurgence
can be attributed to a shift in focus from perceptions of accentedness to broader
aspects of performance, primarily intelligibility and comprehensibility.
Since the mid 1990s there has been enormous growth in research on L2 pronun-
ciation. Pronunciation is an essential aspect of the assessment of oral skills because it
helps us understand the fundamentals in the process of the construction of spoken
discourse in L2 performance; that is, listeners begin by processing individual sounds
constructed by L2 speakers to arrive at an interpretation for a stream of speech. The
discrete sounds of speech remain a critical area of investigation as listeners tend to
attribute native/nonnative speaker status on the basis of pronunciation (Luoma,
2004). Pronunciation is also an important facet of proficiency on which most L2
learners have ready views and clear motivations (Leather, 1999). In recognition
of such significance, the 4th Pronunciation in Second Language Teaching and
Learning (PSLLT) 2012 conference held in Vancouver, British Columbia, focused
its theme on pronunciation and assessment. In fact, it is PSLLT 2012 that initially
motivated the idea of this edited volume.
However, the history of L2 pronunciation has been compared to a pendulum
swinging back and forth between times when it has been completely ignored,
and times when it has been of primary importance. As the role of pronunciation
in general L2 language learning has been a history of extremes (Levis, 2005), the
role of assessment has equally fluctuated with the times. In some cases, assessment
has focused on the accuracy of segmentals, in others, on the approximation or the
2â•… Okim Kang and April Ginther

mastery of suprasegmentals. Since about the year 2005, however, L2 pronuncia-

tion research has gleaned a renewed focus on intelligibility and comprehensibility,
and accentedness. The assessment of pronunciation is a reflection of these historical
perspectives, and the methods of assessing pronunciation have reflected the times.
Thomson examines the representation of accentedness, comprehensibility, and
intelligibility – criterial components of pronunciation assessment. He traces the
transformative influence of Munro and Derwing who have addressed, through
a long line of related investigations, both the definition and inter-relatedness of
the constructs of accentedness, comprehensibility, and intelligibility. These studies
have transformed the field of L2 pronunciation. Notions that accentedness, com-
prehensibility, and intelligibility are related but partially independent constructs
have paved the way for pronunciation teaching and research to put accent in its
place (Derwing & Munro, 2009) and emphasize the end goal of comprehensibility.
Interest in these inter-related constructs has led researchers to seek and discover
how they are best operationalized and measured, how they affect listeners’ rat-
ings, and how they should be addressed in the ESL/EFL classroom. Thomson’s
comprehensive discussion on methodological approaches to the measurement and
evaluation of leading pronunciation constructs should be of use to teachers and
researchers in the field.
One of the critical issues involved in assessing pronunciation is determining
to what extent the measures of pronunciation constructs are valid and reliable.
Harding argues that L2 pronunciation assessment presents unique challenges in
drawing valid inferences from performance, to score assignment, to the ulti-
mate decisions for which a pronunciation assessment was intended, and to the
pedagogical and social consequences beyond. He raises legitimate questions: Are
administration and scoring procedures accurate and consistent? Does the task used
in assessment elicit relevant target features? Does the test yield a score which is fit
for decision-making purposes? Is the assessment fair? While such inquiries can be
generally perceived across all types of language assessments, some of them are cen-
tral to the concept of validity in L2 pronunciation.
The discussion of validity is directly related to the context of World Englishes
(WE), where the reliance on prestigious inner circle norms of native English
has been challenged. This validity issue examines the sociolinguistic realities of
diverse language learners’ actual use (Elder & Harding, 2008). Given that defining
a standard norm is problematic in the era of globalization, where new norms are
emerging, so the assessment of L2 pronunciation must address more questions now
than ever before. Should the pronunciation assessment focus on accuracy with
respect to a norm or on mutual intelligibility? What role does accentedness play
with respect to mutual intelligibility? Providing that within native-speaker varieties
of English (e.g., British, American, New Zealand English or Australian English)
variability exists, the focus on accuracy (i.e., deviations from a native-speaker
norm) introduces the complexity of identification and selection of multiple norms
for teaching and assessment: Whose norm, which norm is the most appropriate?
Furthermore, research in L2 pronunciation now gears toward setting a realistic goal
 Introductionâ•… 3

for pronunciation acquisition, i.e., intelligibility and comprehensibility – rather

than native-like pronunciation (Munro & Derwing, 2011, 2015). While most
large-scale language assessments still appeal to established standard varieties, the
“educated native speaker” has receded into the background, and comprehensibility
and intelligibility have become the focus.
In all pronunciation endeavors, the importance of listeners cannot be under-
estimated, and background characteristics that listeners bring to assessment tasks
have been found to influence their evaluations of accentedness, comprehensibility,
and intelligibility. In this volume, two research chapters examine listeners’ speech
judgments and the background factors that affect their evaluations’ speech. Because
idiosyncratic listener judgments pose a threat to both the reliability and validity of
operational tests, listeners become raters when they are trained to rate to a scale.
Yan and Ginther discuss the differences between listeners and raters, and the effects
of rater training in speaking and pronunciation assessment are highlighted.
The use of listeners who apply minimally represented scales in broad, applied
research contexts invites our attention to reliability and scale representation
(validity). While 9-point Likert-type scales are used widely, idiosyncratic appli-
cation may be masked by the over-generality of the scales. Identifying differences
across raters in their application of a scale creates challenges when there are only
minimal specifications that are provided. Raters in fact have reported some dif-
ficulty when attempting to differentiate middle scale points (Isaacs & Thomson,
2013). Empirical evidence is needed to better understand how listeners may
differentially interpret scales based on their own underlying representations of
different pronunciation constructs.
On a more technical side, L2 pronunciation involves many acoustic features
of the speech stream such as the quality of vowels and consonants, the presence
and placement of pauses, as well as broader, measurable components of prosody:
stress, rhythm, and intonation. Understanding how acoustic features influence
human rating of pronunciation presents a rich domain for research. Thanks to
advances in speech science, computer-assisted instruments can aid in examining
some elements of the acoustic properties of L2 pronunciation. The knowledge
of these instrumentally analyzed pronunciation properties has the potential to
advance our understanding of speech production and inform rubric development
and rater training in oral proficiency testing. The physical properties of the acoustic
measures can also build bases for speech recognition and processing techniques,
which has increasingly attracted the attention of language testers. As topics on the
improvement of automated speech recognition (ASR) effectiveness for L2 speech
continues to be of interest to L2 researchers (e.g., Oh, Yoon & Kim, 2007), better
understanding of this technology-driven approach to pronunciation assessment is
of benefit to all interested parties (Van Moere & Downey, 2016).
Overall, despite researchers’ and language teachers’ growing interest in the
issues mentioned above, there are limited resources that address these issues in
a systematic and comprehensive manner. A reliable and wide-ranging source of
information that reflects current knowledge of such topics is beneficial. In the
4â•… Okim Kang and April Ginther

current volume, we have attempted to serve this purpose. Our contributors

offer their specialized understanding on the importance of pronunciation in the
assessment context. This volume provides a bridge – highlighting both com-
mon concerns and differences for researchers in both domains. We hope that this
volume can be used by students and instructors at upper-level undergraduate,
graduate, and post-graduate courses as well as by established and emerging scholars
involved in research on L2 speech and language assessment.
This volume offers detailed accounts of issues related to L2 pronunciation and
assessment. It is divided into two parts containing eight chapters written both by
applied linguists who specialize in language assessment together with researchers
that expertize in the topic of L2 pronunciation and speaking. Some of the topics
that guide the chapter selections include:

•â•¢ measurement and evaluation of pronunciation constructs;

•â•¢ validity and reliability of pronunciation assessment;
•â•¢ World Englishes and the assessment of pronunciation;
•â•¢ listeners’ individual variation and their background characteristics in L2 accent;
•â•¢ pronunciation features to inform rating criteria;
•â•¢ assessing pronunciation via speech technology.

Beginning in Part I, “Current issues in pronunciation assessment,” Thomson’s

“Measurement of accentedness, intelligibility, and comprehensibility” (Chapter 1)
surveys methodological approaches to the measurement and evaluation of leading
constructs associated with L2 pronunciation, with the goal of providing a detailed
description for applied linguists who are just entering this area of research and for
established researchers looking for a clear overview. The chapter begins by outlining
what the author considers to be valid reasons for measuring L2 pronunciation. Three
dominant L2 pronunciation constructs are then introduced: intelligibility, compre-
hensibility, and accentedness. While these terms are widely used in the literature,
they are not always applied consistently. Thomson describes key findings of research
investigating accent, intelligibility, and comprehensibility and suggests new direc-
tions for L2 pronunciation assessment and recommendations for further research.
In Chapter 2, “Validity in pronunciation assessment,” Harding draws on recent
developments in validity theory to illuminate the issues facing all test developers
but particularly those who are interested in pronunciation assessment. Threats to
validity are identified with reference to pronunciation assessment, and research,
which has investigated such threats, is discussed. He also outlines common research
methods in conducting pronunciation assessment validation research and suggests
new directions for validity-related research in pronunciation assessment, with a
recommendation that pronunciation assessment is a case where effect-driven test
design is clearly warranted.
Dimova, in “Pronunciation assessment in the context of World Englishes”
(Chapter 3), begins her chapter by outlining the early WE conceptualizations of
pronunciation through the model of understanding in cross-cultural communication.
16â•… Ron Thomson

Others’ contributions and research

Foreign accent
There is relative agreement about what constitutes a foreign accent. Referring to
foreign accent as a perceptual phenomenon on the part of a listener. Scovel (1969)
puts it plainly, “the existence of foreign accents is dependent upon the ability of
native speakers to recognize them” (p. 248). This ability to recognize accent is
acute. For example, Flege (1984) demonstrated that listeners could reliably detect
the presence of a foreign accent in speech samples as short as 30 milliseconds, while
Munro, Derwing, and Burgess (2010) found listeners could detect a foreign accent
in speech recordings that were played backwards.
Most researchers define foreign accent in terms of how much L2 speech devi-
ates from a target variety. For example, Kennedy and Trofimovich (2008) define
it as “how closely the pronunciation of an utterance approaches that of a native
speaker” (p. 461), a definition reiterated by O’Brien (2014). Similarly, Isaacs and
Thomson (2013) describe foreign accent as “how different the speaker sounds from
a NS” (p. 141), while Jułkowska and Cebrian (2015) define foreign accentedness
“as the listener’s perception of how closely the pronunciation of an L2 speaker
mirrors the pronunciation of a native speaker of a given language” (p. 212).
While the definitions above all focus on the extent to which a foreign accent
differs from a target native variety, many of those differences are rooted in patterns
carried over from speakers’ first languages. Thus, instead of comparing accented

TABLE 1.1╇ Sample of studies measuring strength of foreign accent and their
operationalization

Study Operationalization of foreign accent

Kennedy & Scalar ratings of T/F sentences; semantically meaningful/

Trofimovich (2008) anomalous sentences: 1 = no non-native accent, 9 = strong
non-native accent
Trofimovich & Isaacs Scalar ratings of extemporaneous speech samples: 1 = heavily
(2012) accented, 9 = not accented at all
Isaacs & Thomson Scalar ratings of extemporaneous speech samples: 1 = heavily
(2013) accented, 9 = not accented at all and a 5-point scale with the
same endpoints
Kraut & Wulff (2013) Scalar ratings of academic passage read-alouds: How strong is the
speaker’s accent from 1–7?
O’Brien (2014) Scalar ratings of extemporaneous speech samples: 1 = no accent,
9 = extremely strong accent
Jułkowska & Cebrian Scalar ratings of a 20-item sentence read-aloud task: 1 = no
(2015) foreign accent, 9 = strong foreign accent
Saito, Trofimovich, & Scalar ratings of extemporaneous speech: Digital slider was used
Isaacs (2015) with sensitivity from 0–1,000 with endpoints not reported
Saito, Trofimovich, & Scalar ratings of extemporaneous speech samples: 1 = no accent,
Isaacs (2016) 9 = heavily accented
Another random document with
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 Fig. 154—Young larva of Stylops on a bee's-hair. Greatly magnified.
(After Newport.)

There is no exact information as to how the young triungulins find

their way to the bee-larvae they live in. Here again the discrepancy
of opinion that prevails is probably due to great difference really
existing as to the method. When a Stylops carried by an Insect (a
Hymenopteron, be it noted, for we have no information whatever as
to Hemiptera) produces young, they cover the body of the host as if
it were powdered, being excessively minute and their numbers very
great; many hundreds, if not thousands, of young being produced by
a single Stylops. The species of the wasp genus Polistes are
specially subject to the attacks of Stylops; they are social Insects,
and a stylopised specimen being sickly does not as a rule leave the
nest; in this case the Stylops larva may therefore have but little
difficulty in finding its way to a Hymenopterous larva, for even though
it may have to live for months before it has the chance of attaching
itself to a nest-building female, yet it is clearly in the right
neighbourhood. The bee genus Andrena has, however, quite
different habits; normally a single female makes her nest
underground; but in the case of a stylopised female it is certain that
no nest is built, and no larvae produced by a stylopised example, so
that the young triungulins must leave the body of the bee in order to
come near their prey. They can be active, and have great powers of
leaping, so that it is perhaps in this way possible for them to attach
themselves to a healthy female bee.
 Fig. 155.—Portion of early stages of Xenos rossii. (After von Siebold.)
A, Small male larva; B, small female larva; C, full-grown male
larva; D, full-grown female larva; E, the so-called "cephalothorax"
and adjacent segment of adult female. (The newly-hatched larva
is very much like that of Stylops shown in Fig. 154.)

We have still only very imperfect knowledge as to the structure and

development of Strepsiptera. Indeed but little information has been
obtained since 1843.[160] Before that time the mature female was
supposed to be a larva, and the triungulins found in it to be
parasites. Although the erroneous character of these views has been
made clear, the problems that have been suggested present great
difficulties. Apparently the change from the triungulin condition (Fig.
154) to the parasitic larvae (Fig. 155, A, B) is extremely great and
abrupt, and it appears also that during the larval growth considerable
sexual differentiation occurs (Fig. 155, C, D); details are, however,
wanting, and there exists but little information as to the later stages.
Hence it is scarcely a matter for surprise that authorities differ as to
which is the head and which the anal extremity of the adult female.
Von Siebold apparently entertained no doubt as to the part of the
female that is extruded being the anterior extremity; indeed he called
it a cephalothorax. Supposing this view to be correct, we are met by
the extraordinary facts that the female extrudes the head for
copulatory purposes, that the genital orifice is placed thereon, and
that the young escape by it. Meinert[161] contends that the so-called
cephalothorax of the adult is the anal extremity, and that fertilisation
and the escape of the young are effected by the natural passages,
the anterior parts of the body being affected by a complete
degeneration. Nassonoff, in controversion of Meinert, has recently
pointed out that the "cephalothorax" of the young is shown by the
nervous system to be the anterior extremity. It still remains, however,
to be shewn that the "cephalothorax" of the adult female
corresponds with that of the young, and we shall not be surprised if
Meinert prove to be correct. The internal anatomy and the processes
of oogenesis appear to be of a very unusual character, but their
details are far from clear. Brandt has given some particulars as to the
nervous system; though he does not say whether taken from the
male or female, we may presume it to be from the former; there is a
supra-oesophageal ganglion, and near it a large mass which
consists of two parts, the anterior representing the sub-oesophageal
and the first thoracic ganglia, while the posterior represents two of
the thoracic and most of the abdominal ganglia of other Insects; at
the posterior extremity, connected with the other ganglia by a very
long and slender commissure, there is another abdominal ganglion.
[162]

It is a matter of great difficulty to procure material for the prosecution

of this study; the fact that the instars to be observed exist only in the
interior of a few Hymenopterous larvae, which in the case of the bee,
Andrena, are concealed under ground; and in the case of the wasps,
Polistes, placed in cells in a nest of wasps, adds greatly to the
difficulty. It is therefore of interest to know that Strepsiptera occur in
Insects with incomplete metamorphosis. They have been observed
in several species of Homoptera; and the writer has a large
Pentatomid bug of the genus Callidea, which bears a female
Strepsipteron apparently of large size. This bug[163] is abundant and
widely distributed in Eastern Asia, and it may prove comparatively
easy to keep stylopised examples under observation. Both v. Siebold
and Nassonoff think parthenogenesis occurs in Strepsiptera, but
there appear to be no facts to warrant this supposition. Von Siebold
speaks of the phenomena of Strepsipterous reproduction as
paedogenesis, or pseudo-paedogenesis, but we must agree with
Meinert that they cannot be so classed.

The males of Strepsiptera live for only a very short time, and are
very difficult of observation. According to Hubbard the males of
Xenos dash about so rapidly that the eye cannot see them, and they
create great agitation amongst the wasps in the colonies of which
they are bred. Apparently they are produced in great numbers, and
their life consists of only fifteen or twenty minutes of fiery energy.
The males of Stylops are not exposed to such dangers as those of
Xenos, and apparently live somewhat longer—a day or two, and
even three days are on record. The individuals of Andrena
parasitised by Stylops are apparently greatly affected in their
economy and appear earlier in the season than other individuals; this
perhaps may be a reason, coupled with their short lives, for their
being comparatively rarely met with by entomologists.

Fig. 156.—Abdomen of a wasp (Polistes hebraeus) with a

Strepsipteron (Xenos ♀) in position, one of the dorsal plates of the
wasp's abdomen being removed. a, Projection of part of the
parasite; b, line indicating the position of the removed dorsal
plate.

It is not possible at present to form a valid opinion as to whether

Stylopidae are a division of Coleoptera or a separate Order. Von
Siebold considered them a distinct Order, and Nassonoff, who has
recently discussed the question, is also of that opinion.
 CHAPTER VI

LEPIDOPTERA—OR BUTTERFLIES AND MOTHS

Order VI. Lepidoptera.

Wings four; body and wings covered with scales usually

variegate in colour, and on the body frequently more or less like
hair: nervures moderate in number, at the periphery of one wing
not exceeding fifteen, but little irregular; cross-nervules not more
than four, there being usually only one or two closed cells on
each wing, occasionally none. Imago with mouth incapable of
biting, usually forming a long coiled proboscis capable of
protrusion. Metamorphosis great and abrupt; the wings
developed inside the body; the larva with large or moderate
head and strong mandibles. Pupa with the appendages usually
adpressed and cemented to the body so that it presents a more
or less even, horny exterior, occasionally varied by projections
that are not the appendages and that may make the form very
irregular: in many of the smaller forms the appendages are only
imperfectly cemented to the body.

Lepidoptera, or butterflies and moths, are so far as ornament is

concerned the highest of the Insect world. In respect of intelligence
the Order is inferior to the Hymenoptera, in the mechanical
adaptation of the parts of the body it is inferior to Coleoptera, and in
perfection of metamorphosis it is second to Diptera. The mouth of
Lepidoptera is quite peculiar; the proboscis—the part of the
apparatus for the prehension of food—is anatomically very different
from the proboscis of the other Insects that suck, and finds its
nearest analogue in the extreme elongation of the maxillae of certain
Coleoptera, e.g. Nemognatha. The female has no gonapophyses,
though in certain exceptional forms of Tineidae, there are
modifications of structure connected with the terminal segments, that
have as yet been only imperfectly investigated. As a rule, the egg is
simply deposited on some living vegetable and fastened thereto.
Lepidoptera are the most exclusively vegetarian of all the Orders of
Insects; a certain number of their larvae prey on Insects that are
themselves filled with vegetable juices (Coccidae, Aphidae) and a
very small number (Tinea, etc.) eat animal matter. In general the
nutriment appears to be drawn exclusively from the fluids of the
vegetables, the solid matter passing from the alimentary canal in
large quantity in the form of little pellets usually dry, and called frass.
Hence the quantity of food ingested is large, and when the
individuals unduly increase in number, forest trees over large areas
are sometimes completely defoliated by the caterpillars.

Fig. 157.—Metamorphosis of a Lepidopteron (Rhegmatophila alpina,

Notodontidae). (After Poujade, Ann. Soc. ent. France, 1891.)
Europe. A, Egg; B, young larva, about to moult; C, adult larva; D,
head and first body-segment of adult larva, magnified; E, pupa, ×
2⁄1; F, male moth in repose; G, female moth in repose.

Lepidoptera pass a larger portion of their lives in the pupal stage

than most other Insects do; frequently during nine months of the year
the Lepidopteron may be a pupa. In other Orders of Insects it would
appear that the tendency of the higher forms is to shorten the pupal
period, and when much time has to be passed between the end of
the feeding up of the larva and the appearance of the imago, to pass
this time as much as possible in the form of a resting-larva, and as
little as may be in the form of a pupa; in Lepidoptera the reverse is
the case; the resting-larva period being usually reduced to a day or
two. Hence we can understand the importance of a hard skin to the
pupa. There are, however, numerous Lepidopterous pupae where
the skin does not attain the condition of hardness that is secured for
the higher forms by the chitinous exudation we have mentioned; and
there are also cases where there is a prolonged resting-larva period:
for instance Galleria mellonella spins a cocoon in the autumn and
remains in it as a resting larva all the winter, becoming a pupa only in
the spring. In many of these cases the resting-larva is protected by a
cocoon. It is probable that the chief advantage of the perfect
chitinous exudation of the Lepidopterous pupa is to prevent the tiny,
complex organisation from the effects of undue transpiration.
Bataillon has suggested that the relation of the fluid contents of the
pupa to air and moisture are of great importance in the physiology of
metamorphosis.

The duration of life is very different in various forms of Lepidoptera. It

is known that certain species (Ephestia kuehniella, e.g.) may go
through at least five generations a year. On the other hand, certain
species that feed on wood or roots may take three years to complete
their life-history; and it is probable that some of the forms of
Hepialidae are even longer lived than this.

Lepidoptera have always been a favourite Order with entomologists,

but no good list of the species has ever been made, and it would be
a difficult matter to say how many species are at present known, but
it can scarcely be less than 50,000. In Britain we have about 2000
species.

The close affinity of the Order with Trichoptera has long been
recognised: Réaumur considered the latter to be practically
Lepidoptera with aquatic habits, and Speyer pointed out the
existence of very numerous points of similarity between the two.
Brauer emphasised the existence of mandibles in the nymph of
Trichoptera as an important distinction: the pupa of Micropteryx (Fig.
211) has however been recently shown to be similar to that of
Trichoptera, so that unless it should be decided to transfer
Micropteryx to Trichoptera, and then define Lepidoptera and
Trichoptera as distinguished by the condition of the pupa, it would
appear to be very difficult to retain the two groups as distinct.

Structure of Imago.—The head of a Lepidopteron is in large part

made up of the compound eyes; in addition to these it frequently
bears at the top a pair of small, simple eyes so much concealed by
the scales as to cause us to wonder if seeing be carried on by them.
The larger part of the front of the head is formed by the clypeus,
which is separated by a well-marked line from the epicranium, the
antennae being inserted on the latter near its point of junction with
the former. There is sometimes (Saturnia, Castnia) on each side of
the clypeus a deep pocket projecting into the head-cavity. The other
parts of the head are but small. The occipital foramen is very large.
[164]

Fig. 158—External structure of a female butterfly, Anosia plexippus.

(After Scudder.) a, Base of antenna; b, pronotum; b2, scutum of
mesothorax; c, clypeus; cx, coxa; d, scutellum; d1, scutellum of
metathorax; e, post-scutellum (= base of phragma); em,
epimeron; ep, episternum; f, scutum of metathorax; m, basal part
of proboscis (= maxilla); o, eye; p, labial palp; r, mesosternum; s,
prothoracic spiracle; t, tegula; tr, trochanter; 1-9, dorsal plates of
abdomen.

The antennae are always conspicuous, and are very various in form;
they are composed of numerous segments, and in the males of
many species attain a very complex structure, especially in
Bombyces and Psychidae; they doubtless function in such cases as
sense-organs for the discovery of the female.

The largest and most important of the mouth-parts are the maxillae
and the labial palpi, the other parts being so small as to render their
detection difficult. The labrum is a very short, comparatively broad
piece, visible on the front edge of the clypeus; its lateral part usually
forms a prominence which has often been mistaken for a mandible;
Kellogg has applied the term "pilifer" to this part. In the middle of the
labrum a small angular or tongue-like projection is seen just over the
middle of the base of the proboscis; this little piece is considered by
several authorities to be an epipharynx.

Fig. 159—Mouth of Lepidoptera. Tiger-moth, Arctia caja. A, Seen from

front; B, from front and below, a, Clypeus; b, labrum; c,
epipharynx; d, mandibular area; d′, prominence beneath
mandibular area; e, one side of haustellum or proboscis; f,
maxillary palp; g, labial palp.

Mandibles.—Savigny, Westwood, and others considered the parts

of the labrum recently designated pilifers by Kellogg to be the
rudimentary mandibles, but Walter has shown that this is not the
case.[165] The mandibles are usually indistinguishable, though they,
or some prominence possibly connected with them,[166] may
frequently be detected in the neighbourhood of the pilifers; they are,
according to Walter, largest and most perfectly developed in
Eriocephala, a genus that was not distinguished by him from
Micropteryx and was therefore termed "niedere Micropteryginen," i.e.
lower Micropteryges. The opinion entertained by Walter that
Micropteryx proper (his "höhere Micropteryginen") also possesses
rudimentary mandibles is considered by Dr. Chapman, no doubt with
reason, to be erroneous.[167] The mandibles, however, in the vast
majority of Lepidoptera can scarcely be said to exist at all in the
imago; there being only an obtuse projection—without trace of
articulation—on each side of the labrum; and even this projection is
usually absent. Meinert recognised these projections as mandibles in
Smerinthus populi, and Kellogg in Protoparce carolina, another large
Sphinx moth. They appear to be unusually well developed in that
group. In Castnia they are even more definite than they are in
Sphingidae.

The Maxillae are chiefly devoted to the formation of the proboscis.

Their basal portions are anatomically very indefinite, though they
exist very intimately connected with the labium. Each usually bears a
small tubercle or a segmented process, the representative of the
maxillary palpus. The proboscis itself consists of the terminal, or
outer, parts of the two maxillae, which parts are closely and
beautifully coadapted to form the spirally coiled organ, that is
sometimes, though incorrectly, called the tongue. The exact
morphology of the Lepidopterous proboscis has not been
established. The condition existing in the curious family Prodoxidae
(see p. 432), where a proboscis coexists with another structure
called a maxillary tentacle, suggests a correspondence between the
latter and the galea of a typical maxilla; and between the proboscis
and the lacinia or inner lobe of a maxilla: but J. B. Smith is of opinion
that the tentacle in question is a prolongation of the stipes. The
condition of the parts in this anomalous family (Prodoxidae) has not,
however, been thoroughly investigated, and Packard takes a
different view of the proboscis; he considers that "it is the two galeae
which become elongated, united and highly specialised to form the
so-called tongue or glossa of all Lepidoptera above the
Eriocephalidae."[168] The proboscis in some cases becomes very
remarkable, and in certain Sphingidae is said to attain, when
unrolled, a length of ten inches. In some cases the maxillary lobes
do not form a proboscis, but exist as delicate structures, pendulous
from the mouth, without coadaptation (Zeuzera aesculi, the Wood-
leopard moth). In other forms they are absent altogether (Cossus,
e.g.), and in Hepialus we have failed to detect any evidence of the
existence of the maxillae. On the other hand, in Micropteryx the
maxillae are much more like those of a mandibulate Insect; and
various other Microlepidoptera approach more or less a similar
condition. In the genus last mentioned the maxillary palpi are largely
developed, flexible and slender. According to Walter various forms of
palpus intermediate between that of Micropteryx and the condition of
rudimentary tubercle may be found amongst the Microlepidoptera.
[169]

Labium.—The labial palpi are usually largely developed, though but

little flexible; they form conspicuous processes densely covered with
scales or hairs, and curve forwards or upwards, rarely downwards,
from the under side of the head, somewhat in the fashion of tusks.
The other parts of the labium are frequently represented merely by a
membranous structure, united with the maxillae and obstructing the
cavity of the pharynx. Where the proboscis is absent it is difficult to
find any orifice leading to the alimentary canal, such opening as may
exist being concealed by the overhanging clypeus and labium. In
some forms, Saturnia, e.g., there appears to be no buccal orifice
whatever. In Hepialus the labium is in a very unusual condition; it
projects externally in the position usually occupied by the labial palpi,
these organs being themselves extremely short. It is very difficult to
form an opinion as to the structure of the labium and other mouth-
parts when the maxillae are not developed, as in these cases the
parts are of a delicate membranous nature, and shrivel after death.
This is the explanation of the fact that in descriptive works we find
vague terms in use such as "mouth aborted" or "tongue absent."

The mouth of the Lepidopterous imago is a paradoxical structure; it

differs very greatly from that of the larva, the changes during
metamorphosis being extreme. We should thus be led to infer that it
is of great importance to the creatures; but, on the other hand, the
various structures that make up the mouth, as we have remarked,
are frequently absent or reduced to insignificant proportions; and
even in forms where the apparatus is highly developed the
individuals seem to be able to accomplish oviposition without taking
food, or after taking only very minute quantities. It is therefore difficult
to understand why so great a change should occur during the
metamorphosis of the Insects of this Order. It has been ascertained
that in some forms where the mouth is atrophied the stomach is in a
correlative condition; but we are not aware that any investigations
have been made as to whether this correspondence is general or
exceptional.

The exact mode in which the proboscis acts is in several respects

still obscure, the views of Burmeister and Newport being in some
points erroneous. Towards the tip of the proboscis there are some
minute but complex structures considered by Fritz Müller to be
sense-organs, and by Breitenbach to be mechanical instruments for
irritating or lacerating the delicate tissues of blossoms. It is probable
that Müller's view will prove to be correct. Nevertheless the
proboscis has considerable power of penetration; there being a
moth, "Ophideres fullonica" that causes considerable damage to
crops of oranges by inserting its trunk through the peel so as to suck
the juices.[170] The canal formed by each maxilla opens into a cavity
inside the front part of the head. This cavity, according to Burgess,
[171] is a sort of sac connected with five muscles, and by the aid of
this apparatus the act of suction is performed: the diverticulum of the
alimentary canal, usually called a sucking-stomach, not really
possessing the function formerly attributed to it.

The Prothorax is very small, being reduced to a collar, between

the head and the alitrunk, of just sufficient size to bear the front pair
of legs. Its most remarkable feature is a pair of processes, frequently
existing on the upper surface, called "patagia." These in many cases
(especially in Noctuidae) are lobes capable of considerable
movement, being attached only by a narrow base. In Hepialus, on
the contrary, they are not free, but are merely indicated by curved
marks on the dorsum. The patagia are styled by many writers
"tegulae." They are of some interest in connection with the question
of wing-like appendages on the prothorax of Palaeozoic insects, and
they have been considered by some writers[172] to be the
equivalents of true wings. The Mesothorax is very large, especially
its upper face, the notum, which is more or less convex, and in the
higher forms attains a great extension from before backwards. The
notum consists in greater part of a large anterior piece, the meso-
scutum, and a smaller part, the meso-scutellum behind. In front of
the scutum there is a piece termed prae-scutum by Burgess. It is
usually small and concealed by the front part of the scutum; but in
Hepialus it is large and horizontal in position. It is of importance as
being the chief point of articulation with the prothorax. The scutellum
is more or less irregularly rhomboidal in form; its hinder margin
usually looks as if it were a lobe or fold placed in front of the base of
the abdomen or metathorax, according to whether the latter is
concealed or visible. In some of the higher forms this meso-scutellar
lobe is prominent, and there may be seen under its projection a
piece that has been called the post-scutellum, and is really the base
of the great mesophragma, a chitinous piece that descends far down
into the interior of the body. In addition to the front pair of wings the
mesothorax bears on its upper surface another pair of appendages,
the tegulae: in the higher forms they are of large size; they are
fastened on the front of the mesothorax, and extend backwards over
the joint of the wing with the body, being densely covered with scales
so that they are but little conspicuous. These appendages are
frequently erroneously called patagia, but have also been called
scapulae, pterygodes, paraptera, and shoulder-tufts, or shoulder-
lappets. The lower surface of the mesothorax is much concealed by
the large and prominent coxae, but the sternum and the two pleural
pieces on each side, episternum and epimeron, are easily detected.
The area for attachment of the anterior wing on each side is
considerable, and appears to be of rather complex structure; its
anatomy has been, however, but little studied.

The Metathorax is small in comparison with the preceding

segment, to which it is intimately co-adapted, though the two are
really connected only by delicate membrane, and can consequently
be separated with ease by dissection. The metanotum consists of (1)
the scutum, which usually appears externally as an anterior piece on
each side; (2) the scutellum, forming a median piece placed behind
the scutum, which it tends to separate into two parts by its own
extension forwards. In order to understand the structure of the
metathorax it is desirable to dissect it off from the larger anterior
segment, and it will then be found that its appearance when
undissected is deceptive, owing to its being greatly arched, or folded
in the antero-posterior direction. A broad, but short phragma
descends from the hind margin of the metascutellum into the interior
of the body. It should be noted that though the metanotum is forced,
as it were, backwards by the great extension of the mesonotum in
the middle line of the body, yet at the sides the metanotum creeps
forward so as to keep the points of attachment of the hind wings
near to those of the front wings. In many forms of Hesperiidae,
Sphingidae, Noctuidae, etc. the true structure of the metanotum is
further concealed by the back of the mesoscutellum reposing on,
and covering it.

Difference of opinion exists as to the thoracic Spiracles; there is one

conspicuous enough in the membrane behind the pronotum, and it is
thought by some writers that no other exists. Westwood and
Scudder, however, speak of a mesothoracic spiracle, and Dr.
Chapman considers that one exists. Minot describes[173] a structure
behind the anterior wing, and thinks it may be an imperfect spiracle,
and we have found a similar stigma in Saturnia pavonia. At the back
of the thorax there is on each side in some Lepidoptera (Noctuidae,
Arctia, etc.), a curious large cavity formed by a projection backwards
from the sides of the metasternum, and a corresponding
development of the pleura of the first abdominal segment. Minot and
others have suggested that this may be an organ of hearing.

The Abdomen differs according to the sex. In the female seven

segments are conspicuous dorsally, but only six ventrally, because
the first segment is entirely membranous beneath, and is concealed
between the second abdominal ventral plate and the posterior
coxae. Besides these segments there are at the hind end two others
smaller, more or less completely withdrawn into the body, and in
certain cases forming an ovipositor. These nine segments are
usually considered to constitute the abdomen; but according to
Peytoureau,[174] a tenth dorsal plate is represented on either side of
the anal orifice, though there is no trace of a corresponding ventral
plate. In the male the segments, externally conspicuous, are one
more than in the female. According to the authority quoted,[175] this
sex has also truly ten abdominal segments, the ninth segment being
withdrawn to a greater or less extent to the inside of the body, and
modified to form part of a copulatory apparatus; its dorsal portion
bears a process called the "uncus"; the anal orifice opens on the
inner face of this process, and below it there is another process—
developed to a greater or less extent—called the "scaphium." The
ventral portion of the ninth segment bears a lobe, the "saccus"
(Peytoureau, l.c.). On each side of the ninth abdominal segment
there is a process called the "valve," the internal wall of which bears
some hook-like or other processes called "harpes"; it is continued as
a membrane surrounding the "oedeagus," or penis, and—bearing
more or less distinct prominences—connects with the scaphium. In
many forms the parts alluded to, other than the valves, are
concealed by the latter, which come together when closed, and may
be covered externally with scales like the rest of the abdomen.
Peytoureau considers that the uncus is really the dorsal plate of a
tenth segment, and that the scaphium is the tenth ventral plate.
Thus, according to this view, the ninth segment is extensive and
complex, being very highly modified in all its parts: while the tenth
segment is greatly reduced. The structure of the male organs is
simpler in Lepidoptera, and less varied than it is in the other great
Orders of Insects. There are seven pairs of abdominal spiracles on
the upper parts of the membranous pleurae.

Fig. 160—Acherontia atropos. The termination of ♂ body, one side

removed. IX, Ninth dorsal plate; IX’, ninth ventral; s, lobe, saccus,
of ninth ventral plate; X, tenth dorsal plate, or uncus; sc,
scaphium, or tenth ventral plate; a, position of anus; b, chitinised
band of scaphium; V, valve or clasper; c, hooks, or harpes, of
clasper; p, penis (or oedeagus). (After Peytoureau.)

Legs.—The legs are long, slender, covered with scales, and chiefly
remarkable from the fact that the tibiae sometimes bear articulated
spurs on their middle as well as at the tip. The front tibia usually
possesses on its inner aspect a peculiar mobile pad; this seems to
be in some cases a combing organ; it also often acts as a cover to
peculiar scales. The tarsi are five-jointed, with two small claws and a
small apparatus, the functional importance of which is unknown,
between the claws.

Wings.—The wings are the most remarkable feature of this Order; it

is to them that butterflies owe their beauty, the surfaces of the wings
being frequently adorned with colours and patterns of the most
charming and effective nature. These effects are due to minute
scales that are implanted in the wing-membrane in an overlapping
manner, somewhat similar to the arrangement of slates on the roof of
a house. The scales are very readily displaced, and have the
appearance of a silky dust. We shall describe their structure and
allude to their development subsequently. The wings are usually of
large size in comparison with the Insect's body: in the genus
Morpho, the most gorgeous of the butterflies, they are enormous,
though the body is small; so that when deprived of these floats the
Insect is insignificant. The great expanse of wing is not correlative
with great powers of flight, though it is perhaps indicative of flying
with little exertion; for the small-winged Lepidoptera, Sphingidae,
etc., have much greater powers of aërial evolution than the large-
winged forms. The area of the wing is increased somewhat by the
fact that the scales on the outer margin, and on a part or on the
whole of the inner margin, project beyond the edges of the
membrane that bears them: these projecting marginal scales are
called fringes. In many of the very small moths the actual size of the
wing-membranes is much reduced, but in such cases the fringes
may be very long, so as to form the larger part of the surface,
especially of that of the hind wings. Frequently the hind wings are of
remarkable shape, being prolonged into processes or tails, some of
which are almost as remarkable as those of Nemoptera in the Order
Neuroptera.
The wings are very rarely absent in Lepidoptera; this occurs only in
the female sex, no male Lepidopterous imago destitute of wings
having been discovered. Although but little is known of the
physiology of flight of Lepidoptera, yet it is clearly important that the
two wings of the same side should be perfectly coadapted or
correlated. This is effected largely by the front wing overlapping the
hind one to a considerable extent, and by the two contiguous
surfaces being pressed, as it were, together. This is the system
found in butterflies and in some of the large moths, such as
Lasiocampidae and Saturniidae; in these cases the hind wing always
has a large shoulder, or area, anterior to its point of insertion. In most
moths this shoulder is absent, but in its place there are one or more
stiff bristles projecting forwards and outwards, and passing under a
little membranous flap, or a tuft of thick scales on the under face of
the front wing; the bristle is called the "frenulum," the structure that
retains it a "retinaculum." In Castnia (Fig. 162) and in some
Sphingidae there is the unusual condition of a highly-developed
shoulder (s) coexisting with a perfect frenulum (f) and retinaculum
(r). The frenulum and retinaculum usually differ in structure, and the
retinaculum in position, in the two sexes of the same moth; the male,
which in moths has superior powers of flight, having the better
retaining organs. Hampson says "the form of the frenulum is of great
use in determining sex, as in the males of all the forms that possess
it, it consists of hairs firmly soldered together so as to form a single
bristle, whilst in nearly all females it consists of three or more bristles
which are shorter than that of the male; in one female Cossid I have
found as many as nine. Also in the large majority of moths the
retinaculum descends from the costal nervure in the male, while in
the female it ascends from the median nervure."[176] This sexual
difference in a structure for the discharge of a function common to
the two sexes is a very remarkable fact. There are a few—very few
—moths in which the bases of the hind wings are not well coadapted
with the front wings, and do not possess a frenulum, and these
species possess a small more or less free lobe at the base of the
front wing that droops towards the hind wing, and may thus help to
keep up an imperfect connexion between the pair; this lobe has been
named a jugum by Professor Comstock. Occasionally there is a
jugum on the hind as well as on the front wing. There is usually a
very great difference between the front and the hind wings; for
whereas in the front wing the anterior portion is doubtless of great
importance in the act of flight and is provided with numerous veins,
in the hind wing, on the other hand, the corresponding part has not a
similar function, being covered by the front wing; hence the hind
wing is provided with fewer nervures in the anterior region, the
divisions of the subcostal being less numerous than they are in the
front wing. In the moths possessing a jugum the two wings differ but
little from one another, and it is probable that they function almost as
four separate wings instead of as two pairs.

Wing-nervures.—The nervures or ribs of the wings are of great

importance in Lepidoptera, as at present they furnish the chief
characters for classification and for the discussions of phylogeny that
are so numerous in entomological literature. On looking at wings that
have been deprived of their scales it will be noticed (Fig. 161) that
the ribs are much more numerous at the outer margins than they are
near the points of attachment of the wings, and that there is usually
but one cell (or area completely enclosed by ribs). This latter point is
one of the chief peculiarities of the Lepidopterous wing; in Insect-
wings generally the number of cells in proportion to the area of the
wings and to the number of nervures is greater than it is in
Lepidoptera, for in the latter there are few or no cross-nervures.
Hence there is sometimes no closed cell at all on the wing (Fig. 161,
II. B). The maximum number of closed cells is six; this is found in
some species of Micropteryx, while in Hepialus there may be three
or four; but the rule is that there is only one cell in the Lepidopterous
wing. When the number of cells is increased this is not necessarily
due to an increase in the cross-nervures; and in fact it is generally
due to irregular forking or to the sinuous form of the longitudinal
nervures themselves (see wing of Castnia, Fig. 162, A.). Some
authorities consider that all transverse or cross-veins in Lepidoptera
are merely portions of longitudinal veins having diverted courses.
When a portion of a nervure beyond the basal or primary portion
serves as a common piece to two forked parts external to it, it is
called a stalk (Fig. 162, A, e). There are cases in which the furcation
takes place in the opposite direction, so that a nervure is double at
the base of the wing (Fig. 161, I, A, 1a, and B, 1b). This important
condition has not yet been adequately discussed.

Fig. 161.—Wing-nervuration of Lepidoptera. I, Diagram of moths' wings

(after Hampson); II, of a butterfly's wings (Morpho menelaus ♂ ,
after Staudinger and Schatz). A, front, B, hind wing. I.—c, costal;
sc, subcostal; m, median; 1a, 1b, 1c, internal nervures; f,
frenulum; 2, 3, 4, branches of median nervure; 5, lower radial; 6,
upper radial; 7-11, divisions of the subcostal; 12, termination of
costal; c, cell; d, discocellular nervure. II.—C, costal; SC,
subcostal; M, median; SM and SN, submedian nervures; 1A,
inner-margin nervure; UR, lower radial; OR, upper radial; SC1 to
SC5, divisions of subcostal; M1 to M3, divisions of median nervure;
C, cell; DC, discocellulars.

Turning to the mode of designation of the nervures,[177] we may

commence by remarking that no system satisfactory from a practical
as well as from a theoretical point of view has yet been devised. The
diagrams given in figure 161 will enable us to explain the methods
actually in vogue; I. representing the system, dating from the time of
Herrich-Schaeffer, chiefly used by British naturalists, and II. that
adopted by Staudinger and Schatz in their recent great work on the
Butterflies of the world. The three anterior nervures in both front and
hind wings correspond fairly well, and are called, looking at them
where they commence at the base of the wing, "costal," "subcostal,"
and "median" nervures. The nervures near the inner margin of the
wing (that is the lower part in our figures) differ much in the front and
hind wings, consisting either of two or of three separate portions not
joined even at the base. British entomologists call these "branches
or divisions of the internal nervure": the Germans call the more
anterior of them the "submedian," and the more internal the "inner-
margin nervure"; they are also frequently called anal nervures. The
cross-nervure that closes the cell is called discocellular; when
apparently composed of two or three parts joined so as to form
angles, the parts are called, according to position, upper, lower, and
middle discocellulars. One or more short spurs may exist on the front
part of the basal portion of the hind wing; these are called
praecostal. The branches or terminal divisions of the nervures
should be called nervules; they are usually mentioned by the
numbers shewn in the diagram (Fig. 161, I.). In addition to this, it is
only necessary to remember that number 2 is always assigned to the
posterior division of the median nervure, the nervules below this
being all called 1, and distinguished by the addition of a, b, c when
requisite. This course is necessary, because if it were not adopted
the corresponding nervules on the front and hind wings would bear
different numbers.

The use of this system of numbers for the nervules is becoming

general, and it answers fairly well for practical purposes. On the
other hand, extreme discrepancy exists as to the nomenclature of
the nervures and nervules, and there are almost as many systems
as there are authorities.

The normal number of nervules is, on the front wing, 11 + 1 or 2

inner marginal, and on the hind wing 7 + 2 or 3 inner marginal. In the
aberrant moths of the genus Castnia the nervuration is unusually
complex and irregular (Fig. 162), and an analogous condition occurs
in our common Goat-moth (Cossus ligniperda). In Hepialus and
Micropteryx (the jugate moths of Comstock) the hind wings are less
dissimilar in nervuration from the front wings than they are in other
Lepidoptera.[178]