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Sílvio Vaz Jr. Editor

Sustainable
Agrochemistry
A Compendium of Technologies
Sustainable Agrochemistry
Sílvio Vaz Jr.
Editor

Sustainable Agrochemistry
A Compendium of Technologies

123
Editor
Sílvio Vaz Jr.
National Research Center for Agroenergy
Brazilian Agricultural Research Corporation
Brasilia, Brazil

ISBN 978-3-030-17890-1 ISBN 978-3-030-17891-8 (eBook)


https://doi.org/10.1007/978-3-030-17891-8
© Springer Nature Switzerland AG 2019
This work is subject to copyright. All rights are reserved by the Publisher, whether the whole or part
of the material is concerned, specifically the rights of translation, reprinting, reuse of illustrations,
recitation, broadcasting, reproduction on microfilms or in any other physical way, and transmission
or information storage and retrieval, electronic adaptation, computer software, or by similar or dissimilar
methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this
publication does not imply, even in the absence of a specific statement, that such names are exempt from
the relevant protective laws and regulations and therefore free for general use.
The publisher, the authors and the editors are safe to assume that the advice and information in this
book are believed to be true and accurate at the date of publication. Neither the publisher nor the
authors or the editors give a warranty, expressed or implied, with respect to the material contained
herein or for any errors or omissions that may have been made. The publisher remains neutral with regard
to jurisdictional claims in published maps and institutional affiliations.

This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
I would like to dedicate this book to my wife
Ana and to my daughter Elena. They are my
source of inspiration.
Preface

Agrochemistry was seen during many years as a source of environmental and health
concerns due to, mainly, the pesticides used in agriculture around the world.
Conventional pesticides produced negative impacts on the environment as the biota
pollution and destruction near to the crop systems; furthermore, agricultural
workers were contaminated by these agrochemicals, causing serious illness and
death.
Nowadays, the demand from the modern society for sustainable production of
food has promoted the development of a sustainable agrochemistry considering
aspects such as reducing negative impacts on the environment- and health-friendly
materials and molecules, bioactive compounds, etc.
Agriculture remains one of the most strategic sectors for the global economy and
well-being. In this way, innovations and new paradigms are necessary for their
sustainable exploitation, and agrochemistry can certainly continue to contribute to
the generation of agricultural wealth for modern society.
This book intends to present a large variety of technologies for a sustainable
agrochemistry, such as semiochemicals for pest management, nanomaterials, green
chemistry principles for agriculture, among others, with their respective case study.
Chapter 1 defines sustainable agrochemistry, the main classes of agrochemicals
as well as their usages and dynamic in the agriculture and the environment.
Furthermore, some relevant aspects of sustainability in agriculture are presented and
discussed. Chapter 2 presents historical aspects of crop protection and the use of
pesticides to guarantee sustainable food supplies. Chapter 3 presents the principles
of semiochemical use for monitoring and controlling pests and the way in which
these natural molecules work is presented and discussed. Chapter 4 describes the
major concepts related to nanoscience and nanotechnology, role of green nan-
otechnology as an essential part of a sustainable future of agriculture, and its
applicability for the development of innovative solutions to challenging issues.
Chapter 5 deals with the use of magnetic resonance techniques to improve agri-
cultural systems, highlighting the obtaining of structural information about indus-
trial biomass and soil organic matter for scientific and technological usages.
Chapter 6 talks about chemical analyses and their important role in agriculture, as

vii
viii Preface

supporting technologies at all stages of agro-industrial chains as grains, forests,


pulp and paper, waste and agricultural residues, among others sources of agricul-
tural products; furthermore, a set of relevant analytical techniques are discussed in
accordance with their application in the agriculture. Chapter 7 treats about the
recent acknowledgement of the supramolecular nature of soil humic substances,
that allowed to devise a fractionation procedure, called Humeomics, that enables
detailed characterization of the structure of humic molecules in soil for their
application in the improvement of soil functionality. Chapter 8 addresses some
general aspects about the chemistry action of crop protection products against pest
attack and the implications of these agrochemicals on the environment in order to
produce food sustainably for a constantly growing world population. Chapter 9
deals with the more relevant information about the toxicity of agrochemicals for the
biota and the human health. From this, the understanding of the impact from
agrochemicals use. Chapter 10 presents and discusses the green chemistry princi-
ples, highlighting their application in agriculture. Moreover, from this, the under-
standing of methods to change the development and production of green
agrochemicals. Chapter 11 deals with the more relevant tools for the ecological risk
assessment of agrochemicals, what clarify the paramount importance of monitoring
and control of agrochemicals in the environment using mathematical models.
Finally, Chap. 12 deals with the more relevant strategies for the management of
agrochemical residues in soil and water. Furthermore, the most advanced treatment
technologies will be explored.
The presentation of case studies aims to expand the reader’s knowledge and
demonstrate examples of application of the technologies proposed by the authors.
Good lecture!

Acknowledgements The editor would like to thank all the authors for the efforts to prepare a
high-quality text—this is not a simple task. Thanks also to Springer team for enabling the trans-
formation of an idea into a book. Finally, thanks to colleagues from Embrapa Agroenergy for
refining and improving some parts of the manuscript.

Brasilia, Brazil Sílvio Vaz Jr.


Contents

1 Introduction to Sustainable Agrochemistry . . . . . . . . . . . . . . . . . . . 1


Sílvio Vaz Jr.
2 The Battle for a Sustainable Food Supply . . . . . . . . . . . . . . . . . . . 13
John Unsworth, Yoshiaki Nakagawa, Caroline Harris and Gijs Kleter
3 Semiochemicals for Integrated Pest Management . . . . . . . . . . . . . . 85
Maria C. Blassioli-Moraes, Raúl A. Laumann, Mirian F. F. Michereff
and Miguel Borges
4 Green Nanotechnology for Sustained Release of Eco-Friendly
Agrochemicals . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 113
Luciano Paulino Silva and Cínthia Caetano Bonatto
5 Magnetic Resonance Spectroscopy Techniques to Improve
Agricultural Systems . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 131
Sílvio Vaz Jr., Etelvino Henrique Novotny and Luiz Alberto Colnago
6 Chemical Analyses for Agriculture . . . . . . . . . . . . . . . . . . . . . . . . . 147
Sílvio Vaz Jr.
7 The Soil Humeome: Chemical Structure, Functions
and Technological Perspectives . . . . . . . . . . . . . . . . . . . . . . . . . . . . 183
Alessandro Piccolo, Riccardo Spaccini, Davide Savy, Marios Drosos
and Vincenza Cozzolino
8 Synthesis of New Agrochemicals . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
Paulo Marcos Donate and Daniel Frederico
9 Toxicological Aspects of Pesticides . . . . . . . . . . . . . . . . . . . . . . . . . 275
Eloisa Dutra Caldas
10 Green Chemistry and Agrochemistry . . . . . . . . . . . . . . . . . . . . . . . 307
Sílvio Vaz Jr.

ix
x Contents

11 Exposure Characterization Tools for Ecological Risk Assessment


of Pesticides in Water . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 321
Claudio A. Spadotto and Rafael Mingoti
12 Management of Agrochemical Residues in the Environment . . . . . 361
Sílvio Vaz Jr. and Luciano Gebler
Contributors

Maria C. Blassioli-Moraes Brazilian Agricultural Research Corporation, National


Research Center for Genetic Resources and Biotechnology—Laboratory for
Semiochemicals, Brasilia, Distrito Federal, Brazil
Cínthia Caetano Bonatto Brazilian Agricultural Research Corporation,
Laboratory of Nanobiotechnology (LNANO), Embrapa Genetic Resources and
Biotechnology, Brasilia, Distrito Federal, Brazil;
NanoDiversity, Applied Research, TecSinapse, São Paulo, SP, Brazil
Miguel Borges Brazilian Agricultural Research Corporation, National Research
Center for Genetic Resources and Biotechnology—Laboratory for Semiochemicals,
Brasilia, Distrito Federal, Brazil
Eloisa Dutra Caldas Department of Pharmacy, Faculty of Health Sciences,
University of Brasilia, Brasilia, Distrito Federal, Brazil
Luiz Alberto Colnago Brazilian Agricultural Research Corporation, Embrapa
Instrumentation, São Carlos, SP, Brazil
Vincenza Cozzolino Department of Agricultural Sciences, Interdepartmental
Research Centre on Nuclear Magnetic Resonance for the Environment, Agro-Food
and New Materials (CERMANU), University of Napoli Federico II, Portici, Italy
Paulo Marcos Donate Departamento de Química da Faculdade de Filosofia,
Ciências e Letras, Universidade de São Paulo, Ribeirão Preto, SP, Brazil
Marios Drosos Faculty of Biology and Environment, Institute of Resource,
Ecosystem and Environment of Agriculture (IREEA), Nanjing Agricultural
University, Nanjing, China
Daniel Frederico Dinagro Agropecuária Ltda., Ribeirão Preto, SP, Brazil
Luciano Gebler Brazilian Agricultural Research Corporation, National Research
Center for Grape and Wine (Embrapa Grape and Wine), Bento Gonçalves, Rio
Grande do Sul, Brazil

xi
xii Contributors

Caroline Harris Exponent International Limited, Harrogate, North Yorkshire, UK


Gijs Kleter Wageningen Food Safety Research, Part of Wageningen University &
Research, Wageningen, The Netherlands
Raúl A. Laumann Brazilian Agricultural Research Corporation, National
Research Center for Genetic Resources and Biotechnology—Laboratory for
Semiochemicals, Brasilia, Distrito Federal, Brazil
Mirian F. F. Michereff Brazilian Agricultural Research Corporation, National
Research Center for Genetic Resources and Biotechnology—Laboratory for
Semiochemicals, Brasilia, Distrito Federal, Brazil
Rafael Mingoti Brazilian Agricultural Research Corporation, Embrapa Territorial,
Campinas, SP, Brazil
Yoshiaki Nakagawa Graduate School of Agriculture, Kyoto University, Kyoto,
Japan
Etelvino Henrique Novotny Brazilian Agricultural Research Corporation,
Embrapa Soils, Rio de Janeiro, Brazil
Alessandro Piccolo Department of Agricultural Sciences, Interdepartmental
Research Centre on Nuclear Magnetic Resonance for the Environment, Agro-Food
and New Materials (CERMANU), University of Napoli Federico II, Portici, Italy
Davide Savy Plant Biology Laboratory, University of Liège, Gembloux Agro-Bio
Tech, Gembloux, Belgium
Luciano Paulino Silva Brazilian Agricultural Research Corporation, Laboratory
of Nanobiotechnology (LNANO), Embrapa Genetic Resources and Biotechnology,
Brasilia, Distrito Federal, Brazil
Riccardo Spaccini Department of Agricultural Sciences, Interdepartmental
Research Centre on Nuclear Magnetic Resonance for the Environment, Agro-Food
and New Materials (CERMANU), University of Napoli Federico II, Portici, Italy
Claudio A. Spadotto Brazilian Agricultural Research Corporation, Embrapa,
Parque Estação Biológica, Brasilia, Distrito Federal, Brazil
John Unsworth Consultant, Chelmsford, UK
Sílvio Vaz Jr. Brazilian Agricultural Research Corporation, National Research
Center for Agroenergy (Embrapa Agroenergy), Embrapa Agroenergia, Parque
Estação Biológica, Brasilia, Distrito Federal, Brazil
Chapter 1
Introduction to Sustainable
Agrochemistry

Sílvio Vaz Jr.

Abstract This chapter defines sustainable agrochemistry, the main classes of agro-
chemicals as well as their usages and dynamics in the agriculture and the environment.
Furthermore, some relevant aspects of sustainability in agriculture are presented and
discussed.

Keywords Agriculture · Environment · Impacts

1.1 Introduction

The practice of agriculture is one of the oldest activities developed by humans. In


the Neolithic period, the constitution of the first techniques and materials used for
the cultivation of plants and the confinement of animals was the main cause for what
was denominated as the sedentarization of human. It allowed fixing residence in a
given locality, although collection and hunting have long coexisted side by side with
agriculture.
The development of agriculture, therefore, was directly associated with the forma-
tion of the first civilizations, which helps us understand the importance of techniques
and the environment in the process of building societies and their geographical spaces.
In that sense, as these societies further advanced their techniques and technologies,
the more the evolution of agriculture was benefited.
Originally, the practice of farming was developed in the vicinity of large rivers,
notably the Tigris and Euphrates, as well as the Nile, the Ganges and others. Not coin-
cidentally, it was in these localities that the first great known civilizations emerged,
because the practice of agriculture allowed the development of trade to a surplus in
production.

S. Vaz Jr. (B)


Brazilian Agricultural Research Corporation, National Research Center for
Agroenergy (Embrapa Agroenergy), Embrapa Agroenergia, Parque Estação Biológica,
s/n, Av. W3 Norte (final), Brasília, DF 70770-901, Brazil
e-mail: [email protected]

© Springer Nature Switzerland AG 2019 1


S. Vaz Jr. (ed.), Sustainable Agrochemistry,
https://doi.org/10.1007/978-3-030-17891-8_1
2 S. Vaz Jr.

One of the most important moments in the process of agricultural evolution


throughout history was, without doubt, what became known as the Agricultural
Revolution (British Broadcasting Corporation 2017). We can say that, over time,
several Agricultural Revolutions have succeeded, but the main ones occurred after
the Industrial Revolution.
The process of industrialization of societies allowed the transformation of the
geographical space in the rural environment, which occurred thanks to the insertion
of greater technological apparatuses in the agricultural production, allowing a greater
mechanization of the field. This transformation materialized from the supply of inputs
from the industry to agriculture, such as machinery, fertilizers and technical objects
and practices in general.
The influence of the Agricultural Revolution in the world was also directly associ-
ated with the European maritime–colonial expansion, in which the European people
disseminated their different cultures through the world by means of introducing new
crops and novel agricultural practices. It is worth remembering that the interaction
between settlers and colonizers also contributed to the agricultural evolution, inas-
much as previously little known techniques were applied and disseminated, such as
the terracing practised in both ancient China and pre-Columbian civilizations.
As an effect of this revolution, Chemistry starts to have a fundamental role in the
agriculture expansion and technological development by means of fertilizers, pest
controllers, scientific knowledge, among other aspects.
In the twentieth century, more precisely after the Second World War, the evolution
of agriculture reached one of its most important hallmarks, in what became known as
the Green Revolution. It was based on a set of measures and promotion of techniques
based on the introduction of genetic improvements in plants and the evolution of
agricultural production apparatus to expand, above all, food production (Pingali
2012).
The introduction of techniques from the Green Revolution has led to a large-scale
increase in grain and cereal production, significantly reducing the need for food in
various regions of Asia, Africa and Latin America, even though hunger has not been
eradicated, since its existence is not only due to lack of food. The impact on the world
was so wide that the American agronomist Norman Borlaug, considered the “father”
of the Green Revolution, was awarded the Nobel Peace Prize in 1990 (Nobel Prize
2018).
Although the Green Revolution is heavily criticized for its environmental impacts
and the process of land concentration that accompanied its evolution due to policies
that were used to promote rapid intensification of agricultural systems and increase
food supplies (Pingali 2012), its importance for the development of agriculture in
the world is undeniable. Furthermore, in the following decades, the improvements
resulting from technology in the field, such as biotechnology, have increased in the
following decades, which has been further increasing the productivity.
Figure 1.1 depicts a cultivated area with corn (Zea mays L.) in the Midwest
Brazilian region. This is a region typically affected by the Green Revolution. The
Brazilian Savannah was an inhospitable and low productive land during a couple
1 Introduction to Sustainable Agrochemistry 3

Fig. 1.1 Corn cropping in the Midwest region of Brazil, one of the largest producing regions of
cereal in the world. Courtesy of Climatempo, São Paulo

Table 1.1 Global production Crop Production, in thousand tonnes


of the five major crops,
according to Food and Sugarcane 1,877,110
Agriculture Organization of Maize (corn) 1,016,740
the United Nations (2015)
Rice 745,710
Wheat 713,183
Potatoes 368,096

of centuries, but, nowadays, it is responsible for Brazil to be the major agricultural


player in South America, and one biggest worldwide.
Table 1.1 describes the main crops cultivated around the world and their produc-
tion. These values would not hardly be achieved without the use of agrochemicals,
brought to the scene by the Green Revolution.
Nowadays, agriculture is constantly required to become more sustainable, with
reduction of its negative impacts on environment allied to an increasing in their
positive impacts on society and economy. These are challenges and, at same time,
opportunities for new production systems.
4 S. Vaz Jr.

1.2 Chemistry and Agriculture

Chemistry is a science closely related to agriculture since its origin. The term agri-
cultural chemistry rises with the publication of Elements of Agricultural Chemistry
by the chemist Humphry Davy (Fig. 1.2) in 1813 in England (Encyclopaedia Bri-
tannica 2018). The book was traduced to French, Italian and German. It was also
published with great prestige in U.S. because it boosted the American agricultural
community, which was based on efficient techniques that had taken into account
scientific principles and observations.
Nowadays, we can define agrochemistry as the application of chemistry in agricul-
ture. Its action, object of study and technical means are not only concerned with the
production of agrochemicals but also with the analysis and prevention of the harmful
effects of chemical substances on both crops and humans (farmers and consumers),
and their impacts on the environment.
The contribution of chemistry goes back to the nineteenth century, with the syn-
thesis of inorganic fertilizers and, by the middle of the last century, of a great number
of compounds synthesized to control insects, diseases and weeds (Pinto-Zaervallos
and Zarbin 2013).
This contribution can be seen clearly and decisively in the cycle of nitrogen,
an essential element to some molecules that integrate the organic matter. Plants,
with some exceptions, do not have the capacity to absorb this element from the

Fig. 1.2 Sir Humphry Davy,


detail of an oil painting.
Courtesy of the National
Portrait Gallery, London
1 Introduction to Sustainable Agrochemistry 5

atmosphere (with 78% nitrogen), the opposite of what occurs with another essential
element, carbon, which is absorbed as CO2 by means the photosynthesis. So, the
only natural way to close the nitrogen cycle is through the decomposition of organic
material from dead animal or plant, or the excretion of living things, and this form
of replenishment is naturally limited.
The capture and use of atmospheric nitrogen in the soil was only possible eco-
nomically via the works of the German chemists Fritz Haber and Carl Bosch. They
developed the Haber–Bosch reaction or process at the beginning of the twentieth
century (Ritter 2008), which allows the ammonia synthesis from the small reac-
tive atmospheric nitrogen and other abundant element, hydrogen, in industrial scale.
Curiously, the incentive that led to this essential innovation was not initially the pro-
duction of fertilizers, but the production of nitrates for military purposes (explosives)
to be used in World War I.

Nanotechnology,
biotechnology,
agronomy

Analytical &
environmental Food security
chemistry and Agrochemistry and enviornment
technological
chemistry

Sustainability

Fig. 1.3 Relation among agrochemistry and related areas of study and technological development
6 S. Vaz Jr.

Ammonia, a molecule with about 82% by weight of nitrogen, can be absorbed by


plants through the soil. However, for reasons of ease of application, it is preferred to
use as nitrogenous fertilizers solid substances derived from ammonia, such as urea
and ammonium nitrate. The world production of ammonia today reaches about 140
million tons per year (US Geological Survey 2017), and almost all global production
is destined to the synthesis of industrial fertilizers. The percentage of the world
population whose food depends on the use of synthetic nitrogen fertilizers is estimated
at 53% (Liu et al. 2016).
Taking the nitrogen fertilizers as an example, we can also conclude that so-called
organic farming, which advocates the exclusion of synthetic fertilizers, can function
as a niche market in societies of abundance, but it is certainly not an alternative to
feed the humanity. Then, we can see clearly the contribution of the agrochemistry to
the well-being of modern society. Figure 1.3 depicts a relation among agrochemistry
and related scientific and technological fields as an interdisciplinary theme.
Analytical and environmental chemistry allied to technological chemistry input
techniques, technologies and knowledge to analyse, produce and monitoring agro-
chemicals. Nanotechnology and biotechnology are new technological approaches
to be incorporated to the agrochemicals for the best agronomic usages. Food secu-
rity and environment are closely related to the agrochemical uses and near to the
consumer, implying in laws, market restrictions and public opinion. Finally, sustain-
ability is a demand of the society for greater quality of life and greater transparency
in the productive chains.

1.3 Classes of Agrochemicals and Their Uses

According to International Union of Pure and Applied Chemistry (2006), an agro-


chemical is an “agricultural chemical used in crop and food production including
pesticide, feed additive, chemical fertilizer, veterinary drug and related compounds”.
Currently, we can observe several agrochemical classes according to their uses in
agriculture:
• Fertilizers—any kind of substance applied to soil or plant tissues to provide one
or more nutrients essential to plant growth;
• Plant growth regulators—(also called plant hormones)—several chemical sub-
stances that profoundly influence the growth and differentiation of plant cells,
tissues and organs;
• Phytosanitary products, pesticides or correctives:

– Herbicides—agents, usually chemicals, used for killing or inhibiting the growth


of unwanted plants (i.e., weeds);
– Insecticides—pesticides formulated to kill, harm, repel or mitigate one or more
species of insects;
– Fungicides—pesticides that kill or prevent the growth of fungi and their spores;
1 Introduction to Sustainable Agrochemistry 7

– Acaricides—pesticides that kill members of the arachnid subclass Acari, which


includes ticks and mites;
– Bactericides—a substance that kills bacteria. Bactericides are disinfectants,
antiseptics, or antibiotics;
– Rodenticides—pesticides that kill rodents. Rodents include not only rats and
mice, but also squirrels, woodchucks, chipmunks, porcupines, etc.;
– Nematicides—a type of chemical pesticide used to kill plant-parasitic nema-
todes;
– Repellents—chemicals that can help reduce the risk of being bitten by insects
and therefore reduce the risk of getting a disease carried by mosquitos or ticks;
– Fumigants—any volatile, poisonous substance used to kill insects, nematodes,
and other animals or plants that damage stored foods or seeds;
– Disinfectants—antimicrobial agents that are applied to the surface of non-living
objects to destroy microorganisms that are living on such objects;
– Antibiotics—powerful drugs that fight bacterial infections; as a highlighted risk,
their continuous use may result in resistance to them by some microorganisms;
– Defoliants—a chemical dust or spray applied to plants to cause their leaves to
drop off prematurely;
– Algaecides—or algicide—a biocide used for killing and preventing the growth
of algae.

Agrochemicals move a huge global market, and it is expected that these markets
achieve 250.5 billion USD by 2020 (Statista 2018). However, they are one of the
main classes of chemical pollutants, with serious negative impacts on public health
and environment. The search for alternatives to conventional agrochemicals presents

Fig. 1.4 Fertilizer consumption in nutrients per ha of arable land, according to Food and Agriculture
Organization of the United Nations (2015)
8

Fig. 1.5 Pesticides per ha of arable land (kg ha−1 ), according to Food and Agriculture Organization of the United Nations (2015)
S. Vaz Jr.
1 Introduction to Sustainable Agrochemistry 9

itself as an excellent opportunity for the development of sustainable agricultural


technologies and for opening new businesses.
As an example of the demand for agrochemicals by the agricultural sector, in
Fig. 1.4 it is presented the world consumption of fertilizers. Asia and Americas have
higher fertilizer consumption because the most agricultural production is located in
these continents.
In Fig. 1.5, the world pesticide is shown. Americas, Europe and part of Africa and
Asia have higher pesticide consumption aiming plagues’ control. Interestingly, small
countries like Japan and Portugal show consumption comparable to larger countries
such as Argentina.

1.4 Impacts of Agriculture on the Environment and Health

Governments, farmers and consumers show increasing concerns with the negative
impacts on the environment and health caused by the large amount of inputs applied
to produce different crops worldwide. Agrochemicals have a direct correlation with
damages from agriculture, with pesticides being the main representative class with
toxicological implications.
The main negatives from the agriculture impacts on the environment are:
• Water and air pollution due to the use of pesticides;
• Extinction of water bodies, due to high water demand;
• Erosion and soil degradation due to inadequate management during cultivation;
• Change in biota due to factors already listed;
• Changes in the quality of environmental resources, also due to factors already
listed;
• Ecological risks for insects, plants and animals associated with the change of
environment;
• Climate change due to the deforestation and biomass combustion.
Regarding those impacts on health, it can be highlighted:
• Poisoning due to pesticide use and food contaminated consumption;
• Occupational risks to farmers due to the exposition to pesticides;
• Human infections, or emerging infectious diseases, that do not respond to treatment
due to the use of antimicrobials in agriculture (Grace 2018).
From these negative impacts, it is becoming paramount the development of a more
environmentally and healthy friendly agriculture.
10 S. Vaz Jr.

1.5 Sustainability and Agrochemistry

Agricultural chemistry is, undoubtedly, one of the fields of research and business
whose impact is felt throughout the world, since we all need to eat to survive. Added
to this is the fact that, increasingly, technology is intertwining with modern agri-
culture, both with regard to new production strategies and the reduction of negative
environmental impacts (Herman 2015).
Sustainability can be seen and understood by means of its three components:
• Environmental impacts;
• Economic impacts;
• Societal impacts.
Impacts can be positive or negative according their direct or indirect effects upon
the environment, economy and society. Considering that agrochemistry is the appli-
cation of chemistry and its concepts and technologies to promote a better agriculture,
economic and societal impacts are expected to be positive, specially the economic
impacts. On the other hand, and due to a historic of environmental incidents at global
level (e.g., water pollution), environmental impacts are expected to be negative; nev-
ertheless, it could be positive if modern technologies and good agricultural practices
are used. A more detailed evaluation of sustainability in agriculture can be seen in
Quintero-Angel and González-Acevedo (2018).
Sustainable chemistry, a recent branch of chemistry, was defined as “(…) a sci-
entific concept that seeks to improve the efficiency with which natural resources are
used to meet human needs for chemical products and services. Sustainable chemistry
encompasses the design, manufacture and use of efficient, effective, safe and more
environmentally benign chemical products and processes.” (Organization of Eco-
nomic Co-operation and Development 2018). From these statements, a relationship
with agrochemistry can be constructed by means of design, manufacture and use of
efficient, effective, safe and more environmentally benign agrochemicals, that is, by
the establishment of a strong innovation drive in agriculture for the next decades as
we can read in this book.

References

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www.bbc.co.uk/history/british/empire_seapower/agricultural_revolution_01.shtml. Accessed
Aug 2018
Encyclopaedia Britannica (2018) Sir Humphry Davy, Baronet. https://www.britannica.com/
biography/Sir-Humphry-Davy-Baronet#ref172328. Accessed Aug 2018
Food and Agriculture Organization of the United Nations (2015) FAO statistical pocketbook. http://
www.fao.org/3/a-i4691e.pdf. Accessed Aug 2018
Grace D (2018) Infectious diseases and agriculture. Reference module in food science. https://doi.
org/10.1016/B978-0-08-100596-5.21570-9
1 Introduction to Sustainable Agrochemistry 11

Herman C (2015) Agricultural chemistry: new strategies and environmental perspectives to feed a
growing global population. American Chemical Society, Washington
International Union of Pure and Applied Chemistry (2006) Glossary of terms related to pesticides.
In: Stephenson GR, Ferris IG, Holland PT, Norberg M (eds) Pure and applied chemistry, vol 78,
pp 2075–2154
Liu J, Ma K, Ciais P, Polasky S (2016) Reducing human nitrogen use for food production. Scientific
reports. https://doi.org/10.1038/srep30104
Nobel Prize (2018) Norman Borlaug—biographical. https://www.nobelprize.org/nobel_prizes/
peace/laureates/1970/borlaug-bio.html. Accessed Aug 2018
Organization of Economic Co-operation and Development (2018) Sustainable chemistry. http://
www.oecd.org/env/ehs/risk-management/sustainablechemistry.htm. Accessed Aug 2018
Pingali D (2012) Green revolution: impacts, limits, and the path ahead. Proc Natl Acad Sci U S A
109:12302–12308
Pinto-Zaervallos DM, Zarbin PHG (2013) Química na agricultura: perspectivas para o desenvolvi-
mento de tecnologias sustentáveis [Chemistry in agriculture: perspectives for the development of
sustainable technologies]. Quim Nova 36:1509–1513
Quintero-Angel M, González-Acevedo A (2018) Tendencies and challenges for the assessment of
agricultural sustainability. Agr Ecosyst Environ 254:273–281
Ritter SK (2008) The Harber-Bosh reaction: an early chemical impact on sustainability. Chemical &
Engineering News. https://pubs.acs.org/cen/coverstory/86/8633cover3box2.html. Accessed Aug
2018
Statista (2018) Worldwide agrochemical market value in 2014 and 2020 (in billion
U.S. dollars). https://www.statista.com/statistics/311957/global-agrochemical-market-revenue-
projection/. Accessed in Aug 2018
U.S. Geological Survey (2017) USGS Minerals information: nitrogen. https://minerals.usgs.gov/
minerals/pubs/commodity/nitrogen/. Accessed Aug 2018
Chapter 2
The Battle for a Sustainable Food Supply

John Unsworth, Yoshiaki Nakagawa, Caroline Harris and Gijs Kleter

Abstract Since the time that Homo sapiens took up farming, a battle has been
waged against pests and diseases which can cause significant losses in crop yield
and threaten a sustainable food supply. Initially, early control techniques included
religious practices or folk magic, hand removal of weeds and insects, and “chemical”
techniques such as smokes, easily available minerals, oils and plant extracts known
to have pesticidal activity. But it was not until the early twentieth century that real
progress was made when a large number of compounds became available for testing
as pesticides due to the upsurge in organic chemistry. The period after the 1940s
saw the introduction of important families of chemicals, such as the phenoxy acid
herbicides, the organochlorine insecticides and the dithiocarbamate fungicides. The
introduction of new pesticides led to significant yield increases, but concern arose
over their possible negative effects on human health and the environment. In time,
resistance started to occur, making these pesticides less effective. This led agrochem-
ical companies putting in place research looking for new modes of action and giving
less toxic and more environmentally friendly products. These research programmes
gave rise to new pesticide families, such as the sulfonylurea herbicides, the strobilurin
fungicides and the neonicotinoid insecticide classes.

This work formed part of a project (Project No. 2012-019-1-600) supported by the International
Union of Pure and Applied Chemistry (IUPAC).

J. Unsworth (B)
Consultant, Vellacotts,
Chelmsford CM1 7EA, UK
e-mail: [email protected]
Y. Nakagawa
Graduate School of Agriculture, Kyoto University, Kyoto 606-8502, Japan
C. Harris
Exponent International Limited, The Lenz, Hornbeam Business Park, Harrogate,
North Yorkshire HG2 8RE, UK
G. Kleter
Wageningen Food Safety Research, Part of Wageningen University & Research,
Akkermaalsbos 2, 6708 WB Wageningen, The Netherlands

© Springer Nature Switzerland AG 2019 13


S. Vaz Jr. (ed.), Sustainable Agrochemistry,
https://doi.org/10.1007/978-3-030-17891-8_2
14 J. Unsworth et al.

Keywords Pest management · Crop yield · Natural pesticides · Inorganic


pesticides · Synthetic organic pesticides · Pest resistance

2.1 Introduction

Homo sapiens, or Latin for wise man, was a term first applied in 1758 by Linnaeus for
modern humans. They are thought to have first appeared roughly 200,000 years ago
in what is now Ethiopia (McDougall et al. 2005), although there is some evidence that
they emerged substantially earlier than this (Hublin et al. 2017). Until the practice of
agriculture some 190,000 years later, the only way to obtain a sufficient food supply
was by foraging, i.e., hunting wild animals and gathering edible plants and seeds.
Hunter-gatherer societies were generally nomadic, moving several times each year
to new areas when the returns of forays from their current camp dropped below those
to be expected from another camp (Kelly 1992). The transition from hunter-gatherer
to agriculture can be considered as having occurred in four steps (Olsson 2001):
(i) Wild plant–food procurement—hunter-gatherers in this stage occasionally burn
the vegetation, gather and protect useful plants and fruit.
(ii) Wild plant–food production with some tillage—maintenance of plant popula-
tions in the wild is carried out, and both planting and sowing of wild plants
are undertaken, as well as weeding. Seeds from selected plants with desirable
characteristics are propagated in new habitats. When harvesting is completed,
some of the seed is stored for future use.
(iii) Cultivation with systematic tillage—land is cleared and the food-producing
activities that were introduced in the previous stage (sowing, weeding, propa-
gation of plants, etc.) are intensified.
(iv) Plant domestication—as the selective cultivation of plants proceeds, new geno-
types eventually appear that more efficiently serve human needs.
Several theories have been put forward to explain why the change from hunter-
gatherers to farmers took place (Price and Bar-Yosef 2011; Weisdorf 2005), including
changes in climate (Richerson et al. 2001; Dow et al. 2005; Bettinger et al. 2009),
population density and technological sophistication (Baker 2008), over-hunting of
large animals leading to their extinction (Smith 1975) and social evolution (Bender
1978). It has also been postulated that as societies began to use ever more varied
equipment for foraging with containers, stone mortars, baskets, fishnets and traps,
drying racks, etc., plus starting to store food in pits and embarking on agriculture, the
necessary equipment became increasingly burdensome. Moving regularly from one
area to another became more difficult and mobility was thus reduced, with people
tending to stay in one area. This increasing and practically enforced sedentarism of
hunter-gatherers increased their involvement in agriculture (de Saulieu and Testart
2015). Whatever the reason, or combination of reasons, agriculture, as in the cultiva-
tion of plants for food, was adopted between 5000 and 12,000 years ago. Independent
points of origin included the Near East, north and south China, sub-Saharan Africa,
2 The Battle for a Sustainable Food Supply 15

Central Mexico, South-Central Andes and the eastern USA, with subsequent diffu-
sion to most parts of the globe (Richerson et al. 2001). Once adopted, agriculturally
based communities increased significantly in size (Gignouxa et al. 2011) with the
corresponding requirement to increase the food supply.
The practice of agriculture first began in the Fertile Crescent of Mesopotamia
(part of present-day Iraq, Turkey, Syria and Jordan) where edible seeds were initially
gathered, as in stage (i) above, by a population of hunter-gatherers (Kislev et al.
2004). Interestingly, the change from hunting and gathering to farming as a subsis-
tence method does not seem to be a logical choice since, with a limited population
and plentiful resources, hunting and gathering are less demanding work than farm-
ing. Agriculture requires cooperation, whilst living in settlements has social impacts
such as diseases, social inequality and division of labour (Hirst 2017). The wide-
scale transition of many human cultures from a lifestyle of hunting and gathering
to one of agriculture and settlement, allowing the ability to support an increasingly
large population, has been called the Neolithic Revolution or Neolithic Demographic
Transition (Bellwood and Oxenham 2008). The adoption of agriculture in the Stone
Age is one of the major curiosities in human cultural history since studies carried out
among contemporary primitive people suggest that the first farmers worked harder
to attain subsistence than their foraging predecessors (Weisdorf 2009). Although the
population explosion that followed the Neolithic Revolution was initially explained
by improved health for agriculturalists, empirical studies of societies shifting subsis-
tence from foraging to primary food production have found evidence for deteriorat-
ing health from an increase in infectious and dental disease and a rise in nutritional
deficiencies (Latham 2013). Hence, early farmers appear to have been smaller and
less healthy than hunter-gatherers (Cohen and Crane-Kramer 2007; Armelagos et al.
1991). This counterintuitive increase in nutritional diseases resulted from factors
such as seasonal hunger, reliance on single crops deficient in essential nutrients, crop
blights and social inequalities. The impact of agriculture, accompanied by increas-
ing population density and a rise in infectious disease, was observed to decrease
stature in populations across the entire globe, regardless of the temporal period dur-
ing which agriculture was adopted, including Europe, Africa, the Middle East, Asia,
South America and North America (Mummert et al. 2011).
As the population became more settled, and farming became the way of life, it
has been suggested that eight plants, the so-called eight founder crops formed the
basis of the origins of agriculture, with all eight coming from the Fertile Crescent
region. The eight crops include three cereals (einkorn wheat, emmer wheat and
barley), four legumes (lentil, pea, chickpea and bitter vetch) and one oil and fibre
crop—flax or linseed (Hirst 2018; Weiss and Zohary 2011). Somewhat later, rice
and millet were domesticated in China (Deng et al. 2015), whilst about 7500 years
ago rice and sorghum were being farmed in the Sahel region of Africa. Local crops
were domesticated independently in West Africa and possibly in New Guinea and
Ethiopia, whilst several crop plants were domesticated in the Americas including
corn, squashes, potato and sunflowers (Petruzzello 2019). It is clear that farmed
crops could suffer from pests and diseases causing a large loss in yield, with the
16 J. Unsworth et al.

ever-present possibility of famine for the population. There was thus a great incentive
to find ways of overcoming the problems caused by pests and diseases.
There are a large number of pests and diseases that can affect crops, thereby
reducing crop yield with the concomitant possibility of hunger and in extreme cases
famine for the population. It has been estimated that there are over 70,000 different
pest species that can damage agricultural crops, including 9000 species of insects and
mites, 50,000 species of plant pathogens and 8000 species of weeds. Although, in
general, less than 10% of these organisms are considered to be major pests, this still
leaves a considerable number that can have a significant effect on crop productivity
(Pimental 2009). In addition, farmers also have to contend with damage caused by
nematodes, slugs, snails, birds and mammals. Damage to crops can occur in different
ways, and pests and diseases can be classified according to the types of damage that
they cause (Boote et al. 1983). These classifications are:
• Stand reducers—e.g. damping-off fungi;
• Photosynthetic rate reducers—e.g. fungi, bacteria, viruses;
• Leaf senescence—e.g. various pathogens;
• Light stealers—e.g. weeds, some leaf pathogens;
• Assimilate sappers—e.g. nematodes, pathogens, sucking arthropods;
• Tissue consumers—e.g. chewing animals, necrotrophic pathogens;
• Turgor reducers—e.g. nematodes, root-feeding insects, root rot pathogens.
Initially, pest control was probably a manual occupation, e.g. weeding or removal
of caterpillars and insects by hand, followed by attempts to protect crops by means of
religion, folk magic and the use of what may be termed early chemical methods. For
example, pyrethrum is recorded as being used in China 2000 years ago (McLaughlin
Gormley King Company 2010) and is still used today (Market Research.com 2015).
Up until the mid-nineteenth century, various plant, animal or mineral derivatives
(Smith and Secoy 1975, 1976a) were used for pest control; these were later augmented
by compounds from the dyestuff industry, e.g. Paris Green, and by-products from
the burgeoning chemical industry based on coal tar with phenol-based chemicals
(Gray 1916). The growth in synthetic pesticides accelerated in the 1940s with the
discovery of the pesticidal effects of numerous organic chemicals, which led to the
development of, for example, chlorinated hydrocarbons such as DDT as insecticides
and phenoxy acid derivatives like 2,4-D as herbicides. As the use of these “new”
pesticides increased, concerns for their effect on human health and the environment
were raised and attention was given to the development of low-rate, lower-risk organic
compounds as pesticides. It can be argued that there have been five significant phases
in the evolution of pesticides. With some inevitable overlap of the individual phases,
they can be summarised as:
• Before 1000—early pest management;
• 1000–1850—plant, animal or mineral derivatives;
• 1850–1940—inorganic products, industrial by-products;
• 1940–1970—synthetic organic compounds;
• 1970–present—lower-risk synthetic organic compounds.
2 The Battle for a Sustainable Food Supply 17

Given the significant number of pests and diseases that can impact on crop yields,
and the variety of ways by which this can happen, it is not surprising that since the
dawn of agriculture, farmers have been engaged in a constant battle against them.
This battle continues today, and it has been estimated that potential crop losses
worldwide would be between 50 and 82%, depending on the crop, in the absence
of pest control (Oerke 2006), and the economic loss has been estimated at US $250
billion (Agropages 2015).
As an example of the deleterious effects of weeds on crop yield, it has been
estimated that at a density of only two weeds per foot of row corn yield was reduced
10% by giant foxtail, 11% by common lambsquarters, 18% by velvetleaf and 22%
by common cocklebur (Moechnig et al. 2013). Given the fact that approximately 1
in 9 of the global population is suffering from undernourishment (FAO 2018) and
that the global population is estimated to rise from 7.3 billion today to 9 billion in
2050, it is essential that food shortages caused by reductions in crop yields due to
pests and diseases are kept to an absolute minimum. Chemistry will continue to have
a significant role in the control of fungal pathogens, insect pests and weeds which,
together with modified farming practices, new crop varieties, increased irrigation and
optimum fertiliser use, will play a key role in achieving increased and sustainable
productivity.

2.2 Early Efforts at Pest Management

Ancient farmers would have tried every method available to them at the time to
maintain or increase crop yields. For example, in the Mediterranean region repeated
fertilisation of the soil, careful tillage, intelligent selection of crops suited to climate
and soil, importation of foreign seed and plants, painstaking seed selection, with
crop protection techniques as part of the mix were all used (Semple 1928). However,
despite this, the yield per hectare remained relatively low. In Mesopotamia, irrigated
cereal crops have been reported as giving a yield of almost 2 tonnes per hectare,
although this declined to only about one-third as much by 1700 BC, probably due to
increasing salinity of the irrigated areas (Evans 1980). A maximum return known in
Roman Italy about 50 BC gave a yield of eight to tenfold of planted seed. This was
the yield of winter wheat from rich volcanic soils in eastern Sicily, and it amounted to
20 or 24 bushels to the acre, or approximately 1.4–1.7 tonnes per hectare. A yield of
15-fold or 36 bushels of wheat to the acre (approximately 2.5 tonnes per hectare) was
quite extraordinary and restricted to a few favoured spots (Semple 1928), although
wheat yields of up to 3.6 tonnes per hectare were probably obtained in Galilee around
the beginning of the first century AD (Evans 1980). These figures can be compared
with about 5 tonnes/hectare achieved in Greece and Italy and 7 tonnes per hectare
in Egypt today (World Bank 2018). Nevertheless, whilst reasonable yields could be
obtained under some favourable conditions, the possibility of disease significantly
reducing crop yield was always present. Cereal stem rust was particularly feared and
could lead to 100% crop loss. In the Bible, there are several references to epidemics
18 J. Unsworth et al.

of rusts and smuts inflicted on the Israelites as a punishment for their sins (Leonard
and Szabo 2005).

2.2.1 Religion and Folk Magic

In many agricultural societies, there was a widespread belief that pests were evil and
were sent as a punishment from the gods; as a result, many early pest management
strategies were based on religious practices. Thus, the Sumerian goddess Ninkilim
was invoked to protect growing barley from damage by rodents, and a Babylonian
tablet contains an incantation asking the sun and moon gods to prevent ergot from
developing on ears of grain (Secoy and Smith 1977). In Egypt, reliefs in tombs dating
as early as the Old Kingdom, about 2649–2150 BC, show that illiterate shepherds
and farmers could recite spells to protect herds and crops (Zucconi 2007). In ancient
Rome, the deity Robigus and the goddess Flora were believed to be responsible
for rust and mildew, but when they were appeased, these diseases would not harm
the grain and trees. In honour of Robigus, the solemn feast of the Robigalia was
thus established, during which a rust coloured dog was sacrificed, while for Flora,
there were games called Floralia (Smith and Secoy 1975). The Greek god Apollo
was believed to give protection to crops against summer heat, blight, mildew and
other pests such as field mice and grasshoppers (Secoy and Smith 1977). In India the
Atharvaveda, a sacred Hindu text, described the chanting of mantras to protect crops
from insects such as grasshoppers; one such mantra from around 400 BC evokes the
Hindu deity Rama to protect grain. The mantra had to be written with the red lac-dye
on a leaf and tied in the field (Goyal 2003). Similarly, in Sri Lanka, mantras have been
used in agriculture to obtain higher yields and also to protect crops from damage by
pests and wild animals (Upawansa and Wagachchi 2018). In Japan, farmers would
entreat deities through ceremonies such as mushi-oi or mushi-okuri, meaning driving
away noxious insects. For this, the local population walked around rice paddies in
procession shouting, with bells and torches to drive away rice pests and pray for
that year’s farming. In addition, farmers would place beef or phallic-shaped ritual
equipment in the water inlets to fields to ward off insect pests (Ohta 2013). Besides
the invocation of deities to keep pests from crops, there were a host of folk remedies
that could also be practiced. Thus, in Roman times, it was thought that trees could
be protected against caterpillars by touching the tops with the gall of a green lizard
and that a crayfish hung up in the middle of a garden would afford protection against
caterpillars. Mildew would be controlled by placing branches of laurel in the ground,
near to growing wheat, thus causing the mildew to pass from the fields to the foliage
of the laurels. Diseases of millet could be prevented by carrying a toad around the
field at night before burying the creature in the middle of the field in a pot (Smith and
Secoy 1975). According to Columella (4–70 AD), the Etruscans set the flayed head
of an ass at the edge of a field to protect grain from rust (Secoy and Smith 1977). It
was also hoped that amulets, sometimes in the form of a protective deity, and others
shaped like the pest itself, would ward off a danger, e.g. locust amulets which have
2 The Battle for a Sustainable Food Supply 19

been discovered in ancient Egyptian tombs (Kenawy and Abdel-Hamid 2015). Many
other religious and folk remedies have been described, and more information can be
found in the above references.

2.2.2 Manual and Physical Techniques

Whilst religious and folk remedies were of dubious value other management prac-
tices involving, for example, removal of weeds by hand, hoeing or ploughing and
the picking of insects from crops would have been successful, if time-consuming.
Chasing or throwing stones at birds and small mammals would have been effective,
and, although probably less effective, scarecrows were also used to deter these pests.
The first scarecrows are reported in Ancient Egypt, dating back over 3000 years,
where they were used to protect wheat fields along the river Nile from quail. The
Greeks, Romans, Japanese and many other cultures also used scarecrows to protect
their crops.
The practice of burning cereal stubble after harvest would also have contributed
to the destruction of weed seeds. In addition, physical methods such as the use of
tar on tree trunks to trap crawling insects and the adding of fine sand to stored grain
would prevent much of the damage caused by insects. In stored grain, the addition of
fine sand inhibits movement of the insects and reduces the oxygen content, thereby
making it hard for the insects to get enough oxygen to survive; in addition, the sand
is abrasive and would scratch the hard covering that surrounds the bodies of beetles,
leading to desiccation and death (De Groot 2004). This method is still used today,
particularly in Africa, and was used in ancient Egypt. It can be assumed that it was
sufficiently effective to exterminate a pest such as the lesser grain borer, recently
arrived in the region (Pharaonic Egypt 2000). Similarly in 750 BC, Homer noted the
use of wood ash spread on land as a form of pest control where it acts as a physical
poison, usually causing abrasion of epicuticular waxes and thus exposing pests to
death through desiccation. It also interferes with the chemical signals emanating from
the host plants, thus obstructing the initial host location from pests. When foliage is
treated with wood ash, it also becomes unpalatable for foliage feeders like cutworms,
caterpillars and grasshoppers (Verma 1998).
Biological methods were also used; for example, mention is made in a docu-
ment reputed to have been written in 304 AD of the use in ancient China of citrus
ants against insects found on citrus trees (Huang and Yang 1987). Similarly, around
1000 AD in Arabia, ants were moved from nearby mountains to oases to control
phytophagous ants which attack date palms.
Another random document with
no related content on Scribd:
Fig. 124—Cis melliei. Martinique. A, Perfect Insect; B, pupa; C, larva;
D, terminal portion of body of larva. (After Coquerel.)

Fam. 51. Sphindidae.—This family of half a dozen species of rare


and small Insects, differs from Cioidae by the tarsi being five-jointed
at any rate on the front and middle feet, opinions differing as to
whether the number of joints of the hind tarsi is four or five. These
Insects live in fungi growing in wood, e.g. Reticularia hortensis, that
are at first pulpy and afterwards become powder. The larvae of both
of our British genera, Sphindus and Aspidiphorus, have been
described by Perris, who considers them allied to the fungivorous
Silphidae and Latridiidae. The systematic position of these Insects
has been the subject of doubt since the days of Latreille.

Fam. 52. Bostrichidae (Apatidae of some authors).—Tarsi five-


jointed, but the first joint very short and imperfectly separated from
the second; front coxae prominent, contiguous, very little extended
transversely; five visible ventral segments. The Bostrichidae attack
dry wood, and sometimes in such large numbers that timber is
entirely destroyed by them; most of them make cylindrical burrows
into the wood. The larvae have the posterior part of the body
incurved, and resemble the wood-boring larvae of Anobiidae rather
than the predaceous larvae of Cleridae. We follow Leconte and Horn
in placing Lyctides as a division of Bostrichidae; although differing
very much in appearance, they have similar habits and larvae. The
typical Bostrichides are remarkable for their variety of sculpture and
for the shapes of the posterior part of the body; this part is more or
less conspicuously truncate, and furnished with small prominences.
Dinapate wrightii, found in the stems of a species of Yucca in the
Mojave desert of California, attains a length of nearly two inches; its
larva is extremely similar to that of A. capucina. Some of the forms
(Phonapate) stridulate in a manner peculiar to themselves, by
rubbing the front leg against some projections at the hind angle of
the prothorax. Upwards of 200 species of the family are known. In
Britain we have only four small and aberrant forms.

Fig. 125—Apate capucina. Europe. A, Larva (after Perris); B, perfect


Insect.

Fam. 53. Ptinidae.—Tarsi five-jointed, first joint not reduced in size,


often longer than second; front and middle coxae small, not
transversely extended, the former slightly prominent; five visible
ventral segments; prosternum very short. Here are included two sub-
families, Ptinides and Anobiides; they are considered as distinct
families by many authors, but in the present imperfect state of
knowledge[124] it is not necessary to treat them separately.

Fig. 126—"Biscuit-weevil." Anobium paniceum.


Fig. 127.—Early stages of Anobium paniceum. A, Eggs, variable in
form; B, larva; C, pupa; D, asymmetrical processes terminating
body of pupa. [This larva is probably the "book-worm" of
librarians].

Ptinidae are sometimes very destructive to dried animal matter, and


attack specimens in museums; Anobiides bore into wood, and
apparently emerge as perfect Insects only for a very brief period;
Anobium (Sitodrepa) paniceum is, however, by no means restricted
in its tastes; it must possess extraordinary powers of digestion, as
we have known it to pass several consecutive generations on a diet
of opium; it has also been reported to thrive on tablets of dried
compressed meat; in India it is said to disintegrate books; a more
usual food of the Insect is, however, hard biscuits; weevilly biscuits
are known to every sailor, and the so-called "weevil" is usually the
larva of A. paniceum (Fig. 127, B). In the case of this Insect we have
not detected more than one spiracle (situate on the first thoracic
segment); the other known larvae of Anobiides are said to possess
eight abdominal spiracles. The skeleton in some of this sub-family is
extremely modified, so as to allow the Insects to pack themselves up
in repose; the head is folded in over the chest, and a cavity existing
on the breast is thus closed by the head; in this cavity the antennae
and the prominent mouth-parts are received and protected; the legs
shut together in an equally perfect manner, so that no roughness or
chink remains, and the creature looks like a little hard seed. Anobium
striatum is a common Insect in houses, and makes little round holes
in furniture, which is then said to be "worm-eaten." A. (Xestobium)
tessellatum, a much larger Insect, has proved very destructive to
beams in churches, libraries, etc. These species are the "death-
watches" or "greater death-watches" that have been associated with
the most ridiculous superstitions (as we have mentioned in Volume
V., when speaking of the lesser death-watches, or Psocidae). The
ticking of these Insects is really connected with sex, and is made by
striking the head rapidly against the wood on which the Insect is
standing.
The very anomalous genus Ectrephes (Fig. 128) is found in ants'
nests in Australia. Westwood placed it in Ptinidae. Wasmann has
recently treated it as a distinct family, Ectrephidae, associating it with
Polyplocotes and Diplocotes, and treating them as allied to
Scydmaenidae.

Fig. 128—Ectrephes kingi. West Australia. (After Westwood.)

Fam. 54. Malacodermidae.—Seven (or even eight) visible ventral


segments, the basal one not co-adapted in form with the coxae; tarsi
five-jointed. Integument softer than usual, the parts of the body not
accurately co-adapted. This important family includes a variety of
forms: viz. Lycides, Drilides, Lampyrides, Telephorides; though they
are very different in appearance, classifiers have not yet agreed on
separating them as families. Of these the Lampyrides, or glow-
worms, are of special interest, as most of their members give off a
phosphorescent light when alive; in many of them the female is
apterous and like a larva, and then the light it gives is usually
conspicuous, frequently much more so than that of its mate; in other
cases the males are the most brilliant. The exact importance of these
characters in the creatures' lives is not yet clear, but it appears
probable that in the first class of cases the light of the female serves
as an attraction to the male, while in the second class the very
brilliant lights of the male serve as an amusement, or as an
incitement to rivalry amongst the individuals of this sex.
Fig. 129—Phengodes hieronymi. Cordoba, South America. (After
Haase.) A, Male; B, female. l, l, Positions of luminous spots; ls,
spiracles. About × 3.

The well-known fire-flies (Luciola) of Southern Europe are an


example of the latter condition. They are gregarious, and on calm,
warm nights crowds of them may be seen moving and sparkling in a
charming manner. These individuals are all, or nearly all, males; so
rare indeed is the female that few entomologists have even noticed
it. The writer once assisted in a large gathering of Luciola italica in
the Val Anzasca, which consisted of many hundreds of specimens;
all of those he caught, either on the wing or displaying their lights on
the bushes, were males, but he found a solitary female on the
ground. This sex possesses ordinary, small eyes instead of the
large, convex organs of the male, and its antennae and legs are
much more feeble, so that though provided with elytra and wings it is
altogether a more imperfect creature. Emery has given an account of
his observations and experiments on this Insect, but they do not give
any clear idea as to the exact function of the light.[125] In our British
glow-worm the female is entirely apterous—hence the name glow-
worm—but the male has elytra and ample wings, and frequently flies
at night into lighted apartments. Although so little has been
ascertained as to the light of Lampyridae, there are two facts that
justify us in supposing that it is in some way of importance to the
species. These are: (1) that in a great many species the eyes have a
magnificent and unusual development; (2) that the habits of the
creatures are in nearly all cases nocturnal. It is true that the little
Phosphaenus hemipterus is said to be diurnal in habits, but it is
altogether an exceptional form, being destitute of wings in both
sexes, and possessed of only very feeble light-giving powers, and
we have, moreover, very little real knowledge as to its natural history;
it is said that the female is of the utmost rarity, though the male is not
uncommon.

The nature of the luminosity of Lampyris has given rise to many


contradictory statements; the light looks somewhat like that given off
by phosphorus, and is frequently spoken of as phosphorescence; but
luminescence is a better term. The egg, larva, pupa, and male are
luminous as well as the female (at any rate in L. noctiluca); the
luminescence is, however, most marked in the female imago, in
which it is concentrated near the extremity of the abdomen; here
there are two strata of cells, and many fine capillary tracheae are
scattered through the luminous substance. Wielowiejski concludes
that the light-producing power is inherent in the cells of the luminous
organ, and is produced by the slow oxidation of a substance formed
under the influence of the nervous system. The cells are considered
to be essentially similar to those of the fat-body.[126] The
luminescence of Lampyridae is very intermittent, that is to say, it is
subject to rapid diminutions and increases of its brilliancy; various
reasons have been assigned for this, but all are guesses, and all that
can be said is that the changes are possibly due to diminution or
increase of the air-supply in the luminous organ, but of the way in
which this is controlled there seems to be no evidence. Considerable
difference of opinion has existed as to the luminescence of the eggs
of Lampyris. If it exist in the matter contained in the egg, it is evident
that it is independent of the existence of tracheae or of a nervous
system. Newport and others believed that the light given by the egg
depended merely on matter on its exterior. The observations of
Dubois[127] show, however, that it exists in the matter in the egg; he
has even found it in the interior of eggs that had been deposited
unfertilised.

From time to time, since the commencement of the nineteenth


century, there have appeared imperfect accounts of extraordinary
light-giving larvae found in South America, of various sizes, but
attaining in some cases a length, it is said, of three inches; they are
reported as giving a strong red light from the two extremities of the
body, and a green light from numerous points along the sides of the
body, and hence are called, it is said, in Paraguay the railway-beetle.
We may refer the reader to Haase's paper[128] on the subject of
these "larvae," as we can here only say that it appears probable that
most of these creatures may prove to be adult females of the
extraordinary group Phengodini, in which it would appear that the
imago of the female sex is in a more larva-like state than it is in any
other Insects. The males, however, are well-developed beetles;
unlike the males of Lampyrides, in general they have not peculiar
eyes, but on the other hand they possess antennae which are
amongst the most highly developed known, the joints being
furnished on each side with a long appendage densely covered with
pubescence of a remarkable character. There is no reason to doubt
that Haase was correct in treating the Insect we figure (Fig. 129, B)
as a perfect Insect; he is, indeed, corroborated by Riley.[129] The
distinctions between the larva and female imago are that the latter
has two claws on the feet instead of one, a greater number of joints
in the antennae, and less imperfect eyes; the female is in fact a
larva, making a slightly greater change at the last ecdysis, than at
those previous. It is much to be regretted that we have so very small
a knowledge of these most interesting Insects. Malacodermidae are
probably the most imperfect or primitive of all beetles, and it is a
point of some interest to find that in one of them the phenomena of
metamorphosis are reduced in one sex to a minimum, while in the
other they are—presumably at least—normal in character.

Numerous larvae of most extraordinary, though diverse, shapes,


bearing long processes at the sides of the body, and having a head
capable of complete withdrawal into a slender cavity of the thorax,
have long been known in several parts of the world, and Dr. Willey
recently found in New Britain a species having these body-processes
articulated. Though they are doubtless larvae of Lampyrides, none of
them have ever been reared or exactly identified.

A very remarkable Ceylonese Insect, Dioptoma adamsi Pascoe, is


placed in Lampyrides, but can scarcely belong there, as apparently it
has but five or six visible ventral segments; this Insect has two pairs
of eyes, a large pair, with coarse facets on the under side of the
head, and a moderate-sized pair with fine facets on the upper side.
Nothing is known as to the habits of this curiosity, not even whether it
is luminous in one or both sexes.

It is believed that the perfect instar of Lampyrides takes no food at


all. The larvae were formerly supposed to be vegetarian, but it
appears probable that nearly all are carnivorous, the chief food being
Mollusca either living or dead. The larvae are active, and in many
species look almost as much like perfect Insects as do the imagos.

The other divisions of Malacodermidae—Lycides, Drilides,


Telephorides—also have predaceous, carnivorous larvae. All these
groups are extensive. Though much neglected by collectors and
naturalists, some 1500 species of the family Malacodermidae have
been detected. We have about 50 in Britain, and many of them are
amongst the most widely distributed and abundant of our native
Insects. Thus, however near they may be to the primitive condition of
Coleoptera, it is highly probable that they will continue to exist
alongside of the primitive Cockroaches and Aptera, long after the
more highly endowed forms of Insect-life have been extinguished
wholesale by the operations of mankind on the face of the earth.

Fig. 130—Malachius aeneus. Britain. A, Larva (after Perris); B, female


imago.

Fam. 55. Melyridae (or Malachiidae).—Six visible and moveable


ventral abdominal segments; the basal part more or less distinctly
co-adapted with the coxae. These Insects are extremely numerous,
but have been very little studied. In many works they are classified
with Malacodermidae, but were correctly separated by Leconte and
Horn, and this view is also taken by Dr. Verhoeff, the latest
investigator. The smaller number of visible ventral segments appears
to be due to a change at the base correlative with an adaptation
between the base of the abdomen and the hind coxae. The
characters are singularly parallel with those of Silphidae; but in
Melyridae the antennae are filiform or serrate, not clavate. The
habits in the two families are different, as the Melyridae are
frequenters of flowers. Many of the Melyridae have the integument
soft, but in the forms placed at the end of the family—e.g. Zygia—
they are much firmer. Thus these Insects establish a transition from
the Malacodermidae to ordinary Coleoptera. Although the imagos
are believed to consume some products of the flowers they frequent,
yet very little is really known, and it is not improbable that they are to
some extent carnivorous. This is the case with the larvae that are
known (Fig. 130, larva of Malachius aeneus). These are said by
Perris to bear a great resemblance to those of the genus Telephorus,
belonging to the Malacodermidae.

Fam. 56. Cleridae.—Tarsi five-jointed; but the basal joint of the


posterior very indistinct, usually very small above, and closely united
with the second by an oblique splice; the apices of joints two to four
usually prolonged as membranous flaps; anterior coxae prominent,
usually contiguous, rather large, but their cavities not prolonged
externally; labial palpi usually with large hatchet-shaped terminal
joint; ventral segments five or six, very mobile. The Cleridae are very
varied in form and colours; the antennae are usually more or less
clubbed at the tip, and not at all serrate, but in Cylidrus and a few
others they are not clubbed, and in Cylidrus have seven flattened
joints. The student should be very cautious in deciding as to the
number of joints in the feet in this family, as the small basal joint is
often scarcely distinguishable, owing to the obliteration of its suture
with the second joint. The little Alpine Laricobius has the anterior
coxal cavities prolonged externally, and the coxae receive the
femora to some extent, so that it connects Cleridae and
Derodontidae. The Cleridae are predaceous, and their larvae are
very active; they are specially fond of wood-boring Insects; that of
Tillus elongatus (Fig. 131) enters the burrows of Ptilinus pectinicornis
in search of the larva. The members of the group Corynetides
frequent animal matter, carcases, bones, etc., and, it is said, feed
thereon, but Perris's recent investigations[130] make it probable that
the larvae really eat the innumerable Dipterous larvae found in such
refuse; it is also said that the larvae of Cleridae spin cocoons for
their metamorphosis; but Perris has also shown that the larvae of
Necrobia ruficollis really use the puparia formed by Diptera. Some of
the species of Necrobia have been spread by commercial
intercourse, and N. rufipes appears to be now one of the most
cosmopolitan of Insects. The beautifully coloured Corynetes
coeruleus is often found in our houses, and is useful, as it destroys
the death-watches (Anobium) that are sometimes very injurious.
Trichodes apiarius, a very lively-coloured red and blue beetle,
destroys the larvae of the honey-bee, and Lampert has reared
Trichodes alvearius from the nests of Chalicodoma muraria, a
mason-bee; he records that one of its larvae, after being full grown,
remained twenty-two months quiescent and then transformed to a
pupa. Still more remarkable is a case of fasting of the larva of
Trichodes ammios recorded by Mayet;[131] this Insect, in its
immature form, destroys Acridium maroccanum; a larva sent from
Algeria to M. Mayet refused such food as was offered to it for a
period of two and a half years, and then accepted mutton and beef
as food; after being fed for about a year and a half thereon, it died.
Some Cleridae bear a great resemblance to Insects of other families,
and it appears probable that they resemble in one or more points the
Insects on which they feed. The species are now very numerous,
about 1000 being known, but they are rare in collections; in Britain
we have only nine species, and some of them are now scarcely ever
met with.
Fig. 131—Larva of Tillus elongatus. (New Forest). A, Head; B, front
leg; C, termination of the body, more magnified.

Fam. 57. Lymexylonidae.—Elongate beetles, with soft integuments,


front and middle coxae exserted, longitudinal in position; tarsi
slender, five-jointed; antennae short, serrate, but rather broad.
Although there are only twenty or thirty species of this family, they
occur in most parts of the world, and are remarkable on account of
their habit of drilling cylindrical holes in hard wood, after the manner
of Anobiidae. The larva of Lymexylon navale was formerly very
injurious to timber used for constructing ships, but of late years its
ravages appear to have been of little importance. The genus
Atractocerus consists of a few species of very abnormal Coleoptera,
the body being elongate and vermiform, the elytra reduced to small,
functionless appendages, while the wings are ample, not folded, but
traversed by strong longitudinal nervures, and with only one or two
transverse nervures. Owing to the destruction of our forests the two
British Lymexylonidae—L. navale and Hylecoetus dermestoides—
are now very rarely met with.

Fig. 132—Hydrocyphon deflexicollis. Britain. A, Larva (after Tournier);


B, imago.

Fam. 58. Dascillidae.—Small or moderate-sized beetles, with rather


flimsy integuments, antennae either serrate, filiform, or even made
flabellate by long appendages; front coxae elongate, greatly
exserted; abdomen with five mobile ventral segments; tarsi five-
jointed. This is one of the most neglected and least known of all the
families of Coleoptera, and one of the most difficult to classify;
though always placed amongst the Serricornia, it is more nearly
allied to Parnidae and Byrrhidae, that are placed in Clavicornia, than
it is to any of the ordinary families of Serricornia. It is probable that
careful study will show that it is not natural as at present constituted,
and that the old families, Dascillidae and Cyphonidae, now
comprised in it, will have to be separated. Only about 400 species
are at present known; but as nearly 100 of these have been detected
in New Zealand, and 17 in Britain, doubtless the numbers in other
parts of the world will prove very considerable, these Insects having
been neglected on account of their unattractive exterior, and fragile
structure. The few larvae known are of three or four kinds. That of
Dascillus cervinus is subterranean, and is believed to live on roots; in
form it is somewhat like a Lamellicorn larva, but is straight, and has a
large head. Those of the Cyphonides are aquatic, and are
remarkable for possessing antennae consisting of a great many
joints (Fig. 132, A). Tournier describes the larva of Helodes as
possessing abdominal but not thoracic spiracles, and as breathing
by coming to the surface of the water and carrying down a bubble of
air adhering to the posterior part of the body; the larva of
Hydrocyphon (Fig. 132, A) possesses several finger-like pouches
that can be exstulpated at the end of the body. It is probable that
these larvae are carnivorous. The imago of this Insect abounds on
the bushes along the banks of some of the rapid waters of Scotland;
according to Tournier, when alarmed, it enters the water and goes
beneath it for shelter. The third form of larva belongs to the genus
Eucinetus, it lives on fungoid matter on wood, and has ordinary
antennae of only four joints.[132] It is very doubtful whether Eucinetus
is related to other Dascillidae; some authorities indeed place it in
Silphidae.

Fam. 59. Rhipiceridae.—Tarsi five-jointed, furnished with a robust


onychium (a straight chitinous process bearing hairs) between the
claws; antennae of the male bearing long processes, and sometimes
consisting of a large number of joints. Mandibles robust, strongly
curved, and almost calliper-like in form. This small family of less than
100 species is widely distributed, though confined to the warmer
regions of the earth, a single species occurring in the extreme south
of Eastern Europe. Very little is known as to the natural history. The
larva of Callirhipis dejeani (Fig. 133, A) is described by Schiödte as
hard, cylindrical in form, and peculiarly truncate behind, so that there
appear to be only eight abdominal segments, the ninth segment
being so short as to look like an operculum at the extremity of the
body. It lives in wood.

Fig. 133—A, Larva of Callirhipis dejeani (after Schiödte); B, Rhipicera


mystacina male, Australia; C, under side of its hind foot.

Fig. 134—Athous rhombeus. New Forest. A, Larva; B, female imago.

Fam. 60. Elateridae (Click-beetles).—Antennae more or less serrate


along the inner margin, frequently pectinate, rarely filiform. Front
coxae small, spherical. Thorax usually with hind angles more or less
prolonged backwards; with a prosternal process that can be received
in, and usually can move in, a mesosternal cavity. Hind coxa with a
plate, above which the femur can be received. Visible ventral
segments usually five, only the terminal one being mobile. Tarsi five-
jointed. This large family of Coleoptera comprises about 7000
species. Most of them are readily known by their peculiar shape, and
by their faculty of resting on the back, stretching themselves out flat,
and then suddenly going off with a click, and thus jerking themselves
into the air. Some, however, do not possess this faculty, and certain
of these are extremely difficult to recognise from a definition of the
family. According to Bertkau[133] our British Lacon murinus is
provided near the tip of the upper side of the abdomen with a pair of
eversible glands, comparable with those that are better known in
Lepidopterous larvae. He states that this Insect does not try to
escape by leaping, but shams death and "stinks away" its enemy.
The glands, it would appear, become exhausted after the operation
has been repeated many times. The extent of the leap executed by
click-beetles differs greatly; in some species it is very slight, and only
just sufficient to turn the Insect right side up when it has been placed
on its back. In some cases the Insects go through the clicking
movements with little or no appreciable result in the way of
consequent propulsion. Although it is difficult to look on this clicking
power as of very great value to the Elateridae, yet their organisation
is profoundly modified so as to permit its accomplishment. The
junction of the prothorax with the after-body involves a large number
of pieces which are all more or less changed, so that the joint is
endowed with greater mobility than usual; while in the position of
repose, on the other hand, the two parts are firmly locked together.
The thoracic stigma is of a highly remarkable nature, and the
extensive membrane in which it is placed appears to be elastic.
Although the mechanics of the act of leaping are still obscure, yet
certain points are clear; the prosternal process possesses a
projection, or notch, on its upper surface near the tip; as a
preliminary to leaping, this projection catches against the edge of the
mesosternal cavity, and as long as this position is maintained the
Insect is quiescent; suddenly, however, the projection slips over the
catch, and the prosternal process is driven with force and rapidity
into the mesosternal cavity pressing against the front wall thereof,
and so giving rise to the leap.

Several larvae are well known; indeed the "wire-worms" that are
sometimes so abundant in cultivated places are larvae of Elateridae.
In this instar the form is usually elongate and nearly cylindrical; the
thoracic segments differ but little from the others except that they
bear rather short legs; the skin is rather hard, and usually bears
punctuation or sculpture; the body frequently terminates in a very
hard process, of irregular shape and bearing peculiar sculpture on its
upper surface, while beneath it the prominent anal orifice is placed:
this is sometimes furnished with hooks, the function of which has not
yet been observed. The majority of these larvae live in decaying
wood, but some are found in the earth; as a rule the growth is
extremely slow, and the life of the larva may extend over two or more
years. Some obscurity has prevailed as to their food; it is now
considered to be chiefly flesh, though some species probably attack
decaying roots; and it is understood that wire-worms destroy the
living roots, or underground stems, of the crops they damage.
Various kinds of Myriapods (see Vol. V. p. 29) are often called "wire-
worm," but they may be recognised by possessing more than six
legs. The larvae of the genus Cardiophorus are very different, being
remarkably elongate without the peculiar terminal structure, but
apparently composed of twenty-three segments.

The genus Pyrophorus includes some of the most remarkable of


light-giving Insects. There are upwards of 100 species, exhibiting
much diversity as to the luminous organs; some are not luminous at
all; but all are peculiar to the New World, with the exception that
there may possibly be luminous species, allied to the American
forms, in the Fiji Islands and the New Hebrides. In the tropics of
America the Pyrophorus, or Cucujos, form one of the most
remarkable of the natural phenomena. The earliest European
travellers in the New World were so impressed by these Insects that
descriptions of their wondrous display occupy a prominent position in
the accounts of writers like Oviedo, whose works are nearly 400
years old. Only one of the species has, however, been investigated.
P. noctilucus is one of the most abundant and largest of the
Pyrophorus, and possesses on each side of the thorax a round
polished space from which light is given forth; these are the organs
called eyes by the older writers. Besides these two eye-like lamps
the Insect possesses a third source of light situate at the base of the
ventral surface of the abdomen; there is no trace of this latter lamp
when the Insect is in repose; but when on the wing the abdomen is
bent away from the breast, and then this source of light is exposed;
hence, when flying, this central luminous body can be alternately
displayed and concealed by means of slight movements of the
abdomen. The young larva of P. noctilucus is luminous, having a
light-giving centre at the junction of the head and thorax; the older
larva has also numerous luminous points along the sides of the body
near the spiracles. It is remarkable that there should be three
successive seats of luminescence in the life of the same individual.
The eggs too are said to be luminous. The light given off by these
Insects is extremely pleasing, and is used by the natives on
nocturnal excursions, and by the women for ornament. The structure
of the light-organs is essentially similar to that of the Lampyridae.
The light is said to be the most economical known; all the energy
that is used being converted into light, without any waste by the
formation of heat or chemical rays. The subject has been
investigated by Dubois,[134] who comes, however, to conclusions as
to the physiology of the luminous processes different from those that
have been reached by Wielowiejski and others in their investigations
on Glow-worms. He considers that the light is produced by the
reactions of two special substances, luciferase and luciferine.
Luciferase is of the nature of an enzyme, and exists only in the
luminous organs, in the form, it is supposed, of extremely minute
granules. Luciferine exists in the blood; and the light is actually
evoked by the entry of blood into the luminous organ.

We have given to this family the extension assigned to it by


Schiödte. Leconte and Horn also adopt this view, except that they
treat Throscides as a distinct family. By most authors Eucnemides,
Throscides, and Cebrionides are all considered distinct families, but
at present it is almost impossible to separate them on satisfactory
lines. The following table from Leconte and Horn exhibits the
characters of the divisions so far as the imago is concerned:—

Posterior coxae laminate; trochanters small.


Labrum concealed; antennae somewhat distant from the eyes, their
insertion narrowing the front .......... Eucnemides.
Labrum visible, free; antennae arising near the eyes under the frontal
margin .......... Elaterides.
Labrum transverse, connate with the front.
Ventral segments six; claws simple; tibial spurs well developed. ..........
Cebrionides.
Ventral segments five; claws serrate; tibial spurs moderate. ..........
Perothopides.
Posterior coxae not laminate; trochanters of middle and posterior legs very
long ..........Cerophytides.

Fig. 135—Larva of Fornax n. sp. Hawaii. A, Upper side; B, under side:


s s, position of spiracles; C, head more enlarged; D, under side of
terminal segment; a, anus.

Throscides are considered to be distinguished by the mesosternum


being impressed on each side in front for the accommodation of the
posterior face of the front coxae. The genus Throscus has the
antennae clavate. The classification of the Elaterides and these
forms is a matter of the greatest difficulty, and, if the larvae are also
considered, becomes even more complex. Cebrionid larvae are
different from those of any of the other divisions, and possess
laminate, not calliper-like, mandibles. The larvae of Eucnemides
(Fig. 135) are very little known, but are highly remarkable, inasmuch
as it is very difficult to find any mouth-opening in some of them, and
they have no legs. The other divisions possess very few species
compared with Elaterides. In Britain we have about sixty species of
Elaterides, four of Throscides and three of Eucnemides; Cerophytum
was probably a native many years ago. Neither Perothopides nor
Cebrionides are represented in our fauna; the former of these two
groups consists only of four or five North American species, and the
Cerophytides are scarcely more numerous.
Fam. 61. Buprestidae.—Antennae serrate, never elongate;
prothorax fitting closely to the after-body, with a process received
into a cavity of the mesosternum so as to permit of no movements of
nutation. Five visible ventral segments, the first usually elongate,
closely united with the second, the others mobile. Tarsi five-jointed,
the first four joints usually with membranous pads beneath. This
family is also of large extent, about 5000 species being known. Many
of them are remarkable for the magnificence of their colour, which is
usually metallic, and often of the greatest brilliancy; hence their wing-
cases are used by our own species for adornment. The elytra of the
eastern kinds of the genus Sternocera are of a very brilliant green
colour, and are used extensively as embroidery for the dresses of
ladies; the bronze elytra of Buprestis (Euchroma) gigantea were
used by the native chieftains in South America as leg-ornaments, a
large number being strung so as to form a circlet. The integument of
the Buprestidae is very thick and hard, so as to increase the
resemblance to metal. The dorsal plates of the abdomen are usually
soft and colourless in beetles, but in Buprestidae they are often
extremely brilliant. The metallic colour in these Insects is not due to
pigment, but to the nature of the surface. Buprestidae appear to
enjoy the hottest sunshine, and are found only where there is much
summer heat. Australia and Madagascar are very rich in species and
in remarkable forms of the family, while in Britain we possess only
ten species, all of which are of small size, and nearly all are
excessively rare. The family is remarkably rich in fossil forms; no
less than 28 per cent of the Mesozoic beetles found by Heer in
Switzerland are referred to Buprestidae.

Fig. 136—A, Larva of Euchroma goliath (after Schiödte); B, imago of


Melanophila decostigma. Europe.
The larvae (Fig. 136, A) find nourishment in living vegetable matter,
the rule being that they form galleries in or under the bark of trees
and bushes, or in roots thereof; some inhabit the stems of
herbaceous plants and one or two of the smaller forms have been
discovered to live in the parenchyma of leaves. A few are said to
inhabit dead wood, and in Australia species of Ethon dwell in galls
on various plants. Buprestid larvae are of very remarkable shape,
the small head being almost entirely withdrawn into the very broad
thorax, while the abdomen is slender.[135] A few, however, depart
from this shape, and have the thoracic region but little or not at all
broader than the other parts. The larvae of Julodis—a genus that
inhabits desert or arid regions—are covered with hair; they have a
great development of the mandibles; it is believed that they are of
subterranean habits, and that the mandibles are used for burrowing
in the earth. Only the newly hatched larva is, however, known.

Series IV. Heteromera.

Tarsi of the front and middle legs with five, those of the hind legs with
four, joints.

This series consists of some 14,000 or 15,000 species. Twelve or


more families are recognised in it, but the majority of the species are
placed in the one great family, Tenebrionidae. The number of visible
ventral segments is nearly always five. Several of the families of the
series are of doubtful validity; indeed beyond that of Tenebrionidae
the taxonomy of this series is scarcely more than a convention. The
larvae may be considered as belonging to three classes; one in
which the body is cylindrical and smooth and the integument harder
than usual in larvae; a second in which it is softer, and frequently
possesses more or less distinct pseudopods, in addition to the six
thoracic legs; and a third group in which hypermetamorphosis
prevails, the young larvae being the creatures long known as
Triungulins, and living temporarily on the bodies of other Insects, so
that they were formerly supposed to be parasites.
Fam. 62. Tenebrionidae.—Front coxae short, not projecting from
the cavities, enclosed behind. Feet destitute of lobed joints. Claws
smooth. This is one of the largest families of Coleoptera, about
10,000 species being already known. A very large portion of the
Tenebrionidae are entirely terrestrial, wings suitable for flight being
absent, and the elytra frequently more or less soldered. Such forms
are described in systematic works as apterous. Unfortunately no
comprehensive study has ever been made of the wings or their
rudiments in these "apterous forms."[136] it is probable that the
wings, or their rudiments or vestiges, always exist, but in various
degrees of development according to the species, and that they are
never used by the great majority of the terrestrial forms. Many of the
wood-feeding Tenebrionidae, and the genera usually placed at the
end of the family, possess wings well adapted for flight. The apterous
forms are chiefly ground-beetles, living in dry places; they are very
numerous in Africa, California, and North Mexico. Their colour is
nearly always black, and this is probably of some physiological
importance; the integuments are thick and hard, and if the wing-
cases are taken off, it will be found that they are usually more or less
yellow on the inner face, even when jet-black externally; the external
skeleton is very closely fitted together, the parts that are covered
consisting of very delicate membrane; the transition between the
hard and the membranous portions of the external skeleton is
remarkably abrupt. These ground-Tenebrionidae form a very
interesting study, though, on account of their unattractive
appearance, they have not received the attention they deserve.

Fig. 137—Tenebrio molitor. Europe, etc. A, Larva (meal-worm); B, pupa


(after Schiödte); C, imago.

Many of the Tenebrionidae, notwithstanding their dark colours, are


diurnal in habits, and some of them run with extreme velocity in

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