A field guide to the tiger beetles of India. Identification and biology of the Cicindelidae [1/1, 1 ed.] 9788121109338

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A FIELD GUIDE TO THE

TIGER

BEETLES OF INDIA Identification and Biology of the Cicindelidae

David L. Pearson | Jürgen Wiesner | Robert E. Acciavatti | V. P. Uniyal Alexander Anichtchenko

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

© 2020, Author and Publisher

All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or transmitted

in any form or by any means, electronic, mechanical, photocopying, recording or otherwise, without prior

permission of the copyright owner. Applications for such permission, with a statement of the purpose and extent of the reproduction, should be addressed to the publisher.

ISBN: 978-81-211-0933-8

Published by Abhimanyu Gahlot for Messrs Bishen Singh Mahendra Pal Singh, 23-A, New Connaught Place, Dehradun, India and Printed at Shiva Offset Press and composed by Doon Phototype Printers, 14, Old Connaught Place, Dehradun, India.

Preface

Tiger beetles are arguably one of the most intensively studied beetle groups in the

world. With nearly 2900 species described worldwide, this group presents enough diversity to pose interesting biological questions for scientists to investigate. Equally

important, this group offers amateurs a life-long hobby to observe and document the species they encounter. Both scientists and amateurs, then, share the challenge

of understanding tiger beetle diversity and the ecological role they play in a habitat or region. There are 241 species of tiger beetles found in India, the third highest number

of any country in the world. Impressively, 118 of these species are found only within the borders of India. Although more than 130 scientific articles have been published

that focus on or include Indian tiger beetles, details of distribution, behaviour and

seasonality for many species remain incomplete. In addition, over the last 30 years 49 new tiger beetle species have been described from India, and undoubtedly more species of these beetles remain to be discovered in corners of India as yet not fully

explored.

Experience shows us that one of the fastest ways to gain natural history

knowledge of a group of species is by recruiting more observers. Field guides such as this one are among the most effective recruiting tools. Increasing numbers of regional field guides about tiger beetles across the world have shown these beetles

to be practical organisms for scientific studies. Just as importantly, these beetles also attract many young people, established naturalists, and outdoor enthusiasts, who,

when given the means to recognize and easily identify tiger beetles, become tiger

beetle aficionados. These enthusiastic amateurs advance our knowledge of Nature around the world to a level that would be impossible for the professionals to do on their own.

We strove to write this field guide in a pleasant and comprehensible style that limits scientific jargon and concepts. Even more basic, however, was resolving

simple problems such as developing an acceptable list of English common names. This alternative is much less likely to distract budding amateurs who often perceive

scientific Latin names as ponderous or even incomprehensible.

We also knew that the colour identification plates would be essential for this field guide to meet its goals. These plates needed to be artistically pleasing as well as scientifically accurate if the field guide was to be useful and attract many new

enthusiasts. With real specimens and the magic of digital photography and Adobe

Photoshop©, we painted the illustrations with pixels in all their glory and colour,

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

but without sacrificing identification details. For a few rare species, however, either

specimens had been lost from museums, or the only available specimens were badly damaged. In these cases we chose to include simple line drawings from their original descriptions to serve as an aid in identification.

Many of the distribution maps included for each species are likely incomplete, but that serves to emphasize the dynamic function of this field guide. As a growing number of professional and especially amateur contributors provide more up to

date information on range extensions, behaviour, habitat details, seasonality, and

perhaps additional species, this field guide will need to be revised. A next edition will be necessary to include all the new information. Thus, beyond serving to make

identification easier, field guides have two other major functions. They help define the level of knowledge for a group of species, and they also promote interactions

between professionals and enthusiastic amateurs. To write this field guide, we relied not only on our combined 150 years of

experience studying tiger beetles, but we unabashedly drafted the help of many

of our colleagues in India, Europe and North America. They generously shared their

knowledge and interest in tiger beetles, so that we could assemble as complete a picture as possible about the identification, distribution, natural history, and habitat details of the 241 species of tiger beetles occurring in India.

Those who provided indispensable data, information and technical assistance,

included: Ditta Amran Balke, Michael Balke, Lutz Behne, Uttaran Chakrabarti, Thierry Deuve, Francois Fournier, Beulah Garner, Hans Huijbregts, Bernd Jaeger,

Charles Kazilek, A. V. Matalin, Karla Moeller, Katja Neven, Arajush Payra, Harald

Schillhammer, Mandy Schróter, Amar Paul Singh, Jeffrey Sneed, Azadeh Taghavian, Emmanuel F. A. Toussaint, Isabelle Zürcher, and many citizen scientists on Facebook-lnsectlndia and Facebook-Friends of Coleóptera at the Natural History

Museum (Entomology). Specimens for study and photography were graciously loaned to us by Arizona State University, Natural History Collections, Tempe, Arizona, USA; Carnegie

Museum of Natural History, Pittsburgh, Pennsylvania, USA; Museum Geneve, Geneva, Switzerland; Muséum National d'Histoire Naturelie, Paris, France; Museum für Naturkunde, Berlin, Germany; Natural History Museum, London, UK; Naturalist Museum, Leiden, Netherlands; Naturhistorisches Museum, Basel, Switzerland;

Senckenberg Deutsches Entomologisches Instituí, Müncheberg, Germany; Societé

d' histoire naturelle Alcide d' Origny, Aubiére, France;

University of Agricultural

Sciences-Bangalore, India; Wiener Naturhistorische Museum, Vienna, Austria; Zoologische Staatssammlung München, Munich, Germany; Zoological Survey of

India, Kolkata, Forest Research Institute, Dehradun and Wildlife Institute of India,

Dehradun. Photographs of live tiger beetles in the field were generously supplied by

Anubhav Agarwal, H. Ashida, V. C. Balakrishnan, C. R. Brown, Jeevan Jose, Milind

Preface

Joshi, Kesavamurthy - Birdwing Photography, Shyamal Lakshminara, E. S. Ross,

Vivek Sarka, Nagaraj Shastri, S. Spomer, Mohith K. Udupi, V. P. Uniyal, and Nagraj Veeraswami. Michael Kippenhan and Karen Prather produced several of the line drawings.

David L. Pearson School of Life Sciences Arizona State University Tempe, Arizona 85287-4501 USA

Jürgen Wiesner Dresdener Ring 11

D-38444 Wolfsburg, Germany

Robert E. Acciavatti Research Associate, Section of Invertebrate Zoology

The Carnegie Museum of Natural History Invertebrate Collection 4400 Forbes Avenue Pittsburgh, Pennsylvania 15213-4080 USA

V. P. Uniyal Wildlife Institute of India Chandrabani, Dehradun, Uttarakhand, India 248 001

Alexander Anichtchenko Daugavpils University, Institute of Life Science and Technologies, Vienibas iela 13-229, Daugavpils, LV-5401 Latvia

Contents

Preface

iii

Chp. 1 The Fascination of Tiger Beetles

01

Chp. 2 Howto Recognize a Tiger Beetle

05

Chp. 3 Phylogeny and Taxonomy

11

Chp. 4 Illustrated Keys to Adult Genera and Species

13

Chp. 5 Identification of Tiger Beetle Larvae

25

Chp. 6 Species Accounts

29

Metallic Tiger Beetles, Genus Grammognatha

30

Arboreal Tiger Beetles, Genus Tricondyla

31

Bark Tiger Beetles, Genus Derocrania

34

Primitive Brush Tiger Beetles, Genus Protocollyris

37

Brush Tiger Beetles, Genus Neocollyris

40

Large Brush Tiger Beetles, Genus Collyris

77

Knobby Tiger Beetles, Genus Therates

80

Wrinkled Tiger Beetles, Genus Rhytidophaena

87

Apothecary Tiger Beetles, Genus Prothyma

88

Jungle Tiger Beetles, Genus Heptodonta Glossy Tiger Beetles, Genus Pronyssa

Beautiful Tiger Beetles, Genus Calochroa

90

91 94

Temperate Tiger Beetles, Genus Cicindela

103

Margined Tiger Beetles, Genus Calomera

111

Ornamented Tiger Beetles, Genus Cosmodela

118

Small-eyed Tiger Beetles, Genus Plutacia

124

Reticulated Tiger Beetles, Genus Lophyra

125

Woolly Tiger Beetles, Genus Chaetodera

134

Tethys Tiger Beetles, Genus Jansenia

136

Pygmy Tiger Beetles, Genus Glomera

162

Lustrous Tiger Beetles, Genus Setinteridenta

164

Rounded-thorax Tiger Beetles, Genus Cylindera

165

Hook-legged Tiger Beetles, Genus Myriochila

197

Hairy-sided Tiger Beetles, Genus Salpingophora

202

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Flat-eyed Coastal Tiger Beetles, Genus Hypaetha

203

Thick-thighed Tiger Beetles, Genus Callytron

205

Enigmatic Tiger Beetles, Genus Apteroessa

207

Chp. 7 Ecology and Behaviour

271

Chp. 8 Biogeography

281

Chp. 9 Conservation

285

Chp. 10 Observing and Studying Tiger Beetles

291

Selected Bibliography

295

Checklist of the Species of Tiger Beetles of India

299

Index

307

The Appeal of Tiger Beetles

1

From North and South America to Asia and Australia a passion for tiger beetles has taken many otherwise normal people into an exciting hobby. These enthusiasts can be found on weekends and holidays searching for tiger beetles along sandy ocean

beaches, on the floor of teak plantations, and across alpine meadows. Others can be found crawling on elbows and knees in the piercing heat of Rajasthan for close-up

photos of a green species. A few sit or kneel patiently on the white sands of a Goa ocean beach taking notes about the foraging behavior of a long-legged tiger beetle

running in amongst sun bathing tourists. Tiger beetles elicit a passion in people that can last a life time. Only a few of these tiger beetle devotees are professional entomologists. Most are non-scientists, such as bankers, lawyers, and policemen, who are passionate enough about their

hobby to fill their weekends, vacations, and any time off with pursuits of six-legged

adventure and discovery. Reasons for this passion about tiger beetles are varied, but all seem to be surrounded by mystery. There is likely to be some kind of physical attraction produced by tiger beetles, but it difficult to define exactly what it is and how it has captured the interest of so many people regardless of whether they are being paid for it or not. All we know is that this depth of interest and curiosity is not logical.

You have to personally encounter tiger beetles to discover if this passion lies inside you, too. For some, it is seeing a group of spectacularly-coloured specimens pinned in perfect rows in a glass-topped insect drawer. For many others, it is noticing a live tiger beetle for the first time as a flash of bright color along a forested path, or

perhaps pursuing several, gaudy individuals along a river bank, or suddenly spying a cryptically coloured specimen in some other place that you may have been taken for granted until this moment. Whatever first caught a person's attention, we know that people with an interest in tiger beetles have felt fulfilled in the pursuit of their passion, even if the uninitiated may see it as little more than an obsession. The main reason we wanted to write this field guide was to persuade others that tiger beetles are special. The book should energize an interest in tiger beetles by amateurs and professionals alike because it serves as the first combined source of illustrated identification, natural history, and distribution for all the species of tiger beetles known to occur in India.

To organize this book we relied on popular styles of tiger beetle field guides previously published from other parts of the world. These guides have proven to be an important means to popularize more technical material developed by specialists and communicate information to a wider audience of amateur readers. The two publications we most heavily relied on were A Field Guide to the Tiger Beetles of the

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

United States and Canada (Oxford University Press) and "Tiger beetles (Coleóptera: Cicindelidae): Their history and future in Indian biological studies" a chapter

published in the book Indian Insects: Diversity and Science (Taylor and Francis Group). Colour images of each species will allow readers to make direct comparison of beetles seen in the field or a mounted specimen in a collection. Although challenging, direct observations of live specimens with binoculars can be very useful when a specimen is spotted. Photographing that specimen while alive in its natural

habitat is another effective way to "capture" a tiger beetle in order to compare it with the field guide pictures. For others who might consider these methods of comparing specimens with pictures too "hit-and-miss," we provide simple keys to work through while examining actual specimens or their images. Larval burrows may be observed

in or near the habitat where adult tiger beetles are seen. However, because the larval stages of Indian tiger beetles are so poorly known, we include only a brief overview of them.

In addition, detailed species accounts are provided, and they include information on behaviour, habitat, similar species, and other data useful for recognizing species

and when to expect to find them in the field. We also indicate which Indian species are so poorly known that any new information would be valuable. As an even greater enticement to explore hidden corners of India, or even in well-known areas, undoubtedly there are new species waiting to be discovered. Distribution maps for each species and subspecies provide valuable information as to the geographical area in which a species has been found. This information is

depicted with dots where their presence has been documented within India. Almost 2900 species of tiger beetles have been described to date, and they are found all over the world's land surface except Antarctica, the Arctic north of 65°

latitude, and some isolated oceanic islands like Hawaii in mid-Pacific Ocean and the Maldives in mid-lndian Ocean. Tiger beetles are present across an elevation range from about 3500 m above sea level to 220 m below sea level. Scientific studies since

the mid-1900s concerning their natural history, population dynamics, communities, patterns of worldwide species richness, and taxonomy of particular subgroups have produced a wealth of biological information. Tiger beetles, as a result, are among the most widely investigated groups of insects. The details of this wealth of biological

information have been synthesized in Tiger Beetles: The Evolution, Ecology, and Diversity of the Cicindelids (Cornell University Press), which serves as a companion book to this field guide. Presently, India, with 241 described species of tiger beetles, has the third highest number in the world. Only Indonesia with 301 and Brazil with 257 have more known species. Five published monographs that compile unique characters, distributional data, habitats, behaviour, revise nomenclature for entire subgroups of Indian tiger beetles have made identification and diversity patterns within India available to scientists and those with access to international libraries. However, for non-professional enthusiasts, this information has not been readily available until this field guide was created. 2

The Appeal of Tiger Beetles

While some limited information and colour photos of a small number of Indian

tiger beetle species are on line, this field guide strives to make information on all the tiger beetles of India easily available to the public, as well as to scientific institutions,

universities, libraries and schools.

In this day of genomic studies, stem cells, and molecular clocks, a field guide to a family of beetles may seem old fashioned to some. However, this field guide

provides the means of spanning the small jump from cellular studies to naming a species and understanding its distribution and natural history. Tiger beetles have already proven themselves as ideal models for understanding many other parts of the biosphere that are themselves so complicated that they hinder advances in our knowledge. Studies of tiger beetle mitochondrial and nuclear DNA have begun to open doors of comprehension that extend to many other organisms. New insights into tiger beetle taxonomy and evolutionary relationships based largely on recent studies of molecules have been included in this field guide.

Investigations of reflectance mechanisms of tiger beetle colours, ultrasound hearing, visual systems, exceptional genetic anomalies, and patterns of species distributions across continents have all been pioneered using tiger beetles. However, without names to attach to species and without basic knowledge of natural history

and distribution, these and other sophisticated studies would be much more difficult if not impossible. There are many who need less justification for a field guide, because they simply want to learn the name of that colourful tiger beetle they have seen, and hopefully even photographed, while it sits on a leaf or moves within their net after its capture. For them, basic procedures, such as putting names on living or dead specimens, can be a means of deepening a relationship between observer and tiger beetle, as well as facilitating communication among enthusiasts about their

experiences. We have also partnered with the Ask-A-Biologist program at Arizona State University in the United States. The website is visited by millions each year and

anyone with an internet connection on a computer or phone can now point their

browser to the new tiger beetle section, askabiologist.asu.edu/explore/tigerbeetles, to keep them up to date or share the link with a new enthusiast.This section has been developed to complement the field guide by providing sections for kids

and recent initiates to engage a new generation of tiger beetle enthusiasts.

HowTo Recognise An Adult Tiger Beetle

2

Although there has been little disagreement that tiger beetles are a distinct group within the beetle Order Coleóptera, how that group fits into the evolutionary tree has been a controversial subject. Experts have interpreted the data variously to show that tiger beetles form the subfamily Cicindelinae, or the supertribe Cicindelitae

or even the tribe Cicindelini within the family of ground beetles called Carabidae.

Others refer to them as Cicindelidae, a separate family of their own. After years of vacillating back and forth, the most recent molecular and morphological analyses place them in their own family, which is how we treat tiger beetles in this field guide. Tiger beetles (Cicindelidae) share many characters with four other closely related families placed together within the Coleóptera suborder called Adephaga. These related families are the predaceous ground beetles (Carabidae), predaceous

diving beetles (Dytiscidae), whirligig beetles (Gyrinidae), and crawling water beetles (Haliplidae).

For the Cicindelidae, there are twenty-seven genera now recognized as occurring in India. All possess the following characters in combination that will reliably distinguish tiger beetles from all other families of Adephaga: 1) long, sickle­ shaped mandibles; 2) simple teeth arranged along the inner side of the mandible with a compound (molar-like) tooth on the inner base of each mandible; 3) long, thin antennae with 11 segments attached to the head between the eye and base of the mandible; 4) long body form with eyes and head together wider than the thorax; 5) long thin running legs; 6) sedentary, tunnel-building behaviour of the larvae; and 7) peculiar forward-facing sets of hooks on the backs of the larvae. Adult tiger beetle species vary primarily in body proportions, size, colour and

shape of markings. In India the smallest species is 5.5 mm long, while the largest is 26 mm. Some species are dull black or brown, but many species are brilliantly emblazoned with bright green, violet, blue, red and yellow colours. All species have long slender legs for fast sprinting across the ground or vegetation. Their body shapes vary from short and stocky to long and slender with streamlined bodies resembling ants. The prominent eyes of these visual hunters are usually so large that

they make the head wider than the relatively narrow thorax. Adults have transparent hind wings that are folded and hidden under the hard front wings, the elytra. In flight, these elytra conveniently open forward to allow the flight wings underneath to unfold

and extend out to the sides. The flight wings are used for short and low escape flights from predators. A few species, however, use their flight wings for long-range dispersal, while a few others have lost these flight wings and are earthbound. On the ground, where most species spend their lives, tiger beetle adults typically run in short, but fast spurts interspersed with brief stops. The stops are necessary

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

because they literally run so fast that they cannot see their prey while running. During their stops, they search for moving insects. If they see a potential prey item, such as an ant, small spider or fly, the tiger beetle quickly turns in that direction and waits for another movement. The tiger beetle then runs the prey down and, if successful, grabs it with its long, thin, sickle-shaped mandibles. These mandibles are used to

chew the prey into a puree. The beetle's mandibular glands near the base of each mandible release enzymes that begin the digestion process. The fluid is conducted from the gland to the mandibular tip and teeth via a groove. This chewing-tobacco­ like substance is also used in defence, and anyone who has collected tiger beetles with a light-coloured net bag soon has a mesh stained with brown spots. A collector who is unlucky enough to have a larger tiger beetle mandible break the skin of his or her thumb also knows this fluid can cause a stinging sensation.

Body Parts Our keys for identifying adult tiger beetle species use simple language and are illustrated wherever possible. However, some basic knowledge of anatomy is necessary, and we include here a simple overview of adult and larval structural

features. These illustrations and brief explanations serve best as a reference dictionary to make technical terms shown in bold more easily understood.

The hard armor-like skin (cuticle) that covers the adult tiger beetle is critical for survival, but it offers features very useful in identification. The outermost layer

(epicuticle) has patterns of tiny pits, larger punctures, ridges and undulations (microsculpture). The differences in these patterns of microsculpture are frequently used to distinguish tiger beetle genera and species. The cuticle is laminated with layers of melanin pigment and translucent waxes that alternately reflect and pass

light. The microscopic distance between these thin alternating layers produces a broad range of metallic colours by bending wavelengths of light in various ways

(reflectance and interference). The degree of uniformity of the cuticular reflector determines the purity of reflected colour. Highly sculptured and non-uniform reflectors produce a broad blend of colours of different wavelengths reflected at various angles from different locations. This type of integument gives rise to dull

green or brown colours similar to those made by pigments in other insects. In bright iridescent species, the cuticular sublayers are more uniform, and the surface is relatively smooth (Plate 31.1). In all-black species, like those of the genera of Arboreal Tiger Beetles (Tricondyla) and Bark Tiger Beetles (Derocrania), the melanin

is deposited in relatively thick and disorganized patterns that absorb most light. In other species, parts of the integument have no pigment deposited, and these areas are pale yellow or white.

The most frequently studied anatomical features of adult tiger beetles include the head (Fig. 2.1) which presents many distinctive structures each with a variety of characteristics, including: 1) antennae long thread-like (filiform) with 11 segments (colour, distribution of hair-like setae, relative overall length) used primarily as 6

HowTo Recognise An Adult Tiger Beetle

Supraorbital seta

Vertex

Antennal segment

Eye Frons Sensory seta Clypeus

Scape

■Gena

•Labrum

_ ___ Maxillary palpus

Submarginal seta, of labrum

Labial palpus

Mandible

Figure 2.1

Median tooth of labrum

Head (front view) of adult tiger beetles with structures identified (Reprinted from Tiger Beetles: The evolution, ecology and diversity of the cicindelids, by Pearson & Vogler © 2001 by. Cornell University. Used by permission of the publisher, Cornell University Press).

tactile sense organs; 2) mandibles (relative length, number and position of "teeth") used for capturing and processing prey and in males for grasping females during mating; 3) upper lip (labrum) (colour, length-width ratio, number and position of "teeth," number and position of setae) used with the mandibles to help grasp and

process prey; 4) first-paired mouthparts under mandibles (labium) (presence and position of setae, relative length and colour of the segments of its finger-like palpi); 5) second-paired mouthparts under mandibles (maxillae) (relative length and colour of the segments of its finger-like palpi) used to manipulate and analyze the quality of food items; 6) compound eyes (degree of bulging, relative size); 7) complexity of surface microsculpturing and depth of grooves (rugae) between the eyes and other parts of the head; 8) distribution of hairs (setae)(white or dark, thin or thick, erect or

flattened) that may function as sense organs and/or insulation against hot surfaces.

Various adult eye sizes and shapes produce variable areas of three dimensional

(stereoscopic) vision. Nocturnal adults such as those of Metallic Tiger Beetles (Grammognatha), have small, relatively flat eyes compared to the large, bulbous eyes of genera active during the light of day.

On the thorax (Fig. 2.2), the most frequently observed distinctive features are found in the relative length and width proportions of the thorax (rectangular, square or

elongate), its shape as viewed from above (cylindrical, parallel, rounded, trapezoid, and so on), the texture (shiny metallic or dull) and colour of its upper surface (pronotum), and the patterns or absence of setae on its side, lower or upper surfaces. Behind the pronotum (Fig. 2.2), are situated the hardened modified front wings

(elytra), that cover both the hind wings (membranous for flight) and top of the abdomen. These elytra are spread and rotated forward in flight where they may

function as airfoils, but do not flap. Elytra offer the most obvious, and therefore, 7

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Figure 2.2 Body (dorsal view) of adult tiger beetles with structures identified ((Reprinted from Tiger Beetles: The evolution, ecology and diversity of the cicindelids, by Pearson & Vogler © 2001 by Cornell University. Used by permission of the publisher, Cornell University Press).

the most commonly used identification characters to distinguish the majority of

tiger beetle species. These elytra characters include: 1) a variety of surface textures

(microsculpture) that include large individual punctures (foveae), patterns of small pits (punctation), grooves (rugae), smooth (impunctate) areas, or raised areas (undulations) and tiny saw-like teeth (microserrations) and spines on the rear edge of the elytra; 2) shape viewed from above (parallel-sided, rounded, oval); 3) profile viewed from the side (domed, flattened); 4) dark background colour and texture

(shiny, metallic, dull); 5) the pattern, colour, and position of spots, lines and stripes (maculations) or their absence (immaculate). The flight wings are membranous with a distinctive framework of thickened

ridges (veins). Modifications of vein structures allow for a triple folding so that the flight wings can be stored completely under the elytra. In some flightless species, these flight wings are shrunk or even totally absent with permanently fused elytra. The prominent and protruding legs (Fig. 2.3) of tiger beetles are thin, and long for fast running. Leg colour, setae abundance (thick or sparse), position along

segments, and type (erect or flattened, straight or hooked) and relative leg size (both to the overall size of the tiger beetle, as well as, between the lengths of the several segments comprising the same leg) are sometimes important as identifying

characters. Males of all species have thickened white pads of long curved setae on several of their segmented feet (tarsi) of the front legs. 8

HowTo Recognise An Adult Tiger Beetle

Figure 2.3 Front (prothoracic) leg of adult tiger beetles with structures identified (Reprinted from Tiger Beetles: The evolution, ecology and diversity of the cicindelids, by Pearson & Vogler © 2001 by Cornell University. Used by permission of the publisher, Cornell University Press).

The form and shape of the mating structures in tiger beetles have been thoroughly studied and

documented, and consequently are well known and have been used

extensively in taxonomic comparisons. The extensible penis (Fig. 2.4) is located within a hardened (sderotized) sheath (aedeagus). The general shape

of the aedeagus, the form of its tip Figure 2.4 Tiger beetle adult male: external and the shape and position of the reproductive structures and aedeagus (Reprinted sclerotized rings and internal elements from Tiger Beetles: The evolution, ecology and diversity of the cicindelids, by Pearson & Vogler © are distinctive for many species. All 2001 by Cornell University. Used by permission of these structures give clues, not only the publisher, Cornell University Press). to species identification, but also to phylogenetic relationships among species, and as a consequence, have formed much of the basis for their classification. When the aedeagus is not extended, it is maintained internally with the right side down. Upon extension it rotates 90° clockwise. At the tip of the extended aedeagus there is a less sclerotized area (internal sac). This internal sac is turned inside out during copulation to extend

beyond the tip of the hard aedeagus, and in this manner the male delivers sperm or packets of sperm (spermatophores) to the female. Due to the difficulty of observing 9

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

characters of the aedeagus and internal sac in a live beetle, few of these structures have been included as part of the identification keys in this field guide.

The female mating structures (Fig. 2.6) also include some specialized characters. The eighth and ninth abdominal segments are modified to form a telescopic ovipositor, which is used to insert eggs, one at a time, into the substrate. Species differ considerably in the form and shape of the ovipositor, especially the terminal portion

(gonapophysis). The differences in these characters have only begun to be studied but offer the potential for many insights into phylogeny, ecology, and behaviour.

Figure 2.5 Tiger beetle adult male: internal aedeagus structures for two, closely related tiger beetle species showing differences in their general shape, and the internal elements, a, arciform piece; b, stylet; c, shield; d, “stick”. Scale bar = 1 mm (Reprinted from Cassola and Pearson 1999 Annali del Museo Civico de Storia Naturale “G. Doria” 7:1-12).

Figure 2.6 Tiger beetle adult female: external reproductive structures and extended ovipositor (Reprinted from Tiger Beetles: The evolution, ecology and diversity of the cicindelids, by Pearson & Vogler © 2001 by Cornell University. Used by permission of the publisher, Cornell University Press).

10

Systematics and Taxonomy

3

Studies of evolutionary relationships help us understand connections among

tiger beetle groups, such as genera and species, based on form, behaviour, and

distribution, a field of study we call systematics. These systematic studies allow researchers to test which groups of tiger beetles are most closely related to other groups. We call this type of extended family tree a phylogeny. In addition to the evolutionary insight they provide, we also use phylogenies to conveniently arrange species into named groups organized and categorized on the basis of their relatedness, a field called taxonomy. This pattern of organization of named groups is termed nomenclature, and it is constantly being tested and modified as more data come in. The advantage of nomenclature is that it provides an effective basis to communicate names that everyone will understand, whether it is the Latin scientific name of genus and species or the common English names that non-scientists often prefer.

As more and more species were described, they were organized into groupings and subgroupings of species that represented evolutionarily related species. Because there are so few tiger beetle fossils on which to base these groups, alternatives were

used, such as shapes and sizes of adult body parts (morphology), similarities in male genitalia, body colour patterns, numbers of sex chromosomes, and more recently

molecules like DNA. For Indian taxa of tiger beetles, their names and branches (clades) are thought to represent their real evolutionary pathways, but as with all scientific endeavors,

they are subject to reinterpretation and change as additional information becomes available. The genera of Arboreal Tiger Beetles (Tricondyla), Bark Tiger Beetles (Derocrania) and Brush Tiger Beetles (Collyris, Protocollyris, and Neocollyris) form distinctive evolutionary groups all considered generally to have been formed from a common ancestor (monophyletic). It is within the evolutionary branch of the Temperate Tiger Beetles (Cicindela) and the numerous related species groups that have been created within this genus, however, that the greatest controversy continues. Should species groups

be considered subgenera within the genus Cicindela or should the species groups actually represent genera of their own? For many taxonomists, this apparent dilemma is as much semantics as science. Here, we have reached a consensus to present the most recent interpretation of species groups as genera and subgenera

based on their scientific names in the latest published catalogues and websites covering Carabidae and tiger beetles. Although modern studies of DNA, molecular clocks, sophisticated statistical

programs, as well as more intense field collecting and observations all provide us with

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

a better idea of the phylogeny and classification of tiger beetles, our interpretation of some relationships will likely change in the future as new information is obtained. With each successive investigation we come one step closer to understanding the actual evolutionary process that resulted in the tiger beetles we encounter within India today.

What are species? There are many definitions about the concept of a species. Even within any of these definitions, however, different interpretations can influence how we might

understand such a concept as applied to populations of organisms. A vital part of this understanding about exactly what constitutes a species is realizing that it is an ongoing and dynamic process of gathering, analyzing, and interpreting biological

data about populations. At any one time, there are examples of many levels of how

different two populations have become in terms of their genes, ecology, behaviour, and physical appearance. At what point these differences are sufficient to distinguish them as separate species is often a personal matter that resides in the eye of the beholder. Because naming a species can involve a range of interpretations, we have decided to use the names most widely accepted for tiger beetles as published by expert taxonomists. However, we also recognize that each of these species decisions is a hypothesis based on the most current information. As more biological details and analytical techniques become available, some of the named species could be "split" or "lumped" because a new, more insightful hypothesis will have replaced the old one.

What are subspecies? Some species, especially those that have broad geographical ranges, often show consistently different colours, sizes or other characters from one area to another within that range. We understand little about the factors that cause these differences or if they inevitably lead to the formation of new species. For species that show these distinctive subpopulations, we often give them names as subspecies. However, some taxonomists argue that so many of these subspecies have been named on the basis of subjective and trivial reasons that the concept of subspecies should be discarded. Fortunately, only a few species in India have subspecies described, and we include them here cautiously.

12

Illustrated keys to adult genera and species

4

Diagnostic Characters for Identification The extensive series of colour plates in this field guide provide an actual image of most Indian tiger beetle species. These images should enable both amateurs and

specialists to identify nearly all of the tiger beetles they may discover throughout

India, especially when combined with the range maps and descriptive information

in the species accounts. However, some species are so similar in colour and marking patterns that a single image will not provide a definitive identification. In these cases, the use of a taxonomic key with more detailed diagnostic characters may be needed. The keys presented in this field guide are based on those developed by Acciavatti and Pearson, Naviaux and Wiesner, but we incorporate some reorganization and modifications that should allow for easier distinctions among the species. We also present less technical terminology to make the key more "user -friendly"for the non­ specialist. These characters are described in the next sections and illustrated by line drawings throughout the key. In several cases, we also rely on diagnostic geographic distribution to separate otherwise similar species. Be aware, however, that a few species, especially those within the same couplet, may be especially difficult to

recognise because their adults exhibit a considerable amount of individual variation.

Elytra and Colour The elytra are the first pair of hardened wings that cover the back part of the thorax and most of the abdomen of beetles, and they provide the structure with the most useful identifying characters. Elytra colour and their distinctive markings (maculations) readily identify many tiger beetle species, especially when combined with habitat and geographic range. Elytra colours are usually consistent within a species, but a range of distinct colour forms may occur within a species. The colour of the underside of the abdomen is usually metallic. The standard pattern of elytra markings consists of three distinct light lines (lunules) on each elytron (the complete or normal pattern), a front maculation (humeral lunule), a middle maculation (middle band) and a rear (apical lunule) maculation at the tip (Fig. 4.1). In some tiger beetle species these markings are reduced to a series of separate dots (Fig. 4.1), but on other species some or all markings may be absent (immaculate), and on still other species at the other extreme, markings can be greatly expanded and fused together to form mostly white elytra.The white line running along the outer edge of the elytra (marginal line) may partially or fully connect the three distinct markings. Within species, subspecies, and populations there can be considerable variation in the extent of these maculations.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Figure 4.1 Tiger beetle adult elytra: left elytron with complete (standard) pattern of light markings and identifying characters; right elytron with reduced markings pattern (Reprinted from A Field Guide to the Tiger Beetles of the United States and Canada Identification, Natural History, and Distribution of the Cicindelinae by Pearson et al. © 2015 by Oxford University Press. Used by permission of the publisher, Oxford Publishing Limited).

The elytra surface may be shiny, metallic, dull, or greasy in appearance depending on the species. These differences are caused by texture, primarily surface pits of variable size (seen under 40 x magnifications). Deep pits will give a shiny appearance, shallow pits or no pits will make the elytra appear dull. Sometimes

larger pits (punctures) are present, usually in rows running along the length of the middle of the elytra. These pits often reflect a different colour than the rest of the elytral surface. Another variation of surface texture is the presence of small raised bumps (granules). Some species have one or two distinctly raised ridges running

longitudinally down the middle of each elytron. Small tooth-like projections

(microserrations), seen only at 40x magnification, are present along the rear margin of the elytra in some species. Distinct expansions or flanges on the outer edge of the elytra are seen in females of a few species.

Setae The numbers and types of hair-like setae on various parts of the body are useful in distinguishing species and some of the genera. Long, thin setae are often present on the antennae, head, labrum, and on the second basal segment of the legs

(trochanters). Some species have long, thin setae bent at their ends to form hooks located along the rear margins of the longest upper segment of the legs (femora). All species have long sensory setae along the outer end of the first antennal segment,

but certain species have additional setae on that segment.

Groups of long erect setae that originate from medium or large pits are especially common in most species of the Tern perate Tiger Beetles (genus Cicindela). These setae may be worn off in older specimens, but evidence of their presence can often be deduced from the presence of these peculiar pits. Males of several species have the 14

Illustrated keys to adult genera and species

fourth antennal segment closest to the head with a few to many clubbed or thickened setae often forming an erect comb-like bundle.Thick, flattened setae densely cover the underside of the body in other groups of Indian tiger beetles, such as the Hairy-sided Tiger Beetles (genus Salpingophora) and Flat-eyed Tiger Beetles (genus Hypaetha).

Other characters The upper lip (labrum) often provides important characters for distinguishing species. These characters include: proportional length, shape, number of teeth

(usually one to seven) on the front edge, middle ridge or flattened surface, number of erect bristles and their position, uniform colour or bicoloured, surface reflection

metallic, leathery, or ivory coloured. Usually the labrum is wide and short, but in a few species it is longer than it is wide.

Body length of adult tiger beetles in India ranges from about 6 mm to over 25 mm. Although many species show considerable variation in body length and females are often significantly larger than males, each tiger beetle species usually falls within a specified range that can be useful for identification.

Identification keys The first key below will allow identification of genera and their subgroups for all tiger beetle adults found in India. Separate keys to the species within each of these genera and subgroups then follow in Chapter 6. A few variable species with distinct subspecies key out at more than one couplet.

Artificial Key to the Genera and Subgroups la.

Entire body form, especially thorax and elytra,

2

elongated and thin with small or no shoulders (Subfamily Collyrinae) (Fig. 4.2) 1b.

Entire body form compact, thorax and elytra broad

6

with distinct shoulders in outline (Subfamilies Megacephilinae and Cicindelinae) (Fig. 4.3)

Figure 4.2 Adult tiger beetle with little or no shoulders at the front, outer angle of elytra (Tricondyla).

Figure 4.3 Adult tiger beetle with distinct shoulders at the front, outer angle of elytra (Therates). 15

h Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

2a. (la.)

Flight

wings

absent

under

fused

elytra,

3

body black, forward portion of elytra sides conspicuously constricted

2b.

Flight wings present under separated elytra,

4

body metallic blue, green violet to rufous, elytra

sides parallel or near parallel along their entire

length (Fig. 4.4)

Figure 4.4 Adult tiger beetle with no shoulders at the front, outer angle of elytra (Neocollyris).

3a. (2a.)

Area between eyes forming a deep depression (Fig. 4.5), body length 16-24 mm

Key I: Arboreal Tiger Beetles, genus

Tricondyla, page 31, Plate 1

3b.

Area between eyes relatively flat (Fig. 4.6), body

length 9- 12 mm

Key II: Bark Tiger Beetles, genus

Derocrania, page 34, Plate 2

Figure 4.5 Top of head between eyes (vertex) deeply depressed (Tricondyla).

4a. (2b.)

Figure 4.6 Top of head between eyes (vertex) shallow to flat (Derocrania).

Vertex short, with a sharp curved line between it and frons (Fig. 4.7)

Key V: Large Brush Tiger Beetles, genus

Collyris, page 77, Plate 9

16

Illustrated keys to adult genera and species

4b.

Vertex long, smoothly confluent with irons (Fig.

5

4.8)

Figure 4.7 Top of head between eyes (vertex)

Figure 4.8 Top of head between eyes (vertex)

deeply depressed and separated from forehead (frons) (Collyris).

blending smoothly into forehead (frons) (Neocollyris and Protocollyris).

5a. (4a.)

Labrum short (Fig. 4.9); body smaller than 9

Key III: Primitive

mm, slender; sculpture of elytra "shallow, dense

Brush Tiger Beetles,

and uniform

genus Protocollyris, page 37, Plate 2

5b.

Key IV: Brush Tiger

Labrum longer (Fig. 4.10)

Beetles, genus

Neocollyris, page 40, Plate 2-9

Figure 4.9 Short labrum on front of tiger

Figure 4.10 Long labrum on front of tiger

beetle head. (Protocollyris).

beetle head. (Neocollyris).

6a. (1b.)

Body form robust, large (length 19-26 mm),

7

head and pronotum wide with head only slightly

narrower, eyes small not bulging outward

6b.

Body form slender, size from small to large (length

8

6 mm to over 22 mm), head variable in width but

always wider than pronotum, eyes of various sizes

and bulging outward on most species

17

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

7a. (6a.)

Body above completely iridescent, surface

Eurasian Metallic

texture fine on head and pronotum, elytra

Tiger Beetle,

elongate with smooth maroon to green and

Grammognatha

reddish brilliant metallic reflections and large,

euphratica, page 30,

broad yellow or tan lunules covering only their

Plate 1

apical ends, flight wings present 7b.

Body above completely dull, dark brownish

Enigmatic Tiger

black without reflections, surface texture of

Beetle, Apteroessa

honey-combed pits on top and side of head

grossa, page 207,

and anterior margin of pronotum, elytra oval

Plate 30

with deep pits in front half and three large

yellow lunules evenly spaced from their bases

to apical ends, flight wings absent. 8a. (6b.)

Body

surface

underside

bright

mostly

and

hairless,

polished

head

with deeply

Key VI: Knobby Tiger Beetles, genus

excavated between large bulging eyes, labrum

Therates, page 80,

with hairs inserted between anterior teeth,

Plate 10-11

pronotum globular, elytra covered with bright

colour patterns in addition to lighter bands and

two or more distinctly rounded humps near the front of each wing cover

8b.

Body surface usually dull but occasionally polished, white hairs on the underside may

be absent, exist only in small patches, or cover extensive areas, head between eyes at most only moderately excavated, labrum with hairs inserted behind anterior teeth,

pronotum square, globular, or elongate, elytra marked with discrete spots, lunules,

elongated bands, or iridescent areas rather than bright colour patterns and never with distinctly rounded humps near the front of each wing cover 9a. (8a.)

Body underside completely without white hairs or with white hairs restricted to a fringe where

the back legs attach to the body, elytra either marked with two or three small white spots on the outer margins from shoulder to tip or

unmarked

18

9

Illustrated keys to adult genera and species

9b.

Body underside with white hairs on head,

13

thorax and abdominal segments to varied extent

from total to sparse coverage, elytra usually

with a variety of markings in a multitude of

arrangements as spots, lunules forming patterns, elongated bands of various widths, or iridescent

areas, and only occasionally unmarked 10a. (9a.)

Body underside completely without white hairs,

11

labrum short or moderately long with five distinct

teeth along entire rounded anterior margin 10b.

Body underside bare except for a fringe of white

12

hairs where back legs attach to thorax, labrum elongated and large with five or seven distinct teeth only at extreme anterior edge

11a. (10a.)

Head and pronotum with distinct wavy or

Key VII: Apothecary

transverse grooves, labrum with four (rarely

Tiger Beetles, genus

five) hairs, pronotum distinctly longer than

Prothyma, page 88,

broad, body dark blue, green, or almost black 11b.

Plate 11

Head and pronotum without distinct wavy

Key VIII: Wrinkled

or transverse grooves, labrum with six hairs,

Tiger Beetles, genus

pronotum square, body bright and shiny blues,

Rhytidophaena,

greens and reds, occasionally with violet tinge

page 87, Plate 11

on elytra shoulders 12a. (10b.)

Body upperside dull olive green, or dark

Key IX: Jungle Tiger

coppery green on elytra, pronotum longer than

Beetles, genus

wide, body length 15-17 mm

Heptodonta, page 90, Plate 11

12b.

Body upperside with extremely bright metallic reflections on elytra, pronotum transversely spheroidal shaped, body length 13-15 mm

Key X: Glossy Tiger Beetles, genus Pronyssa, page 91,

Plate 11-12 13a. (9b.)

Pronotum lateral sutures present on top a

Key XXII: Flat-eyed

short distance inward from the edges; body

Tiger Beetles, genus

underside with thick white hair-like setae

Hypaetha, page 203,

covering segments 13b.

Pronotum lateral sutures only evident along the

Plate 30

14

edges, body underside for some species with more extensive setae, others with fewer setae,

and some with no setae on most segments

19

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

14a. (13b.)

Head broadly concave between two large

15

extremely flat and outward bulging eyes, elytra

markings restricted to a continuous lateral

band with or without short inward projections

or completely missing, pronotum hind angles large, acutely projecting

14b.

Head

narrowly concave

between

upward

16

protruding eyes of varied sizes from small and inconspicuous to large and prominent, elytra markings as wide separate or fused spots, or

narrow lunules with long, often curved, inward projections, rarely lacking markings, pronotum

hind angles small, subacute to obtuse 15a. (14a.)

First large segment of each leg short and stocky,

Hairy-legged

elytra lateral band wide and wavy with short

Tiger Beetle, Salpingophora

inward projections at one or two places

maindroni, page 202, Plate 29 15b.

First large segment of each leg long and slender

thighed Tiger

band uniformly thin and continuous along

Beetles, genus

the entire lateral edge or completely missing

Callytron, page 205,

depending on the species

16a. (14b.)

Key XXIII: Thick­

and swollen at its base, elytra lateral marking

Elytra marked with separate or fused spots or

Plate 30

17

elongated stripes where their bases join, along both margins, and in their centres

16b.

Elytra without markings where their bases join

18

or along their inner margins, markings only

laterally or in middle or both on most species,

markings absent on a few species 17a. (16a.)

Elytra markings as three broadly fused lunules

Key XVI: Reticulated

or widely separated spots against darker

Tiger Beetles, genus

surface, mandible with three or four acute teeth

Lophyra, page 125,

beyond basal molar, head with setae in various

Plate 18-19

patterns as only a cluster of flat setae on top of

head at back or in front of each eye on some species, or entirely absent for others

20

Illustrated keys to adult genera and species

17b.

Elytra marked with complex patterns of lunules

Key XVII: Woolly

obscuring much of the dark surface, mandible

Tiger Beetles, genus

with four acute teeth beyond basal molar, flat

Chaetodera,

white setae cover much of head between eyes

page 134, Plate 19

or restricted to patch on front above antenna

18a. (16b.)

Labrum with only four (rarely six) setae in

legged Tiger

spaced along most of posterior margins on

Beetles, genus

longest segments (femora) of front and middle legs, sparser hooked setae on back femora 18b.

Key XXI: Hook­

one row, hooked setae numerous and closely

Labrum with numerous setae (usually 8 to 15)

Myriochila, page 197, Plate 29 19

setae in one or several rows arising inward from the edge on most species, or between the teeth for one species, hooked setae absent

from longest leg segments of most species, but sparse hooked setae only at bases of front and

middle legs for one species 19a. (18b.)

Labrum short, more than ten labral setae

Key XIII: Margined

arranged in 2 or more irregular rows, elytra with

Tiger Beetles, genus

one wide, elongate or oval marking on apical

Calomera, page 111,

half at lateral margin either behind middle band

Plate 15-16

on most species, or fused to middle band on

some species, or separated from middle band on others, but for all species lateral markings

are anterior to and separated from apical lunule 19b.

Labrum

variable

in

length

and

setae

20

arrangement, but long for species with more than ten setae, various lengths on species with

ten or fewer labral setae, setae arranged in one row (on some species a few setae at middle),

elytra markings on most species at middle of

lateral margin consisting only of middle band or its narrow lateral expansion, no separate

macula at lateral margin behind middle band, a few species with a narrow elongate spot at

lateral margin either fused to middle band or to both it and apical lunule

20a. (19b.)

Medium to large beetles, body length greaterthan

21

12 mm, pronotum bulky, elytra markings large,

most species with bold spots, for some species broad bands, for other species wide stripes, on unmarked species elytra surface smooth and

lacking pits or glossy and iridescent margins 21

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

20b.

Small beetles, body length less than or equal to

24

12 mm, pronotum slender, elytra markings small, most species with oval dots, for some species narrow elongate spots, for other species thin curved lunules complete or divided, for a few

unmarked species elytra surface roughened, pits present, or margins glossy and iridescent

21a. (20a.)

Elytra dull black or greenish black with large

Key XII: Temperate

yellow or white bands, head in front and on top between eyes with numerous long erect setae

Tiger Beetles, genus

scattered over surface and two or more pairs of

Plate 14-15

Cicindela, page 103,

single or multiple setae near each eye

21b.

Elytra shiny with bright solid or polychromatic colours with bold yellow or white spots or

22

bands or unmarked, head in front and on top

without erect setae except for two pairs of single or multiple erect setae near each eye

22a. (21a.)

Labrum shiny black or pale without metallic

Key XV: Small-eyed

reflections and flattened, wide and broad with

Tiger Beetles, genus

six to ten small feeble teeth, irregularly spaced, no ridge down middle of labrum, elytra with

Plutacia, page 124,

Plate 17

middle band partly in apical half and obliquely extended onto centre or almost to suture

22b.

Labrum metallic blue, green, or copper on most

23

species, shiny black or pale with or without a longitudinal ridge at middle on other species, elytra with middle band varied, either missing or present as an oval or elongate spot oriented across or down the centre, in a few species as a

continuous or broken middle stripe

23a. (22b.)

Body size and markings varied by species, one

Key XI: Beautiful

species very large (more than 20 mm) of multiple

Tiger Beetles, genus

colours with three large yellow spots aligned on

Calochroa, page 94,

each elytron, all other species large size (15 to 20

Plate 12-14

mm) without multiple colours and elytra markings in middle as a minute dot or four large spots or

a continuous stripe longitudinally arranged or completely unmarked, labrum varied from long

with five large teeth to short with three minute teeth, pronotum and head contrasting in surface

texture, head between eyes with distinct parallel or wavy ridges in all species and on most species pronotum surface smooth and polished, but on some species finely wrinkled, but lacking parallel

ridges along midline at centre of pronotum 22

Illustrated keys to adult genera and species

23b.

Body large (15 to 20 mm), elytra with bold

Key XIV:

markings superimposed on intermingled multiple colours and iridescence for most

Ornamented Tiger

species, pronotum and head similar in surface texture, both finely to moderately wrinkled

Beetles, genus

Cosmodela, page 118, Plate 16-17

with ridges parallel along midline at centre of pronotum, labrum long with three to five acute teeth about equal in length and usually with a

middle ridge

24a. (20b.)

Elytra markings not a standard pattern, only two or three oval or slightly elongated spots on rear half at or near lateral margins on many species, or bent bands on some species, or totally

Key XVIII: Tethys Tiger Beetles, genus

Jansenia, page 136,

Plate 19-23

missing on a few others, elytra surface often brightly coloured and iridescent as wide lateral margins contrasting or blending with central

areas, elytra texture often rough and irregular with metallic foveae on the inner edges and centres contrasting with surface colour, labrum

of most species long and covering irregularly

toothed mandibles and noticeably inflated and thick segment of labial palpi, head bare and pronotum at most with only sparse setae

24b.

Elytra markings a standard pattern of four

25

or more lunules or oval spots over entire surface extending inward from narrow or fragmented lateral bands on most species,

lunules completely fused on some species, lacking entirely on a few species, elytra surface

only brightly coloured and iridescent within small lateral areas contrasting or blending

with central areas, elytra texture smooth and evenly contoured on all species except one with a rough and unmarked surface, elytra

metallic foveae if present only adjacent inner edge, labrum of most species short, exposing regularly toothed mandibles and long thin labial feelers (palpi), head and pronotum rarely without numerous setae 25a.

Elytra without pale markings only shiny copper-

Key XIX: Pygmy

green and glossy purple-black lateral areas extend inward onto smooth evenly contoured

Tiger Beetles, genus

surface, legs entirely pale and nonmetallic, smallest body size (less than 6.5 mm)

Plate 23

Glomera, page 162,

23

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

25b.

Elytra

markings on

most species distinct

26

although one or more spots may be reduced to faint dots, lateral margins solely glossy black extend inward on one species, unmarked with

rough and unevenly contoured surface on another species, legs completely or partially dark often metallic, larger body size (more than

6.5 mm) 26a.

26b.

Elytra markings reduced to two oval spots

Lustrous

laterally on rear half, elytra surface smooth

Tiger Beetle,

and highly polished, labrum with middle setae

arising at anterior margin from between teeth,

Setinteridenta rhytidopteroides,

eyes large and prominent

page 164, Plate 23

Elytra markings on most species forming more

Key XX: Rounded-

extensive patterns of spots, lunules and bands

pronotum Tiger

laterally or centrally or both but elytra of one

Beetles, genus

species without markings, elytra surface varied

Cylindera, page 165,

in texture either dull with or without non-

Plate 24-28

punctate areas or slightly shiny from punctures or rough and uneven, eyes varied from small

not prominent on some species to large and

prominently raised on many species, labrum with middle setae inserted away from anterior margin behind teeth

Biology of Tiger Beetle Larvae

The larvae of this family are unique among beetles. They all are designed for a similar life in a narrow burrow which they enlarge each time they grow by moulting through three successively larger larval forms (instars). Thus, even though the adults may be nocturnal or diurnal, long and thin or short and wide, the larvae are all white and grub-like (Fig. 5.1), with much of the outer covering of their bodies membranous. A dark armored capsule covers the head, and scattered dark plates are especially noticeable on the top of the thorax (pronotum).They have a large head with up to six small eyes on top and formidable mandibles underneath. A particularly striking and unique feature on the larva's lower back includes a prominent hump with two

pairs of large, darkened hooks that face forward.

Figure 5.1 Side view of a typical tiger beetle larvae and its morphological characters often used for identification. EU, eusternum; IH, inner hook; MH, median hook; MSN, mesonotum; MTN, metanotum; PN, pronotum; PY, pygopod; T3, third abdominal sclerite (Reprinted from Knisley & Pearson 1984 Transactions of the American Entomological Society 110:465-551. Used by permission from the American Entomological Society).

The- larvae, like adult tiger beetles, are predaceous, but unlike the adults the larvae wait

for prey to come to them. Each larva positions itself at the top of a long burrow with its head and thorax flush to the substrate surface and exactly filling the diameter of the burrow entrance (Fig. 5.2). Where larval burrows are constructed depends on the species. Larval burrows for Temperate Tiger Beetles (Cicindela) and closely

related genera are in terrestrial sites like flat soil,

Figure 5.2 Tiger beede larva at mouth of its vertical tunnel.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

vertical clay banks, or among forest leaf litter, whereas, for a tropical forest taxa, such

as, Knobby Tiger Beetles (Therates) burrows are made into rotted wood of branches and twigs, and Brush Tiger Beetles (Neocollyris) in the pith of live bushes. When a prey item approaches the burrow entrance closely, the larva extends its body, anchored by the back hooks into the side of the tunnel, and quickly reaches out backward to grab the prey in its powerful mandibles. The larva then pulls the struggling prey down into the depths of its burrow and dispatches it with a few mighty bites

(Fig. 5.3).

Figure 5.3 Tiger beetle larva foraging from its tunnel. A, waiting at the soil surface entrance hole as prey approaches; B, extending backward to capture prey as it enters the larva’s zone of capture; C, descending to the bottom of the tunnel to consume digestible parts of prey; D, throwing discarded indigestible remains of prey away from tunnel entrance (Reprinted from Faasch, H. 1968. Zoologische Jahrbücher: Abteilung fur Systematik, Geographic und Biologie der Tier 95:477-522.)

The larvae of only a few species of Indian tiger beetles have been described. But the details we do have are intriguing. Unlike most other larval tiger beetles, two Indian species have evolved distinctive differences in their tunnel construction.

Larval Golden-banded Tiger Beetles (Cicindela aurofasciata) construct a low, thick turret over the tunnel entrance above the ground's surface. It is bent over to face its opening easterly into the early morning sun (Fig. 5.4). This construction apparently enables the larva to more quickly warm-up on cool mornings, which in turn increases hunting time throughout the day. Increased food accelerates growth into the pupal

and adult stages.

26

Biology of Tiger Beetle Larvae

Figure 5.4 Curved larval turret over tunnel of the Golden-banded Tiger Beede (Cicindela aurofasciatd) on open, forest floor in peninsular India. Ao, angle of turret from horizontal; Ht, height of larval turret; Rd, reaching distance for the larva when capturing prey; Sd, depth of the pit excavated by the larva adjacent to the turret opening (Reprinted from Shivashankar et al. 1988 Coleopterists Bulletin 42:63-68. Used by permission from the Coleopterists Society).

The

extraordinarily

thin,

elongated

larvae of the arboreal Brush Tiger Beetles

(Neocollyris) burrow into the narrow stems

of undergrowth bushes, including coffee plants on plantations. Here they construct a lengthwise burrow into the pith of the branch and hunt aphids and other prey from the burrow's entrance (Fig. 5.5). Because the head and thorax are usually the same colour and texture as the surrounding soil surface, most larvae are hard to see as they wait at the top of the burrows. Their reaction when disturbed or exposed to danger is to retreat immediately down into their tunnel away from the burrow mouth.

The observer is often made aware of their presence only when a black hole suddenly seems to appear on the surface where before there was none. Larval tiger beetles have fewer and less obvious characters than adults (Fig. 5.1) to

distinguish among species. For temperate

Figure 5.5 Typical larva of the Brush Tiger Beedes (Neocollyris). A, side view, B, dorsal view, C, head; D, detail of one of the back hooks on the 8th segment; E, excavated tunnel in the pith of a stem of a coffee bush occupied by the larva (Reprinted from Shelford 1907. Ecological Entomology 55: 83-90.)

27

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

species, many subtle characters have proven to be important taxonomically for species recognition including: 1) shape and relative size of the inner and median hooks on the back of the fifth abdominal segment; 2) relative size, number and

placement of the simple eyes (stemmata); 3) relative lengths of the segments of the short antennae; 4) size and shape and the presence of ridges on the head and dorsal thoracic plates (nota), mouthparts, and terminal abdominal segment (pygopod); 5) number and position of hair-like setae on the pronotum and throughout the body.

The eyes are the most studied organs of larval tiger beetles.The sedentary larvae have less difficulty detecting prey movement than do the mobile adults. Unlike the grub-like larvae of some insects, which only can achieve a coarse visual pattern with their simple eyes, the eyes of tiger beetle larvae have dense photoreceptors that

permit detailed focusing and 3-dimensional perception.

The pupal stage of tiger beetles has been largely ignored. Species of only 14 genera worldwide have had this stage described. A recent article by Roza and Mermudes standardizes the morphological characters of the pupal stage and summarizes what is known about the tiger beetle species, including published descriptions of the pupal stage for some Indian species.

Species Accounts

6

We use Acciavatti and Pearson's The Tiger beetle genus Cicindela (Coleóptera: Insecta)

from the Indian subcontinent, Naviaux's Les Collyris (Coleóptera, Cicindelidae): révison de genres et description de nouveaux taxons and Les Tricondylina (Coleóptera, Cicindelidae): révison de genres Tricondyla Latreille et Derocrania Chaudoir et descriptions de nouveaux taxons, and Wiesner's Die Gattung Therates Latreille und Ihre

Arten and Checklist of the Tiger Beetles of the World as a taxonomic basis for the status of names. These together with subsequent studies of DNA and morphology are used to provide the most up-to-date species list and their taxonomic groupings possible.

We also promote the use of common English names for tiger beetle species. The common name of each Indian species is paired with its scientific Latin ñame throughout this Chapter and the person(s) who originally described the species are added within the heading of each detailed species account. Subspecies, however, are indicated only by their Latin name.

Distribution maps Using a compilation of published regional recordsand collection data from specimens

in museums and personal collections, we assembled the most complete and current distributional status possible for each tiger beetle species and recognized

subspecies occurring within the political boundaries of India. If the entire known range of a species is confined to India, we refer to it as endemic.

For convenience, we have depicted these geographic ranges on maps showing state boundary lines to help orient users to localities where a given species or subspecies has been recorded. When more than one subspecies for a species has been recorded within India, each of these is included on the same map and distinguished with different symbols. Areas of subspecies intergradation are indicated with intermediate symbols. We acknowledge that the distribution maps for some species are incomplete because field observations and collecting in many areas of India have been limited. The most complete data come from the northeast, the north, and the south where historically many reports have been published on tiger beetle collections. The vastness of India has resulted in major geographical regions, most obvious from

central and northwestern India, where relatively few collections and observations have occurred. One of the major goals of the field guide is to motivate enthusiasts within India to actively seek out these less studied areas and provide information to fill in the gaps for the more common species as well as the rarer species - perhaps

even leading to the discovery of a new species hitherto unknown in the country.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Subfamily Ciclndelinae, Tribe Megacephalini Metallic Tiger Beetles, genus Grammognatha < Many authors treat Megacephala as a Pantropical genus. Others, however, restrict the name Megacephala to several species in Africa. We follow these experts and treat representatives of this group in Asia as a member of the distinct genus Grammognatha. All Metallic Tiger Beetles areyprimarily nocturnal, and adults of most species are flightless or fly only wea^®(dults tend to be highly colourful with bodies of metallic copper, blue ana green predominating. Their legs, antennae and mouth parts are usually yellowish. Only a single species occurs in

Europe, North Africa and southwest Asia. Some species of this group hear sounds of prey calling from underground and have been proposed as natural control of pests such as mole-crickets.

Eurasian MetallicTiger Beetle, Grammognathaeuphratica (Latreilleand Dejean) (Plate 1) [Map 1] Identification: Length 18 - 22.5 mm. Body robust with the upper surface metallic green with maroon reflections. A large ovoid yellow area covers the back one-third

of each elytron. The antennae, legs, and palpi are yellowish. The heavy mandibles are yellow with blackish tips. Unlikely to be confused with any other species in India.

Subspecies and morphological variants: The inland southwest Asia population that exists in northern India as blue individuals is often assigned to the subspecies

G. e. armenica.

Distribution and habitats: Spain and Morocco east across northern Africa and southern Europe to southwestern Asia. In India this species is known only from Agra (Uttar Pradesh). Occurs along coastal salt flats and inland salt marshes and oases.

Behaviour: Adults are active only at night, and although they have wings, they rarely use them. They can most frequently be seen running along wet sand beaches and

muddy flats at night. They are attracted to night lights. The larvae are known only from captive breeding, but presumably both larvae and adults occur in similar habitats.

Seasonality: Adult activity in southwest Asia is limited to the warm, wet monsoon

season from July to September. 30

Map 1 Eurasian Metallic Tiger Beetle, Grammognatha euphratica.

Species Accounts

Subfamily Collyrinae, Tribe Collyrini Arboreal Tiger Beetles, genus Tricondyla These long, narrow beetles are all blackish, flightless with fused elytra, and primarily arboreal and diurnal. They are limited to moist forests and resemble large ants. Some species are remarkably mimicked by late instar grasshoppers. They run up and down the trunks of trees and move to the opposite side of the tree when threatened by a predator. When pressed by danger, they will fall to the ground and remain still in the leaf litter. They also move between trees by walking across the ground. Larval burrows have been found in the bark of large trees, and the larvae are active primarily at night. The 48 species of this genus occur throughout southeastern Asia to northern Australia. Four species are known Atom India, one of which is endemic there.

Key I: Arboreal Tiger Beetles, genus Tricondyla Identification Key to Tricondyla Species la.

Pronotum with the sides parallel as far

2

as the apical constriction (Fig. 6.1)

1b.

Pronotum with the sides more or less

widened

and

rounded,

and

3

more

or less convergent before the apical

Figure 6.1

Sides of thorax (pronotum) relatively parallel and straight-sided.

2a. (la.)

Figure 6.2 Sides of thorax (pronotum) conspicuously widened and rounded.

Pronotum strongly striated transversely

Pleated Arboreal Tiger

Beetle, Tricondyla femorata 2b.

Pronotum finely striated transversely

Satin Arboreal Tiger Beetle,

3a. (1 b.)

Elytral sculpture strong and granular

Lumpy Arboreal Tiger

Tricondyla gounellei

Beetle, Tricondyla tuberculata

3b.

Elytral sculpture moderately shallow,

Thick-necked Arboreal Tiger

formed by transverse wrinkles in the

Beetle, Tricondyla macrodera

anterior half, apex nearly smooth 31

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Pleated Arboreal Tiger Beetle, Tricondyla femorata Walker (Plate 1) [Map 2] Identification: Length 20 - 24 mm; body black and shiny with some metallic reflections; elytra with fine wrinkles running across their width; head broad with relatively shallow excavation between eyes; legs long and dark red; labrum and mouth parts dark red.

Subspecies and morphological variants: Formerly considered a subspecies of Tricondyla granulifera Motschoulsky, which is confined to Sri Lanka.

Distribution and habitats: Southern India (Nilgiri Hills) and Sri Lanka. Forest and forest edge at mid elevations above 300 m.

Behaviour: Runs quickly up and down tree

Map 2 Pleated Arboreal Tiger Beetle,

trunks and occasionally across the leaf litter

Tricondyla femorata.

to the next tree.

Seasonality: Unknown.

Satin Arboreal Tiger Beetle, Tricondyla gounellei Horn (Plate 1) [Map 3] Identification: Length 18-24 mm; body black and velvety; elytra with small, shallow cavities that resemble roof tiles and run

across the width of the elytra to form irregular wavy lines; elytra short and narrow

in the middle; head smooth; legs long and dark red, labrum and first four segments of antennae black, the rest dark red.

Subspecies and morphological variants: No other subspecies have been described from India.

Distribution and habitats: Endemic to Southern India. Forest and forest edge (750 -1200 m).

Behaviour: Runs quickly up and down tree

Map

3 Satin Arboreal Tricondyla gounellei.

trunks and occasionally across the leaf litter t the next tree.

Seasonality: March to November. 32

Tiger

Beetle,

Species Accounts

Thick-necked Arboreal Tiger Beetle, Tricondyla macrodera Chaudoir (Plate 1) [Map 4] Identification: Length 16-20 mm; body dark brownish and moderately shiny; neck

broad and almost as wide as the eyes; elytra with striations across their width

at the front and smooth toward the back; legs with reddish femora and patchy black tibiae and tarsi; labrum and mouth parts

brownish red.

Subspecies and morphological variants: Two other subspecies have been described

from Burma and South China.

Distribution

and

habitats:

Himalayas Bhutan, Nepal, Myanmar, Cambodia, Vietnam, and China.

Map 4 Thick-necked Arboreal Tiger Beetle,

Forest and forest edge on the lower slopes

Tricondyla macrodera.

in

Northeastern

India,

(180-1530 m).

Behaviour: Runs rapidly up and down the trunks of trees searching for prey. Occasionally drops to ground and hides under the leaf litter. Seasonality: April to August.

Lumpy Arboreal Tiger Beetle, Tricondyla tuberculata Chaudoir (Plate 1) [Map 5] Identification: Length 17-19 mm; body dull black; elytra with coarse sculpturing over their entire surface; legs with reddish femora and black tibiae and tarsi; labrum and mouth parts black.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India, Bangladesh, and Myanmar. Forest and forest edge in the lowlands along the lower

slopes of the Himalayas (700 - 1400 m) in

Behaviour: Unknown. Seasonality: April to September.

Map 5 Lumpy Arboreal Tiger Beetle, Tricondyla tuberculata.

33

k Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Bark Tiger Beetles, genus Derocrania These all black species are small counterparts of the closely related genus Tricondyla and closely resemble ants. They are flightless with their elytra fused, and they run rapidly up and down the bases of tree trunks. The larvae are unknown and little has been written about general behaviour and natural history of the adults. Four of the 16 species in this genus are endemic to extreme southern parts of India with the remainder endemic to Sri Lanka.

Key II: Bark Tiger Beetles, genus Derocrania Identification Key to Derocrania Species 1 a.

Pronotum with a distinct neck (collum)

2

(Fig. 6.3)

1 b.

Pronotum conical, without distinct neck

3

(collum) (Fig. 6.4)

Figure 6.3 Thorax (pronotum) with distinct

Figure 6.4 Thorax (pronotum) conical with

neck (collum).

no distinct neck (collum).

2a. (la.)

Sculpture of elytra coarse at base, and

Camel-backed Bark Tiger

with deep longitudinal grooves toward

Beetle, Derocrania longesulcata

the back

2b.

Sculpture of elytra coarse at base, then

Flat-headed Bark Tiger

formed by longitudinal and moderately

Beetle, Derocrania honorei

deep wrinkles, irregularly shaped and more or less confusedly marked

3a. (1b.)

Pronotum short; greatest width of elytra

approximately at 2/3 of the length 3b.

Pronotum long; greatest width of elytra placed in the middle

34

Long-necked Bark Tiger

Beetle, Derocrania dembickyi Short-necked Bark Tiger

Beetle, Derocrania brevicollis

Species Accounts

Camel-backed Bark Tiger Beetle, Derocrania longesulcata (Horn) (Plate 1) [Map 6] Identification: Length 9 - 11.5 mm; body shiny black with blue or bronze reflections; elytra widen abruptly and become much

wider than the pronotum; elytra with regular

deep longitudinal grooves.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to the southern part of the Western Ghats of India. Forests (850 - 1800 m).

Behaviour: Runs like an ant and emits a defensive chemical with a peculiar odour.

Seasonality: April to October.

Map 6 Camel-backed Bark Tiger Beetle, Derocrania longesulcata.

Flat-headed Bark Tiger Beetle, Derocrania honorei Fleutiaux (Plate 2) [Map 7] Identification: Length 9-11.5 mm; body shiny black with bronze reflections; top of

head relatively flat with little excavation between eyes; elytra widen gradually and become somewhat wider than the

pronotum; elytra with coarse sculpturing at

base than

becoming

irregular with

longitudinal wrinkles toward the middle and

end. Similar to the Long-necked Bark Tiger Beetle (D. dembickyi), but the Flat-headed

Bark Tiger Beetle is smaller, elytral grooves irregular, and the female has a shorter and

broader pronotum.

Subspecies and morphological variants:

Map 7 Flat-headed Bark Tiger Beetle,

No subspecies described.

Derocrania honorei.

Distribution and habitats: Endemic to the southern part of the Western Ghats of India. Upland forests (1066 m).

Behaviour: Unknown.

Seasonality: Unknown.

35

k Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Long-necked Bark Tiger Beetle, Derocrania dembickyi Naviaux and Moravec (Plate 2) [Map 8] Identification: Length 10 - 11.5 mm; body shiny black with bronze to green reflections; elytra widen gradually and become somewhat

wider than the pronotum; elytra with welldefined longitudinal grooves. Similar to the Flat-headed Bark Tiger Beetle (D. honorei), but the Long-necked Bark Tiger Beetle is larger with elytral grooves more obvious.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to the Western Ghats of southern India. Forests (900-1000 m). Map 8 Long-necked Bark Tiger Beetle,

Behaviour: Unknown.

Derocrania dembickyi.

Seasonality: Unknown.

Short-necked Bark Tiger Beetle, Derocrania brevicollis Horn (Plate 2) [Map 9] Identification:

Length

10

-

12,

mm;

body shiny black with blue reflections on

pronotum and head and bronze reflections on elytra; pronotum relatively short; elytra widen noticeably at about half of their

length; elytra with irregular sculpturing at

base then forming ambiguous longitudinal grooves toward the middle and end. Resembles Long-necked Bark Tiger Beetle (D. dembickyi), but the Short-necked Bark Tiger Beetle has a shorter pronotum and the elytra widest in the middle.

Subspecies and morphological variants: No subspecies have been described.

Map 9 Short-necked Bark Tiger Beetle,

Distribution and habitats: Endemic to the

Derocrania brevicollis.

Western Ghats of southern India. Forests

(1900 m).

Behaviour: Unknown. Seasonality: July to December. 36

Species Accounts

Brush Tiger Beetles, Subtribe Collyrina Adults of these narrow-bodied species run along leaf surfaces, flowers and small branches of undergrowth bushes and trees to hunt for insect prey and seek mates, but they readily fly from bush to bush to escape danger. Most are dark bluish or green to black. The larvae make their tunnels in dead and decaying stems of

bushes and trees. The 299 species in this subtribe occur from India and Sri Lanka throughout Southeast Asia and central China to Taiwan and the Philippines. Sixty-six species are

found in India, 39 of which are endemic. They have been divided into three provisional genera (Protocollyris, Neocollyris, and Collyris) based on subtle and often confusing morphological characters. The largest genus, Neocollyris, consists of 269 species and has in turn been divided into 12 subgenera based on even more subtle characters. Few extensive molecular studies have been done to more confidently establish relationships among the genera and subgenera. As future studies of DNA and RNA

are conducted, they will doubtlessly force rearrangements of our present concepts.

Primitive Brush Tiger Beetles, genus Protocollyris The four species found in India are among the smallest species in this subtribe (body length 7-12 mm). Three are endemic to south India and one extends into northeast India and Southeast Asia.

Key III: Primitive Brush Tiger Beetles, genus Protocollyris Identification Key to Protocollyris Species la.

Legs completely black

Short-lipped Brush Tiger

Beetle, Protocollyris brevilabris

1b. 2a. (1b.)

Femora partly brownish

Pronotum with a distinct neck (collum)

2 Brittle Brush Tiger Beetle,

Protocollyris fragilis 2b.

Pronotum conical, without distinct neck

3

(collum) 3a. (2b.)

Pronotum finely striated across its width;

Nilgiri Brush Tiger Beetle,

antennae long, reaching half of the

Protocollyris niligiriensis

pronotum, when bent backwards 3b.

Pronotum smooth, not striated; antennae

Stout Brush Tiger Beetle,

short, reaching the first 3rd of the

Protocollyris pacholatkoi

pronotum, when bent backwards

37

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Nilgiri Brush Tiger Beetle, Protocollyris niligiriensis Naviaux (Plate 2) [Map 10] Identification: Length 7.5 - 9.2 mm, body dark, dull blue; pronotum long and narrow; elytra slightly expanded toward the rear and

with dense microsculpturing evenly over entire surface; underside black; legs black with reddish femora. Similar to Patcholatko's Brush Tiger Beetle (Neocollyris patcholatkoi), which has a shorter and thicker pronotum, parallel-sided elytra, and shorter antennae.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from the Nilgiri Hills of Tamil Nadu (900 -

1100 m). Leaves of tall undergrowth bushes and short trees.

10 Nilgiri Brush Protocollyris nilgiriensis.

Map

Tiger

Beetle,

Tiger

Beetle,

Behaviour: Unknown.

Seasonality: May.

Stout Brush Tiger Beetle, Protocollyris pacholatkoi Naviaux (Plate 2) [Map 11] Identification: Length 6.6 - 9 mm; body dark, dull blue; pronotum relatively short and thick; elytra nearly parallel-sided and with shallow microsculpturing over entire surface; underside black; legs black with reddish femora; antennae extremely short. Similar to Nilgiri Brush Tiger Beetle (Neocollyris nilgiriensis), which has a longer and thinner pronotum, elytra whose sides expand noticeably toward the rear, and

longer antennae.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from the Nilgiri Hills of Tamil Nadu (900 - 1100 m).

Behaviour: Unknown.

Seasonality: May. 38

Map

11 Stout Brush Protocollyris pacholatkoi.

Species Accounts

Short-lipped Brush Tiger Beetle, Protocollyris brevilabris Horn (Plate 2) [Map 12] Identification: Length 7 - 8.5 mm; body dark blue to green; femurs of two back pair of legs reddish; short, yellowish labrum; head relatively flat between eyes.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Eastern India, Myanmar, Bhutan, Thailand, Malaysia, and

Indonesia.

Behaviour: Unknown.

Seasonality: Unknown. Map 12 Short-lipped Brush Tiger Beetle, Protocollyris brevilabris.

Brittle Brush Tiger Beetle, Protocollyris fragilis Naviaux (Plate 2) [Map 13] Identification: Length 9.2 mm; slender head and pronotum contrast with relatively broad elytra; somewhat resembles the Longlegged Brush Tiger Beetle (A/, maindroni), but the Brittle Brush Tiger Beetle has a shallower cavity between the eyes, and a much smaller labrum.

Subspecies and morphological variants:

Distribution and habitats: Known only from Kerala in South India.

Behaviour: Unknown.

Seasonality: May. Map

13 Brittle Brush. Tiger Protocollyris fragilis.

Beetle,

39

h Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Brush Tiger Beetles, genus Neocollyris The 56 species of this genus that occur in India make up almost a quarter of its entire tiger beetle fauna. They occur across the country except for the deserts of the northwest and high altitude habitats of the north. Thirty-three species are endemic to India.

Key IV: Brush Tiger Beetles, Neocollyris Identification Key to Neocollyris Species la.

Pronotum long, with slender curved neck (collum) (Fig. 6.5)

Bottle-necked Brush Tiger Beetle, Neocollyris ampullicollis

1 b.

Pronotum shorter, collum, if present, not slender (Fig. 6.6)

2

Figure 6.5 Thorax with long, slender neck

Figure 6.6 Thorax short with little or no

(collum) for more than half its length.

obvious neck (collum).

2a. (1b.)

Antennae very short, reaching back only to the middle of the pronotum (Fig. 6.7); labrum with single row of setae (submarginal) at the lower edge

10

2b.

Antennae longer (Fig. 6.8); labrum with additional setae above the submarginal setae row

3

Figure 6.7 Short antennae reaching back only half the length of the thorax. 40

Figure 6.8

Long antennae reaching back much the length of the thorax; head with an oval impression between eyes.

Species Accounts

3a. (2b.)

Excavation on top of head between the eyes (interocular) with an oval impression (Fig. 6.8); aedeagus with apical knob

14

3b.

Excavation without oval impression; aedeagus without apical knob

4

4a. (3b.)

Body length less than than 9 mm, stout and broad

Maroon-spotted Brush Tiger Beetle, Neocollyris purpureomaculata

4b. 5a. (4b.) 5b.

Body length greater than 9 mm

5

Labrum short (Fig. 6.9)

18

Labrum longer (Fig. 6.10)

Figure 6.9 Short labrum on front of head.

Figure 6.10 Long labrum on front of head.

6a. (5b.)

Pronotum slender, tapered, spindleshaped, or spherical at the rear, with relatively smooth surface

19

6b.

Pronotum broader, with shallow but distinct cross grooves (rugae) (Fig. 6.11)

7

Figure 6.11 Broad thorax (pronotum) with shallow grooves crossing the surface and sides nodular in shape.

7a. (6b.)

7b.

Pronotum nodular (Fig. 6.11) Shape of pronotum not nodular

26

8

8a. (7b.)

Labrum long with elongated teeth (Fig 6.12)

30

8b.

Labrum shorter, with short teeth (Fig 6.13)

9

41

h Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Figure 6.12 Long labrum with elongated teeth. 9a. (8b.)

Pronotum long, slender, with shallow grooves across its width (Fig. 6.14)

34

9b.

Pronotum short, surface relatively smooth and without shallow grooves across its width (Fig. 6.15)

41

Figure 6.14 Thorax (pronotum) with shallow but distinct grooves across it.

Figure 6.15 Thorax (pronotum) with relatively smooth surface.

10a. (2a.)

Head (without labrum) as long as wide (Fig. 6.16)

11

10b.

Head (without labrum) wider than long (Fig. 6.17)

12

Figure 6.16 Head in top dimensions as long as wide.

42

Figure 6.13 Shorter labrum with short teeth.

Species Accounts

11 a. (10a.)

11b.

12a. (10b.)

12b. 13a. (12b.)

13b.

Labrum with fewer than pronotum smooth

10 setae;

Labrum with more than 10 setae; pronotum with shallow furrows

Hairy Brush Tiger beetle, Neocollyris multipilosa

Tapered Brush Tiger Beetle, Neocollyris attenuata

Hind tarsi yellow

Hind tarsi black

Pronotum short and (collum) short

Brown Brush Tiger Beetle, Neocollyris acuteapicalis

13 smooth,

neck

Pronotum long with shallow grooves (striae) across it, neck (collum) longer

Large-necked Brush Tiger Beetle, Neocollyris crassicornis Greyish Brush Tiger Beetle, Neocollyris subclavata

14a. (3a.)

Legs completely yellow

15

14b.

Femora darkened above

16

Antennal segments 4 to 11 yellow

Yellow-legged Brush Tiger Beetle, Neocollyris roeschkei

15b.

Antennal segments 4 to 11 darkened at their far (distal) ends

Ringed Brush Tiger Beetle, Neocollyris annulicornis

16a. (15a.)

Pronotum long, with a distinct, long neck (collum)

Ingrid's Brush Tiger Beetle, Neocollyris ingridae

16b.

Pronotum shorter, without distinct long neck (collum)

17

15a. (14a.)

17a. (16b.)

17b.

18a. (5a.)

18b.'

19a. (6a.)

19b.

20a. (19a.)

20b.

Tibia of hind legs completely yellow

Basal half of hind tibia darkened

Meghalaya Brush Tiger Beetle, Neocollyris pearsoni

Thin Brush Tiger Beetle, Neocollyris macilenta

Centre of labrum yellow

Four-lined Brush Tiger Beetle, Neocollyris quadrisukata

Labrum completely dark metallic

Blue-feeler Brush Tiger Beetle, Neocollyris cyaneipalpis

Labrum partly yellow

20

Labrum completely dark, metallic

22

Middle portion of tibiae dark, yellowish at the base and/or far end (distally) Tibiae completely yellow

Stout Brush Tiger Beetle, Neocollyris pacholatkoi 21

43

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicmdelidae

21a. (20b.)

Outer-most segment of labial palpi drop-shaped (Fig. 6.18); aedeagus S-shaped (sigmoid)

Diamond-headed Brush Tiger Beetle, Neocollyris variicornis

21b.

Outer-most segment of labial palpi triangular (Fig. 6.19); aedeagus curved

Slim Brush Tiger Beetle, Neocollyris brancuccii

Figure 6.18 Last segment of jointed tasters (labial

Figure 6.19 Last segment of jointed tasters

palpi) tear- drop shaped.

(labial palpi) triangular shaped.

22a. (19b.)

Tarsi of hind leg yellow

23

22b.

Tarsi of hind leg black

24

Tibia of hind leg completely yellow

Spindle-necked Brush Tiger Beetle, Neocollyris variitarsis

23b.

Basal half of hind tibia dark

Long-necked Brush Tiger Beetle, Neocollyris brendelli

24a. (22b.)

Femora completely yellow

Short-necked Brush Tiger Beetle, Neocollyris kollari

23a. (22a.)

24b.

Femora darkened above

25

25a. (24b.)

Body colour dark green

Long-legged Brush Tiger Beetle, Neocollyris maindroni

25b.

26a. (8b.)

Pronotum brownish black

Lesser Brush Tiger Beetle, Neocollyris parvula 27

Middle part of pronotum yellowish red

Pitted Brush Tiger Beetle, Neocollyris foveifrons

27a. (26a.)

Elytra with large light brown patches at rear and at the shoulders

Thin-spotted Brush Tiger beetle, Neocollyris smithii

27b.

Elytra without light brown patches, but with a yellow band in the middle

28

28a. (27b.)

Pronotum with strong grooves across its width

Assam Brush Tiger Beetle, Neocollyris assamensis

28b.

Pronotum with smooth surface or barely visible grooves across its width

29

26b.

44

Body colour blue

Species Accounts

29a. (28b.)

Elytra near the central suture with connected rows of punctures (Fig. 6.20); pronotum without neck (collum)

Rufous Brush Tiger Beetle, Neocollyris coapteroides

29b.

Elytra near the central suture with isolated punctures (Fig. 6.21); pronotum with short neck (collum)

Long-backed Brush Tiger Beetle, Neocollyris apteroides

Figure 6.20 Punctures connected across central

Figure 6.21 Punctures isolated on central

part of elytra.

part of elytra.

30a. (8a.)

Tarsi of hind legs black

31

30b.

Tarsi of hind legs yellow

33

31a. (30a.)

Elytra with yellow middle band and yellow rear (apical) line

Brown-kneed Brush Tiger Beetle, Neocollyris nilgirica

31b.

Elytra only with brownish patches at rear

32

32a. (31b.)

Elytra with deep transverse furrows in the middle, nearly smooth at the shoulders and rear (Fig. 6.22)

Chestnut Brush Tiger Beetle, Neocollyris rubens

32b.

Shoulder and rear covered with deep pits (Fig. 6.23)

Narrow-necked Brush Tiger Beetle, Neocollyris compressicollis

Figure 6.22 Elytra with deep furrows in central

Figure 6.23 Elytra with deep pits at shoulders

parts.

and rear ends.

45

A. Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

33a. (30b.)

Body colour black with blue reflections; pronotum shorter; with short neck (collum) less than one third length of pronotum

Small-eyed Brush Tiger Beetle, Neocollyris andrewesi

33b.

Body colour black with green reflections; pronotum longer; with neck (collum) one third of length of pronotum

Charcoal Brush Tiger Beetle, Neocollyris anthracina

Body length more than 15 mm

Blackish Brush Tiger Beetle, Neocollyris juengeri

34a. (9a.) 34b.

35a. (34b.)

35b.

36a. (35b.) 36b.

37a. (36b.)

37b.

Tibia and femora of all legs yellow

Tibia of all legs black, lighter at base in some specimens Elytra blackish blue Elytra greenish

Lower part of labrum yellow

Labrum completely black

35

Smooth-necked Brush Tiger Beetle, Neocollyris subtileflavescens 36 Yellow-spotted Brush Tiger Beetle, Neocollyris conspicua

37 Least Brush Tiger Beetle, Neocollyris flava 38

38a. (37b.)

Pronotum with a distinct and long neck (collum); elytra with a broad yellow middle patch

Greenish Brush Tiger Beetle, Neocollyris vannideki

38b.

Pronotum with no neck (collum) or, if present, collum short and indistinctelytra with a slender yellow middle band

39

39a. (38b.) 39b.

Elytra evenly dark green Elytra with contrasting metallic luster near the yellow middle band

Wrinkle-necked Brush Tiger Beetle, Neocollyris rugata 40

40a. (39b.)

Elytra light green

Furrow-necked Brush Tiger Beetle, Neocollyris plicicollis

40b.

Elytra blue green

Metallic Brush Tiger Beetle, Neocollyris metallica

41a. (9b.)

41b.

42a. (41a.) 42b.

43a. (42a.) 43b.

46

Body length less than 15 mm

Tarsi of hind legs yellow

42

Tarsi of hind legs black

49

Tibia of hind legs completely yellow

43

Tibia of hind legs black or occasionally with outer ends partially yellow

45

Body length more than 13 mm Body length less than 13 mm

Dark-legged Brush Tiger Beetle, Neocollyris fuscitarsis

44

Species Accounts

44a. (43b.)

Last abdominal segment yellow

Husky Brush Tiger Beetle, Neocollyris schaumi

44b.

Last abdominal segment black

Graceful Brush Tiger Beetle, Neocollyris redtenbacheri

45a. (42b.)

Body length more than 16 mm

46

Body length less than 16 mm

47

45b.

46a. (45a.)

Surface of elytra dull and flat with dense pits often running together (Fig. 6.24)

Broad-headed Brush Tiger Beetle, Neocollyris insignis

46b.

Surface of elytra shiny, pits separated (Fig. 6.25)

Eminent Brush Tiger Beetle, Neocollyris egregia

Figure 6.24 Pits on elytra running together.

Figure 6.25 Pits on elytra separated.

47a. (45b.)

Outer quarter or more of tibia on hind legs yellow

48

47b.

Tibia of hind legs all dark to nearly black

Nepal Brush Tiger Beetle, Neocollyris nepalensis

48a. (47a.)

48b.

Elytra blue with dense regular pits (Fig. 6.26)

Blood-spotted Brush Tiger Beetle, Neocollyris cruentata

Elytra black with metallic luster, pits small in basal half, larger and longitudinal in apical half (Fig. 6.27)

Figure 6.26 Pits on elytra dense and regularly-spaced.

Golden Brush Tiger Beetle, Neocollyris orichalcina

Figure 6.27 Pits on elytra small towards the front, but larger towards the rear.

47

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

49a. (41b.)

49b.

50a. (49a.)

Palpi partly yellow, dark brown, or completely yellow

50

Palpi black

52

Body length more than 15 mm

Body length less than 15 mm

50b.

51

51a. (50b.)

Tibia of all legs dark with metallic reflections

Red-tipped Brush Tiger Beetle, Neocollyris rufipalpis

51b.

Basal part of tibia on front and middle legs yellow

Blue-lipped Brush Tiger Beetle, Neocollyris similis

52a. (49b.)

Outer six segments of antennae yellow or reddish

Round-headed Brush Tiger Beetle, Neocollyris batesi

52b.

Outer six segments of antennae dark brown or black

53

53a. (52b.)

Body length more than than 13.5 mm

Body length less than 13.5 mm

53b.

54a. (53b.)

Elytra with a yellow spot at the shoulder

Elytra without shoulder spot

54b.

Sapphire Brush Tiger Beetle, Neocollyris saphyrina

54

Orange-spotted Brush Tiger Beetle, Neocollyris hiekei

55

55a. (54b.)

Pronotum slender (Fig. 6.28) with many punctures; aedeagus length more than 1.7 mm

Long-lipped Brush Tiger Beetle, Neocollyris distincta

55b.

Pronotum stout (Fig. 6.29) with few punctures; aedeagus length less than 1.7 mm

Red-legged Brush Tiger Beetle, Neocollyris bonellii

Figure 6.28 Pronotum slender. 48

Blue Brush Tiger Beetle, Neocollyris smaragdina

Figure 6.29 Pronotum stout.

Species Accounts

Maroon-spotted Brush Tiger Beetle, Neocollyris purpureomaculata (Horn) (Plate 2) [Map 14] Identification: Length 8.7 mm; head and pronotum black with green reflections; elytra bright green with two large maroon to purple bands across each elytron.

Subspecies and morphological variants: The subspecies N. p. borea is the only one known from India.

Distribution and habitats:The subspecies N. p. borea is found (1400 - 2000 m) in northeastern India, Vietnam, Laos Malaysia, Thailand, Myanmar, and south China.

Behaviour: Unknown. Seasonality: April to May.

Map 14 Maroon-spotted Brush Tiger Beetle, Neocollyris purpureomaculata.

Ingrid's Brush Tiger Beetle, Neocollyris ingridae Naviaux (Plate 3) [Map 15] Identification: Length 9.5 - 11.3 mm; small shiny black head; long and slender pronotum shiny dark green; elytra green with small oblong reddish patch on mid­ margin of each elytron.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from northeastern India (500 - 1150 m).

Behaviour: Unknown.

Seasonality: April to May.

Map 15

Ingrids Neocollyris ingridae.

Brush Tiger

Beetle,

49

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Thin Brush Tiger Beetle, Neocollyris macilenta Naviaux (Plate 3) [Map 16] Identification: Length 9 mm; head and pronotum shiny green; elytra parallel­ sided and green with small oblong reddish patch on margin of each elytron; similar to Ingrid's Brush Tiger Beetle (N. ingridae), but the Thin Brush Tiger Beetle has a shorter pronotum and larger head; the Ringed Brush Tiger Beetle (A/, annulicornis) is also similar, but it lacks the reddish spots on

the elytra and the sides of its elytra expand toward the back.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India at 1400 m.

Map 16 Thin Brush Tiger Beetle, Neocollyris macilenta.

Behaviour: Unknown.

Seasonality: April.

Ringed Brush Tiger Beetle, Neocollyris annulicornis Naviaux (Plate 3) [Map 17] Identification: Length 10.2 mm; head and neck short, shiny blue and green; elytra

green and conspicuously widen toward the back. No reddish spots on elytra.

Subspecies and morphological variants: No subspecies have been described

Distribution and habitats: Endemic to northeastern India at 1150 m.

Behaviour: Unknown.

Seasonality: April.

Map

17 Ringed Brush Tiger Neocollyris annulicornis.

50

Beetle,

Species Accounts

Yellow-legged Brush Tiger Beetle, Neocollyris roeschkei (Horn) (Plate 3) [Map 18] Identification:

Length

12

mm;

head

dark violet; pronotum very short and

bright blue; elytra blue-green with violet borders; elytra with triangular shaped

reddish spot on mid-margin of each elytron; bright yellow narrow band across front edge of elytra; rear surface of elytra

heavily punctured; antennae and legs yellow.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from the southern part of West Bengal in northeastern India.

Behaviour: Unknown.

Map 18 Yellow-legged Brush Tiger Beetle, Neocollyris roeschkei.

Seasonality: Unknown.

Meghalaya Brush Tiger Beetle, Neocollyris pearsoni Naviaux (Plate 3) [Map 19] Identification: Length 9.8 mm; body blue; pronotum short and broad; elytra with small yellowish oblong patches on their mid­

margins.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from northeastern India.

Behaviour: Unknown.

Seasonality: April.

Map 19 Meghalaya Brush Tiger Beetle, Neocollyris pearsoni.

51

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Blue-feeler Brush Tiger Beetle, Neocollyris cyaneipalpis Horn (Plate 3) [Map 20] Identification: Length 8.9 mm; labrum short and black with blue reflections; elytra with small diffuse light areas along

the mid-margins; short lengthwise ridge between eyes; long antennae. Similar to the Four-lined Brush Tiger Beetle (N. quadrisuculata) which is distinguished from the Blue-feeler Brush Tiger Beetle by a long lengthwise ridge between the eyes, shorter antennae, and dark labrum with yellow in the centre.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to

Map 20 Blue-feeler Brush Tiger Beetle,

Western Ghats in southern India.

Neocollyris cyaneipalpis.

Behaviour: Unknown.

Seasonality: Unknown.

Four-lined Brush Tiger Beetle, Neocollyris quadrisulcata Horn (Plate 3) [Map 21] Identification: Length 8.8 mm; body blue; long

ridge

lengthwise

between

eyes;

relatively short antennae; labrum black with yellow centre. Similar to Blue-feeler Brush Tiger beetle (N. cyaneipalpis), but which has a short lengthwise ridge between eyes, long antennae, and an all-black labrum.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from Tamil Nadu in southern India.

Behaviour: Unknown.

Seasonality: Unknown.

Map 21 Four-lined Brush Tiger Beetle, Neocollyris quadrisulcata.

52

Species Accounts

Graceful Brush Tiger Beetle, Neocollyris redtenbacheri Horn (Plate 3) [Map 22] Identification: Length 9.5 - 11.5 mm; body bright green to blue green; legs, palpi,

antennae yellow; labrum metallic blue; small

yellow spots on mid-margin of elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Himalayas from northeastern Pakistan to northern India, Nepal, Bhutan, and Myanmar. Most common Brush Tiger Beetle species in secondary

forest, gardens and nurseries (60 - 2500 m).

Behaviour: Occurs on the leaves and stems of bushes 1 - 2 m above the ground. It flie$ spontaneously for short distances and runs rapidly on leaf surfaces in search of insect

Map 22 Graceful Brush Tiger Beetle, Neocollyris redtenbacheri.

prey.

Seasonality: April to July.

Husky Brush Tiger Beetle, Neocollyris schaumi Horn (Plate 3) [Map 23] Identification: Length 9.5 - 11 mm; head blue; pronotum green; elytra bright green with copper reflection and a diffuse reddish soot on the mid-margin; labrum metallic green; similar to the Graceful Brush Tiger Beetle (N. redtenbacheri), which is shorter and has the last abdominal segment yellow.

Subspecies and morphological variants: A variant of this species was described as the variety N. r. chevrolati. It is broader and

shorter with a fine rufous line on the elytral margins at the shoulder and middle.

Distribution and habitats: Endemic to the Andaman Islands.

Behaviour: Unknown.

Map 23 Husky Brush Neocollyris schaumi.

Tiger

Beetle,

Seasonality: Unknown. 53

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Red-legged Brush Tiger Beetle, Neocollyris bonellii Guérin-Méneville (Plate 4) [Map 24] Identification: Length 9-13 mm; head and pronotum dark blue to black; elytra bright blue with no or weak reddish oval mark at mid-margin; trochanters of middle leg yellowish; femora reddish.

Subspecies and morphological variants: Five subspecies have been described, but only N. b. bonnellii is found in India. Another subspecies was described for India (Nepal

and Sikkim), N. b. arunensis, but is now considered to be only a variant within the

population and not a subspecies. It has a wider and longer head, shorter pronotum, and wider elytra than the nominative N. b. bonellii.

Map 24 Red-legged Brush Tiger Beetle,

Distribution and habitats: One of the most

Neocollyris bonellii.

widespread species of Neocollyris, it occurs throughout most of India (except the desert northwest), Nepal, China, Burma, Thailand, Malaysia, Vietnam, Indonesia and the Philippines. Lowland forests (75 - 1000 m)

Behaviour: Found on broad-leafed undergrowth plants in shady places and on foliage along paths and roads in or near subtropical forests.

Seasonality: April to December.

Long-lipped Brush Tiger Beetle, Neocollyris distincta Chaudoir (Plate 4) [Map 25] Identification: Length 10.5 -13.5 mm; body bright blue to green or violaceous; legs red; long labrum black with blue reflections; elytra I

surface with smooth microsculpturing, which together with its overall smaller size separate it from the similar Sapphire Brush Tiger Beetle (N. saphyrina). From the similar Red-legged Brush

Tiger Beetle (N. bonellii) it can be distinguished by its larger size and long labrum.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from southern and northeastern Indian (900 -1000 m). Behaviour: Unknown. Seasonality: May to October. 54

Map 25 Long-lipped Brush Tiger Beetle, Neocollyris distincta.

Species Accounts

Nepal Brush Tiger Beetle, Neocollyris nepalensis Naviaux (Plate 4) [Map 26] Identification: Length 12.4 - 14.2 mm; head black with violet reflections; body long, robust and blue; leg trochanters dark

and femora reddish with black patches. Separated from the similar Red-legged

Brush Tiger beetle (A/, bonellii), which has yellow rather than dark trochanters.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats:

Northeastern

India, eastern Nepal, and Myanmar. Forest

patches in the river plains (500 - 1150 m).

Behaviour: Unknown. Seasonality: April to May.

Map 26 Nepal Brush Neocollyris nepalensis.

Tiger

Beetle,

Orange-spotted Brush Tiger Beetle, Neocollyris hiekei Naviaux (Plate 4) [Map 27] Identification: Length

10.8 mm; Head black with violet reflections; pronotum

bright violet; elytra bright blue-violet with large bright orange spots on mid-margin and at .shoulders; trochanters and femora red. Resembles the Red-legged Brush Tiger Beetle (A/, bonellii), which has bluish elytra with no or faint orange spots.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from the foothills of the Himalayas in Sikkim. Behaviour: Unknown.

Seasonality: Unknown.

Map 27

Orange-spotted Beetle, Neocollyris hiekei.

Brush

Tiger

55

A Field Guide to the Tlyrr Rrrtlr* of Incita; Identification and Biology of the Clclndelldae

Blood-spotted Brush Tiger Beetle, Neocollyris cruentata (Schmidt-Goebel) (Plate 4) [Map 28] Identification: Length 11.6 - 13.4 mm; large head with prominent excavation between eyes; short, thick pronotum; body blue,

green violaceous or brownish; elytra often with dark red stripe on the shoulders and irregular reddish patch at mid-margins; tarsal segments of hind leg light-coloured. Larger than the similar Red-legged Brush Tiger Beetle (N. bonellii) and the excavation

between the eyes more prominent. Light colour of the hind tarsi separate it from the

Red-tipped Brush Tiger Beetle (/V. rufipalpus).

Subspecies and morphological variants: No subspecies have been described.

Map 28 Blood-spotted Brush Tiger Beetle,

Distribution and habitats: Northeastern

Neocollyris cruentata.

India to Myanmar, Thailand, Laos, Vietnam, China, and Indonesia. Broad-leafed undergrowth along paths and roadsides of tropical mountain forests.

Behaviour: Unknown. Seasonality: April to July.

Round-headed Brush Tiger Beetle, Neocollyris batesi Horn (Plate 4) [Map 29] Identification: Length 13-14.5 mm; head black, round and large with eyes relatively small; distinctive excavation between eyes;

pronotum blue-violet; elytra roughly pitted,

blue with no reddish marks. Resembles the

Red-legged Brush Tiger Beetle (N. bonelfi) but is readily distinguished by its large round head. At one time it was considered a subspecies of the Red-legged Brush Tiger Beetle.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern India, Myanmar,

Laos,

Thailand,

Indonesia.

Behaviour: Unknown.

Seasonality: July. 56

Malaysia,

and Map 29 Round-headed Brush Tiger Beetle, Neocollyris batesi.

Species Accounts

Golden Brush Tiger Beetle, Neocollyris orichalcina Horn (Plate 4) [Map 30] Identification: Length 12 -15.5 mm; head and pronotum black with blue and violet reflections;

large head and relatively short pronotum; elytra very shiny violet or blue with large and

irregular microsculpturing. Similar to the Bloodspotted Brush Tiger Beetle (N. cruentata), which has bright violaceus-blue elytra and more

extensive and deeper microsculpturing.

Subspecies and morphological variants: One other subspecies has been described from Yunnan, China.

Distribution and habitats:

Southern to northeastern India, Myanmar, Thailand, Laos, and Vietnam. Higher elevation forests* Map 30 Golden Brush Tiger Beede,

(1000-1500 m).

Neocollyris orichalcina.

Behaviour: Unknown.

Seasonality: June to July.

Dark-legged Brush Tiger Beetle, Neocollyris fuscitarsis (Schmidt-Goebel) (Plate 4) [Map 31 ] Identification: Length 13.5 - 18 mm; body completely brilliant blue but rarely violet or green; pronotum short; elytra with uniform shallow rounded pits and diffusely red, oblong spot at each mid-margin; legs light reddish-

brown with dark tibia and tarsi; trochanters reddish or light brown. Similar to the Redtipped Brush Tiger Beetle (N. rufipalpus), which

also has light reddish femora but with black "knees". Also similartothe Sapphire Brush Tiger Beetle (N. saphyrina), which has a larger head,

shorter pronotum, and black trochanters.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern

Map 31 Dark-legged Brush Tiger Beetle, Neocollyris fuscitarsis.

India (120 - 1150 m), Nepal, Myanmar, Malaysia, Indonesia, Thailand, Cambodia,

Laos, Vietnam, and China. Broad-leafed undergrowth shrubs in shady forests and on foliage along paths and roadways in forests.

Behaviour: Runs on leaves and stems of undergrowth bushes.

Seasonality: April to July.

57

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Broad-headed Brush Tiger Beetle, Neocollyris insignis Chaudoir (Plate 4) [Map 32] Identification: Length 17-21 mm; head and black with blue reflections; pronotum and elytra blue rarely brown with

large

violaceus reflections; elytra long, broaden gradually toward the rear, and central surface

covered with deep labyrinth of connected depressions. Trochanters and femora of legs reddish. Similar to Dark-legged Brush Tiger Beetle (A/, fuscitarsis), which is larger in size and has a relatively longer pronotum. Distinguished from the Blue Brush Tiger Beetle (N. smaragdina) which is larger, has a longer pronotum, and its elytra expand toward the rear. Very similar to the Sapphire Brush Tiger Beetle (/V. saphyrina), which is

Map 32 Broad-headed Brush Tiger Beetle, Neocollyris insignis.

larger, the head less square, and the pronotum more elongate.

Subspecies and morphological variants: No subspecies have been described. Distribution and habitats: Northeastern India (Assam, Meghalaya, Sikkim), Nepal, and Bhutan. Forest and forest edge of the Himalayan foothills (500 - 2000 m)

Behaviour: Runs on leaves of undergrowth bushes. Seasonality: April to July.

Blue Brush Tiger Beetle, Neocollyris smaragdina Horn (Plate 5) [Map 33] Identification: Length 15-17.5 mm; head and pronotum dark blue; elytra bright blue and parallel-sided, dark brown trochanters. At one time considered a subspecies of the Broad-headed Brush Tiger beetle (A/, insignis), but separated by the smaller length and narrower body, parallel-sided elytra, shorter pronotum, and brownish trochanters.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Foothills of the Himalayas (1200 - 1500 m) in northeastern India, Bhutan, Nepal, and Myanmar.

Behaviour: Unknown. Seasonality: July. 58

Map 33 Blue Brush Tiger Beetle, Neocollyris smaragdina.

Species Accounts

Sapphire Brush Tiger Beetle, Neocollyris saphyrina Chaudoir (Plate 5) [Map 34] Identification: Length 13.5-17 mm; large head and short pronotum black with blue

reflections; elytra bright blue; trochanters black; femora reddish with rest of leg dark

brown or black. Similar to the Dark-legged Brush Tiger Beetle (N. fuscitarsis), which has a large head, short and more slender pronotum, and dark trochanters.

Subspecies and morphological variants: No subspecies have been described.

and habitats: Himalayan lowlands (450 - 3600 m) of northeastern India, Nepal, Bhutan, Bangladesh, Myanmar,

Distribution

Thailand, Laos, and possibly China. Foothill forests.

Map 34 Sapphire Brush Tiger Beetle, Neocollyris saphyrina.

Behaviour: Unknown.

Seasonality: June to July.

Eminent Brush Tiger Beetle, Neocollyris egregia Naviaux (Plate 5) [Map 35] Identification: Length 17.5-18 mm; body entirely violet; head rounded and wide;

elytra with short and thin light line at each

mid-margin, deep and convoluted pits in surface at middle of elytra; legs dark.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to the base of the Himalayas in northeastern India. Middle elevation forests (350 -1400 m)

Behaviour: Unknown. Seasonality: April to May. Map 35 Eminent Brush Tiger Beetle, Neocollyris egregia. 59

k Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelldae

Blue-lipped Brush Tiger Beetle, Neocollyris similis Lesne (Plate 5) [Map 36] Identification: Length 9.5-14 mm; Head and pronotum blue-black; labrum long and dark blue; elytra blue with dense pits

forming a confluence of polygons toward the centre of the elytra; trochanters dark yellow to light brown; femora reddish.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Rare species in India with records from Tamil Nadu and northeastern states, Nepal, Myanmar,

Thailand, Laos, Vietnam, and China.

Behaviour: Unknown. Seasonality: Unknown.

Map 36 Blue-lipped Brush Tiger Beetle, Neocollyris similis.

Red-tipped Brush Tiger Beetle, Neocollyris rufipalpis Chaudoir (Plate 5) [Map 37] Identification: Length 11 - 14.5 mm; head and pronotum blue, short and wide; elytra blue-green, short and expanded towards the

rear; labrum long and metallic blue; palpi light brown in males and brown with dark tips in the females. Light brown trochanters and reddish femora with black knees and black tibiae and tarsi separate this species from the similar Dark-legged Brush Tiger Beetle (N. fusdtarsis).

The Red-tipped Brush Tiger Beetle has finer punctuation on the elytra and is smaller than the Sapphire Brush Tiger Beetle (N. saphyrina). The similar Long-lipped Brush Tiger Beetle (A/. distincta) has a wider pronotum and coarser punctuation on the elytral surface.

Map 37 Red-tipped Brush Tiger Beetle, Neocollyris rufipalpis.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India, Nepal, Myanmar,Thailand, Malaysia, Indonesia, Laos, Cambodia, Vietnam, and China. Undergrowth broad-leafed bushes, shrubs and bamboos leaves on the edge of forests

Behaviour: Adults are found running on leaves and stems and making short flights between bushes.

Seasonality: Unknown. 60

Species Accounts

Large-necked Brush Tiger Beetle, Neocollyris crassicornis Dejean (Plate 5) [Map 38] Identification: Length 11 - 15.5 mm; large head black with blue reflections; short, thick,

blue pronotum; parallel-sided elytra blue to

violaceous with their surfaces covered by a network of interlocking punctures; short, thick antennae; trochanters dark blue; femora reddish with black "knees".

Subspecies and morphological variants: The population on the Andaman Islands has been described as the subspecies N. a andamana, and it is larger, more elongate, and with coarser elytral sculpturing than the

mainland population.

Distribution

and

habitats:

Northern, southern and northeastern India, Nepal,

Map 38 Large-necked Brush Tiger Beetle, Neocollyris crassicornis; A, N. c. crassicornis, B, N. c. andamana.

Bangladesh, Bhutan, Sri Lanka, Myanmar, Thailand, Laos, Vietnam, Malaysia, Indonesia, China, and Taiwan. Lowland forests (120 - 915 m) on leaves of shrubs along forest edge, near rivers, swampy areas, and in adjacent cultivated areas.

Behaviour: Unknown.

Seasonality: May to October.

Greyish Brush Tiger Beetle, Neocollyris subclavata Chaudoir (Plate 5) [Map 39] Identification: Length

13.5 -

17

mm;

body blue to grey; head large and round;

pronotum long with distinct lines running from side to side on the top surface; indistinct- reddish patch at mid-margins of elytra; tibia of legs dark brown. Similar to the Large-necked Brush Tiger Beetle (N. crassicornis), but the Greyish Brush Tiger Beetle is slenderer, especially the pronotum.

Subspecies and morphological variants: Two subspecies have been described:

N. s. subclavata - blue coloured body and finer elytral microsculpture.

N. s. continentalis - steel grey colour with green reflections and coarser elytral microsculpture.

Map 39 Greyish Brush Tiger Beetle, Neocollyris subclavata-., A, TV. s. subclavata, B, TV s. continentalis. 61

Ai Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Distribution and habitats: N. s. subdavata - India (northern part of Western Ghats, Karnataka, Goa). N. s. continentalis - southern part of the Western Ghats (Kerala, Tamil Nadu, Andaman

Islands).

Behaviour: Unknown.

Seasonality: July.

Brown Brush Tiger Beetle, Neocollyris acuteapicalis Horn (Plate 6) [Map 40] Identification: Length 13 mm; body dark brown with blue reflections along the sides; pronotum long; antennae short; rufous oval patch on mid-margin of elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India.

Behaviour: Unknown. Seasonality: Unknown.

Map 40

Brown Brush Neocollyris acuteapicalis.

Tiger

Beetle,

Tiger

Beetle,

Hairy Brush Tiger beetle, Neocollyris multipilosa Naviaux (Plate 6) [Map 41] Identification: Length 12.2 mm; entirely dull black; long, thin pronotum covered with white setae; elytra covered evenly with microsculpturing.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to Western Ghats of Karnataka in southern

India.

Behaviour: Very rare. Seasonality: Unknown. 41 Hairy Brush Neocollyris multipilosa. Map

62

Species Accounts

Tapered Brush Tiger Beetle, Neocollyris attenuata Redtenbacher (Plate 6) [Map 42] Identification: Length 9.5 - 12.5 mm; head and pronotum black with blue or violet reflections; palpi thin and yellow with brown tips; elytra bright green with regular and dense sculpturing on surface, expanded in width toward the rear; head long and oval.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern and northeastern India, Pakistan, Nepal, Myanmar, Bhutan, and China. Lowland forest edge and gardens of rural areas (350

- 3400 m),

Map 42 Tapered Brush Tiger Beetle, Neocollyris attenuata.

Behaviour: Unknown. Seasonality: May to September.

Slim Brush Tiger Beetle, Neocollyris brancuccii Naviaux (Plate 6) [Map 43] Identification: Length 10.4 - 11.5 mm; head and pronotum dark green; elytra bright green with copper reflections; head and entire body extremely elongated and thin; thin yellowish mark at mid-margins of the elytra; labrum rounded and black with a pale yellow centre.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India (1300 -1400 m).

Behaviour: Unknown. Seasonality: May.

Map 43 Slim Brush Tiger Beetle, Neocollyris brancuccii.

63

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Long-necked Brush Tiger Beetle, Neocollyris brendelli Naviaux (Plate 6) [Map 44] Identification: Length 13.2 mm; head black with violet reflections and elongated with

relatively small eyes; pronotum black with violet reflections and extremely narrow column at the front and bulbous at the back; elytra narrow and very long; labrum rectangular and brown.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India.

Behaviour: Unknown. Seasonality: Unknown.

Map 44 Long-necked Brush Tiger Beetle, Neocollyris brendelli.

Lesser Brush Tiger Beetle, Neocollyris párvulo Chaudoir (Plate 6) [Map 45] Identification: Length 8-10.5 mm; bright blue-violet; head narrow; pronotum round and narrow in the front and expanded to the rear; elytra widen distinctly toward the rear. Similar to the Long-legged Brush Tiger Beetle (N. maindroni), which is greenish

in colour, larger, has a more constricted pronotum, and longer legs.

Subspecies and morphological variants: No subspecies have been described, but

some individuals are greenish or olive­ green, which may be related to age.

Distribution and habitats: Endemic to southwestern India. Lowland forest edge

(600-4500 m).

Behaviour: Unknown. Seasonality: May to October.

Map

45 Lesser Brush Neocollyris parvula.

Tiger

Beetle,

Species Accounts

Long-legged Brush Tiger Beetle, Neocollyris maindroni Horn (Plate 6) [Map 46] Identification: Length 8-10 mm; dark green to blackish with green to copper reflections; head narrow and elongated; pronotum extremely long and thin with

distinct cross lines; elytra short; tibiae and tarsi significantly longer than similar species such as the Lesser Brush Tiger Beetle (A/. parvula), which also differs in body colour (blue-violet) with a distinctly shorter and stouter pronotum as well as shorter legs.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India (80 m).

Behaviour: Unknown.

Map 46 Long-legged Brush Tiger Beetle, Neocollyris maindroni.

Seasonality: May to July.

Short-necked Brush Tiger Beetle, Neocollyris kollari Horn (Plate 6) [Map 47] Identification: Length 9.5 - 11.5 mm; bright blue; head rounded; pronotum short; elytral surface with uniform small pits. Similar to the Diamond-headed Brush Tiger Beetle (N. variicornis), which has a shorter body length, blue colour, shorter pronotum with less constriction at the front, and metallic blue labrum. The Lesser Brush Tiger Beetle (N. parvula) has a smaller head, longer and slimmer pronotum with a more obvious constriction at the front, and longer elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from southern, central and northeastern

Map 47 Short-necked Brush Tiger Beetle, Neocollyris kollari.

India.

Behaviour: Unknown. Seasonality: July. 65

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Diamond-headed Brush Tiger Beetle, Neocollyris variicornis Chaudoir (Plate 6) [Map 48] Identification: Length 9.5 - 12.5 mm; triangular-shaped head black with violet reflections; long vase-shaped

pronotum

black with green or blue reflections; elytra bright green, dark blue, or purple often with a reddish mark at mid-margins and another small one at the shoulders.

Subspecies and morphological variants: Despite its considerable variation in colour and broad geographical distribution no

subspecies have been described.

Distribution and habitats: Northeastern India and Andaman Islands, Nepal,Thailand,

Vietnam, and Lowland forests (350 - 1400 m). Laos,

Myanmar,

China.

Map 48 Diamond-headed Brush Tiger Beetle, Neocollyris variicornis.

Behaviour: Unknown. Seasonality: April to July.

Pacholatko's Brush Tiger Beetle, Neocollyris pacholatkoi Sawada and Wiesner (Plate 7) [Map 49] Identification: Length 11.3 - 13.3 mm; head bluish black with short yellow-brown antennae that do not reach back to the elytra; pronotum bluish black and shaped like a flask, covered with distinct white hair-like setae; elytra greenish black with small yellow band across middle, punctures form rows across the centre. Similar to the Diamond­

headed Brush Tiger Beetle (A/, variicornis), which has a sigmoid-shaped aedeagus. The Stout Brush Tiger Beetle in contrast has a straight aedeagus that is bent only at its tip.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to

Map 49 Pacholatko’s Brush Tiger Beede, Neocollyris pacholatkoi.

northeastern India. Lowland forests (500 - 950 m).

Behaviour: Unknown. Seasonality: April to May. 66

Species Accounts

Spindle-necked Brush Tiger Beetle, Neocollyris variitarsis Chaudoir (Plate 7) [Map 50] Identification: Length 10.1 - 11.8 mm; elongate head black with violet or blue reflections; pronotum black with violet or blue reflections and expanding from a narrow front to a distinctly expanded rear; elytra blue or rarely green, long, and parallel­ sided with a narrow reddish transverse line at mid-margins.

Subspecies and morphological variants: Over its extensive range there is variation in head length and body size, but only one other subspecies has been described from southern Thailand.

Distribution and habitats: Northeastern

Map 50 Spindle-necked Brush Tiger Beetle, Neocollyris variitarsis.

Laos, Vietnam, and China. Lowland forests (80-1150 m). India,

Nepal,

Myanmar, Thailand,

Behaviour: Forages on leaves of undergrowth vegetation of dense tropical forests and forest edge, especially in light gaps.

Seasonality: May to July.

Bottle-necked Brush Tiger Beetle, Neocollyris ampuHicollis Horn (Plate 7) [Map 51] Identification: Length

13.5 -

17

mm;

black with violet reflections; head round; pronotum extremely long with a unique and distinct constriction in the top and sides that divides the long narrow front from the extremely expanded rear; elytra long.

Subspecies and morphological variants: No subspecies have been described for this rare and localized species.

Distribution and habitats: Endemic to southern India.

Behaviour: Unknown.

Seasonality: Unknown.

Map 51 Bottle-necked Brush Tiger Beetle, Neocollyris ampullicollis.

67

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Furrow-necked Brush Tiger Beetle, Neocollyris plicicollis Horn (Plate 7) [Map 52] Identification: Length 10-13 mm; head and pronotum bright dark blue; pronotum

crossed by numerous distinct lines; elytra bronze to olive with a large, yellow patch on the mid-margins, and elytra significantly wider to the rear than the front. Somewhat similar to the Diamond-headed Brush Tiger Beetle (N. variicornis), which has narrower elytra and a distinctively narrowed neck (collum) giving it a vase-like shape.

Subspecies and morphological variants: No subspecies for this localized species have been described.

Distribution and habitats: Endemic to southern India (300 m).

Map 52 Furrow-necked Brush Tiger Beetle, Neocollyris plicicollis.

Behaviour: Unknown.

Seasonality: Unknown.

Metallic Brush Tiger Beetle, Neocollyris metallica Naviaux (Plate 7) [Map 53] Identification: Length 9.5 - 11.5 mm; shiny and metallic dark green or blue with

violaceous patches on the front part of elytra;

elytra parallel-sided with small, pale spot at

mid-margins. Similar to the Short-lipped Brush Tiger Beetle (N. bevilabris), which is smaller and narrower, metallic coloured, and

lacks the large light patches on the elytra. NOTE: The species Neocollyris shyamrupi was described by Saha and Halder, and it may be the same species as N. metallica. However, the only known specimen of N. shyamrupi could not be located for direct comparison.

Subspecies and morphological variants: No subspecies have been described.

Map 53 Metallic Brush Tiger Beetle, Neocollyris metallica.

Distribution and habitats: Endemic to southern India. Forests (900 - 1000 m).

Behaviour: Unknown. Seasonality: April to September. 68

Species Accounts

Smooth-necked Brush Tiger Beetle, Neocollyris subtileflavescens Horn (Plate 7) [Map 54] Identification: Length 11 - 12.5 mm; bright dark brown-violet; head round; pronotum short and stout; elytra parallel-sided with small light yellow spot at mid-margins. Similar to the Furrow-necked Brush Tiger Beetle (A/, plicicollis), but distinguished from it by the thicker and shorter pronotum and the more parallel-sided elytra.

Subspecies and morphological variants: At one time considered a subspecies of the Furrow-necked Brush Tiger Beetle (A/, plicicollis), but since elevated to a full species.

Distribution and habitats: Endemic to southern India.

Map 54 Smooth-necked Brush Tiger Beetle, Neocollyris subtileflavescens.

Behaviour: Unknown.

Seasonality: Unknown.

Wrinkle-necked Brush Tiger Beetle, Neocollyris rugata Naviaux (Plate 7) [Map 55] Identification: Length 12.3 mm; head black with violet reflections; pronotum and elytra dark green; pronotum long and slender; elytra expanding slightly to the rear with a yellowish spot at each mid-margin. Similar

to the Smooth-necked Brush Tiger Beetle

(N. subtileflavescens), but distinguished by its longer pronotum and more uniform sculpturing on the elytral surface.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to Western Ghats in India. Foothill forests (550 m).

Behaviour: Unknown.

Map 55 Wrinkle-necked

Brush Tiger

Beetle, Neocollyris rugata.

Seasonality: June.

69

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Yellow-spotted Brush Tiger Beetle, Neocollyris conspicua Naviaux (Plate 7) [Map 56] Identification: Length 9 mm; Head and pronotum black with blue reflections; elytra dark brown with a conspicuous, large yellow

patch at mid-margins and a small reddish spot at the shoulders. Resembles the Longlegged Brush Tiger Beetle (N. maindroni), but

differs by the obvious yellow patches on the elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India.

Behaviour: Unknown.

Map 56 Yellow-spotted Brush Tiger Beetle,

Seasonality: September.

Neocollyris conspicua.

Greenish Brush Tiger Beetle, Neocollyris vannideki Naviaux (Plate 7) [Map 57] Identification: Length

10.2 -

11

mm;

coppery green; head round with large eyes; antennae long; extremely long and slender pronotum with a constriction; elytra parallel-sided and bronze with a distinctive patch of yellow-red on the mid-margins. Similar to Furrow-necked Brush Tiger Beetle (N. plicicollis) but differs from it with long antennae, long, thin pronotum with constriction, and parallel-sided elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to Ghats of southern elevation (1000 m) forests.

Western

Behaviour: Unknown.

Seasonality: May to September. 70

India. Mid

Map 57 Greenish Brush Tiger Beetle, Neocollyris vannideki.

Species Accounts

Least Brush Tiger Beetle, Neocollyris flava Naviaux (Plate 8) [Map 58] Identification:

Length 10 mm; head round and bright green; pronotum bright green and short; elytra brown with copper

reflections and a large yellow patch at each mid-margin, sides expanding toward the rear. Somewhat similar to the Greenish Brush Tiger Beetle (A/. vannideki), but it has a larger

head, shorter pronotum, elytra that expand toward the rear and large yellow patches on the elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India.

58 Least Neocollyris flava. Map

Brush

Tiger

Beetle,

Behaviour: Unknown. Seasonality: Unknown.

Blackish Brush Tiger Beetle, Neocollyris juengeri Naviaux (Plate 8) [Map 59] Identification: Length 15.5 - 16 mm; bright blackish grey; head long with large eyes; pronotum long with strong cross striations;

elytra long and expanding slightly toward

rear; surface evenly covered with dense polygons.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India.

Behaviour: Unknown. Seasonality: Unknown. Map 59 Blackish Brush Tiger Beetle, Neocollyris juengeri.

71

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Charcoal Brush Tiger Beetle, Neocollyris anthracina Naviaux (Plate 8) [Map 60] Identification: Length 14-17 mm; head round and pronotum extremely long and constricted, both parts shiny black with green reflections; elytra long, thin, parallel­ sided and matte black with shades of dark green.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to Western Ghats of southern India. Mid­ elevation forests (600 -1000 m).

Behaviour: Unknown.

Seasonality: June.

Map 60 Charcoal Brush Tiger Beetle, Neocollyris anthracina.

Brown-kneed Brush Tiger Beetle, Neocollyris nilgirica Fowler (Plate 8) [Map 61] Identification: Length 14.5 mm; greenishbronze; large head with long antennae; pronotum long and slender with strong

cross striations; elytra long and narrow with yellow mark at each mid-margin.

Subspecies and morphological variants: The type specimen has apparently been lost, and Fowler's description raises some confusion as to how it is distinguished from the Small-eyed Brush Tiger Beetle (N. andrewsi) and the Charcoal Brush Tiger

Beetle (N. anthracina). Neocollyris fowler! was recently described by Naviaux, but it appears to be the same species as the Brown-kneed

Map 61 Brown-kneed Brush Tiger Beetle,

Brush Tiger Beetle (A/, nilgirica).

Neocollyris nilgirica.

Distribution and habitats: Southern India.

Behaviour: Unknown.

Seasonality: Unknown.

72

Species Accounts

Small-eyed Brush Tiger Beetle, Neocollyris andrewesi Horn (Plate 8) [Map 62] Identification: Length 17 - 24 mm; black with blue reflections; head long and eyes relatively small; pronotum long and stout with distinct

constriction; elytra expanding slightly toward rear.

Subspecies and morphological variants: Two subspecies have been described from India: N. a. andrewesi - smaller (17-19.5 mm). N. a. regia - much larger (18 -24 mm) with a

somewhat different pattern of sculpturing on the elytral surface and some minor

differences in mouth parts.

Distribution and habitats: Both subspecies occur in mid-altitude forests (900 - 1500 m):

Map 62 Small-eyed Brush Tiger Beetle,

N. a. andrewesi - Western Ghats of Karnataka.

Neocollyris andrewesi-, Ay N. a. andrewesi, B, N. a. regia.

N. a. regia - Western Ghats of Kerala and Tamil Nadu.

Behaviour: Unknown.

Seasonality: April to November.

Chestnut Brush Tiger Beetle, Neocollyris rubens Bates (Plate 8) [Map 63] Identification: Length 15-17 mm; chestnut brown; head darker and broad; pronotum strongly constricted forming two humps;

elytral surface smooth except for 5 or 6 deep and sharp wrinkles in the middle; elytral edges slightly constricted in the middle and with small indentations; antennae and legs dark brown to blackish. Similar to the Narrow­ necked Brush Tiger Beetle (A/, compressicollis), but distinguished from it by the deep elytral surface sculpturing extending almost the entire length of the elytra in the Narrow­

necked Brush Tiger Beetle.

Subspecies and morphological variants:

Map 63 Chestnut Brush Tiger Beetle,

No subspecies have been described.

Neocollyrts rubens.

Distribution and habitats: Northeastern India and Vietnam. Behaviour: Unknown.

Seasonality: Unknown. 73

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Narrow-necked Brush Tiger Beetle, Neocollyris compressicollis Horn (Plate 8) [Map 64] Identification: Length 15-17 mm; chestnut brown; head darker and broad; pronotum

strongly constricted forming two humps;

elytral surface with polygons and wrinkles almost the entire length; elytral edges slightly constricted in the middle and with small indentations; antennae and legs dark brown to blackish. Similar to the Chestnut Brush Tiger Beetle (N. rubens), but

distinguished from it by the deep elytral surface sculpturing being limited to the

middle of the elytra in the Chestnut Brush Tiger Beetle.

Subspecies and morphological variants:

Map 64 Narrow-necked Brush Tiger Beetle, Neocollyris compressicollis.

No subspecies have been described.

Distribution and habitats: Northeastern India (700 - 1150 m), Laos, Vietnam, and China.

Behaviour: Seen in flight across paths in forest and searching for prey on broadleafed shrubs at forest edge.

Seasonality: April to June.

Long-backed Brush Tiger Beetle, Neocollyris apteroides Horn (Plate 8) [Map 65] Identification: Length 14-18 mm; shiny dark brown; head round; pronotum short with distinct dilation toward the front; elytra long and only slightly expanded toward the rear; elytral surface with deep polygons in the middle and a faint reddish spot at mid­

margin and at the shoulders.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India.

Behaviour: Unknown. Seasonality: Unknown.

Map 65 Long-backed Brush Tiger Beetle, Neocollyris apteroides.

74

Species Accounts

Pitted Brush Tiger Beetle, Neocollyris foveifrons Horn (Plate 9) [Map 66] Identification: Length 15-18 mm; shiny red to matte brown or occasionally bluish; head with exceptionally deep excavation between eyes; pronotum short and stoutlong and expanded toward the

elytra

rear; elytral surface with distinct folds and wrinkles in the middle.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India to Vietnam.

Behaviour: Unknown.

Seasonality: Unknown.

Map

66 Pitted Brush Neocollyris foveifrons.

Tiger

Beetle,

Tiger

Beetle,

Assam Brush Tiger Beetle, Neocollyris assamensis Naviaux (Plate 9) [Map 67] Identification: Length 12-18 mm; dark reddish brown; large round head with distinct excavation between eyes; pronotum long with distinct constriction near frontelytra exceptionally long with moderate

widening at rear; elytral surface with deep

folds restricted to middle.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India. Middle to upper altitude forest edges (500 - 2600 m) along the Himalayan foothills.

Behaviour: Unknown.

Map 67 Assam Brush Neocollyris assamensis.

Seasonality: April to August. 75

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Rufous Brush Tiger Beetle, Neocollyris coapteroides Horn (Plate 9) [Map 68] Identification: Length 16.3 mm; head and pronotum brownish; elytra bright reddish brown, long, slender, and slightly widened at the rear.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India.

Behaviour: Unknown.

Seasonality: April to May.

Map 68

Rufous Brush Neocollyris coapteroides.

liger

Beetle,

Thin-spotted Brush Tiger beetle, Neocollyris smith" Chaudoir (Plate 9) [Map 69] Identification: Length 18.5 - 21 mm; large dark bluish-black head; long and stout dark blue pronotum; elytra long and dark

blue with large light brown patches at the shoulders and rear ends.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India and Myanmar. Lowland forests (500 -

1400 m).

Behaviour: Unknown.

Seasonality: April to May. Map 69 Thin-spotted Brush Tiger Beetle, Neocollyris smithii. t

76

Species Accounts

Large Brush Tiger Beetles, genus Collyris These are the largest of the Brush Tiger Beetles with body lengths from 17-26 mm. They tend to occupy higher parts of the undergrowth and brush vegetation. In flight they are conspicuous by their size and slow and steady rise to cover relatively long distances. Of the ten species currently placed in this genus, five occur in India, of which two are endemic.

Key V: Large Brush Tiger Beetles, Collyris Identification Key to Collyris Species la.

Punctured Brush Tiger

Foot of hind leg almost yellow

Beetle, Collyris dormeri

1b. 2a. (lb.)

2b.

Foot of hind leg completely black

2

Labrum yellow, basal border and the outer

Short-winged Brush Tiger

margins of the five middle teeth darkened

Beetle, Collyris brevipennis

Labrum

completely dark

and

outer

3

margins of the five middle teeth never

darkened 3a. (2b.)

Smooth Brush Tiger Beetle,

Tibia partly brownish

Collyris subtilesculpta 3b.

4a. (3b.)

4

Tibia completely black

Pronotum rather short, with the base wide,

Notch-lipped Brush Tiger

strongly contracted towards the neck

Beetle, Collyris longicollis

(collum) (Fig. 6.30); body length 17-25 mm

4b.

Pronotum elongated, more smoothly and

Violet Brush Tiger Beetle,

less constricted towards the neck (collum)

Collyris dohrnii

(Fig. 6.31); body length 22-26 mm

Figure 6.30 neck (collum).

Pronotum short with distinct

Figure 6.31 Pronotum elongated with less distinct neck (collum).

77

k Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Notch-lipped Brush Tiger Beetle, Collyris longicollis Fabricius (Plate 9) [Map 70] Identification: Length 17-25 mm; blue to dark blue or blackish; head large and short; pronotum relatively short and stout with cross striations; elytra parallel-sided with

metallic blue and rounded pits in the front and rear and polygons in the middle.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern and southern India and Nepal. Foothill forests above 500 m.

Behaviour: Runs on leaves and small branches of tall bushes and lower parts of trees. Reluctant to fly, but when it does, the

Map 70 Notch-lipped Brush Tiger Beetle, Collyris longicollis.

flight can be long (10-100 m). At rest or in

flight resembles a dark blister beetle.

Seasonality: August to October.

Violet Brush Tiger Beetle, Collyris dohrnii Chaudoir (Plate 9) [Map 71] Identification: Length 22 - 23 mm; violaceus brown; large head; short, thick pronotum smooth on top; elytra long and gradually widening toward the rear, trochanters and base of femora dark brown. Distinguished from the Notch-lipped Brush Tiger Beetle (C. longicollis) which is larger, violaceous-brown

colour, longer pronotum, and larger head.The

very similar Short-winged Brush Tiger Beetle (C. brevipennis) has light red trochanters.

Subspecies and morphological variants: The population in India has been named C. d. indica. It is smaller, more purple than black, and its pronotum is smoother than the nominate population found on Sri Lanka.

Map 71 Violet Brush Tiger Beetle, Collyris dohmii.

Distribution and habitats: Southern India in Western Ghats forests (600 - 900 m) and Sri Lanka.

Behaviour: Unknown.

Seasonality: May. 78

Species Accounts

Short-winged Brush Tiger Beetle, Collyris brevipennis Horn

(Plate 9) [Map 72]

Identification: Length 19-24 mm; bright blue; head round with deep excavation between eyes; pronotum long with obvious

cross striations; elytra I surface with deep polygons in the middle and more rounded toward the front and rear. The very similar Violet Brush Tiger Beetle (C. dohrnii) has dark brown trochanters. From the Notch­ lipped Brush Tiger Beetle (C. longicollis) it is

distinguished by a less dilated pronotum with deeper cross striations, and shorter elytra. Differs from the Smooth Brush Tiger Beetle (N. subtilesculpta) by its deeper and more obvious sculpturing on the middle of the elytra.

Map 72 Short-winged Brush Tiger Beetle, Collyris brevipennis.

Subspecies and morphological variants:' No subspecies have been described.

Distribution and habitats: Known only from the Western Ghats of southern India. Forests (1300 m).

Behaviour: Forages on branches and leaves of small bushes. Observed inserting eggs into small branches of these bushes.

Seasonality: October to November.

Smooth Brush Tiger Beetle, Collyris subtilesculpta Horn

(Plate 9) [Map 73]

Identification: Length 21-26 mm; head round, large, and black or violet; pronotum dark blue, long and larger at the base; elytra dark to bright blue, long, parallel-sided,

and with relatively small surface pits along their length. This species was originally described as a variety of the Short-winged Brush Tiger Beetle (N. brevipennis) and is readily distinguished from it by having finer sculpturing on the elytral surface and its more southerly distribution.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only

Map 73 Smooth Brush Tiger Beetle, Collyris subtilesculpta.

from the Western Ghats of southern India. Forests (900 - 1200 m).

Behaviour: Unknown.

Seasonality: Unknown. 79

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Punctured Brush Tiger Beetle, Collyris dormeri Horn (Plate 9) [Map 74] Identification: Length 21-23 mm; head and stout pronotum black with violet reflections; elytra violaceous brown with extensive punctures and folding on the central third of the elytral surface; antennae longer than the other species of Large Brush

Tiger Beetles (Collyris) in India as they reach back almost to the elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India, Myanmar and Laos.

Behaviour: Unknown.

Map 74 Punctured Brush Tiger Beetle,

Seasonality: Unknown.

Collyris dormeri.

Knobby Tiger Beetles, genus Therates Adults of this genus range in size from 5 to 23 mm in length. They vary in colour from all black to various shades of dark blue with orange or yellow patterns on the elytra. Adults typically have or more rounded protuberances (humps) on the surfaceof the forward part ofthe elytra. They also have peculiarly reduced mouth parts except for an immense labrum that hides the closed mandibles. They forage on moist forest floors and quickly fly to leaves of undergrowth bushes to escape danger. The more than 70 species occur from Nepal to the Solomon Islands and Taiwan. Ten species occur in India, all in the northeastern quarter ofthe country. All but one of them are endemic to India.

Key VI: Knobby Tiger Beetles, genus Therates Identification Key to Therates Species 1a.

Basal segments of the hind leg foot

Hennig's Knobby Tiger Beetle,

much thicker than the same ones of

Therates hennigi

the middle leg

1b.

Segments of the hind leg foot of the same size as those ofthe middle leg

80

2

Species Accounts

2a (1b.)

Elytra with a narrow pale yellow stripe

Wrinkled Knobby Tiger

running along each side from the

Beetle, Therates westbengalensis

shoulder two-thirds length of elytra

2b.

Lacks long yellow stripe along length

3

of elytra

3a (2b.)

Brown-vented Knobby Tiger

Underside completely pale

Beetle, Therates dohertyi 3b.

Underside partly to almost dark

4a. (3b.)

Forehead (frons) mostly reddish

4

Red-fronted Knobby Tiger Beetle, Therates waagenorum

4b. 5a. (4b.)

5

Forehead (frons) mostly black

Elytra markings variable but connected

5b.

Brown-bellied Knobby Tiger Beetle, Therates sausai

throughout Elytra markings isolated or at most

6

humeral lunule and basal dot connected at base and/or humeral lunule connected

with central dot 6a. (5b.)

Central dot of elytra subsquare or a

7

trapezoidal band 6b.

Central dot of elytra rounded, not

8

forming a band 7a. (6a.)

Labrum uniformly yellow

Yellow-spotted Knobby Tiger Beetle, Therates nepalensis

7b.

Labrum dark at outer margin

Dark-lipped Knobby Tiger Beetle,

Therates arunachalcolus 8a. (6b.)

Central dot of elytra acutely angled

8b.

9a. (8b.)

9b.

Ingrid's Knobby Tiger Beetle, Therates ingridae

outwards toward the front

Central dot of elytra not as above

9

Humeral lunule long, connected with

Brown-legged Knobby Tiger

the central dot or nearly reaching it

Beetle, Therates annandalei

Humeral

lunule

short,

neither

connected to nor reaching near the

Brown-spotted Knobby Tiger

Beetle, Therates jendeki

central dot

81

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Brown-legged Knobby Tiger Beetle, Therates annandalei Horn (Plate 10) [Map 75] Identification: Length 7.2 - 8.2 mm; Head and pronotum shining greenish-black; elytra

shining black with long brownish yellow front mark, small yellow central dot, and a light yellow rear patch that covers the rear humps; labrum yellowish.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India. Damp, shady places

among undergrowth forests (200 -1500 m).

plants

of foothill

Behaviour: Unknown.

Seasonality: July.

Map 75

Brown-legged Knobby Tiger Beetle, Therates annandalei.

Yellow-spotted Knobby Tiger Beetle, Therates nepalensis Probst and Wiesner (Plate 10) [Map 76] Identification: Length 7.7 - 8.2 mm; head and prothorax shiny greenish-black; elytra shiny greenish or violet black with orangish

markings in front and yellow in the middle and rear; labrum yellowish.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India and Nepal. Foothill forests (1800 m).

Behaviour: Unknown. Seasonality: June. Map 76 Yellow-spotted Knobby Tiger Beetle, Therates nepalensis.

82

Species Accounts

Brown-spotted Knobby Tiger Beetle, Therates jendeki Sawada and Wiesner (Plate 10) [Map 77] Identification: Length 6.7 - 9.1 mm; head and thorax shiny greenish-black; elytra

shiny black with broad brownish yellow

front marking, right-angled brownish yellow central dot, and an obscure brown rear patch; labrum yellowish with blackish outer margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India. Foothill forests (500 1400 m).

Behaviour: Unknown.

Map 77 Brown-spotted Knobby Tiger

Seasonality: April to June.

Beetle, Therates jendeki.

Dark-lipped Knobby Tiger Beetle, Therates arunachalcolus Sawada and Wiesner (Plate 10) [Map 78] Identification: Length 7.8 mm; head and pronotum shiny greenish-black; elytra shiny

brownish-black with long yellow front mark, broad sigmoid-shaped middle band, and yellow rear mark that reaches the humps; labrum yellowish with darkened marginal base.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India. Foothill forests (350 m).

Behaviour: Unknown.

Seasonality: April to May.

Map 78 Dark-lipped Knobby Tiger Beetle, Therates arunachalcolus.

83

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Ingrid's Knobby Tiger Beetle, Therates ingridae Sawada and Wiesner (Plate 10) [Map 79] Identification: Length 4.8 - 6.7 mm; shiny black; elytra with yellow, variable-shaped front mark, yellow central dot, and indistinct

pale crescent at rear; labrum yellowish with

darkened marginal base.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India. Foothill forests (500 950 m)of

Behaviour: Unknown. Seasonality: April. Map 79 Ingrid’s Knobby Tiger Beetle, Therates ingridae.

Brown-bellied Knobby Tiger Beetle, Therates sausai Sawada and Wiesner (Plate 10) [Map 80] Identification: Length 6.2 - 8.1 mm; Head shiny greenish-black with reddish forehead; pronotum greenish-black; elytra shiny brownish-black with large 'T-shaped brownish-yellow front mark, yellow horizontal central dot, and pale yellow rear patch; labrum yellowish with darkened marginal base.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India. Foothill forests (500 -

1300 m).

Behaviour: Unknown. Seasonality: April to May.

84

Map 80

Brown-bellied Beetle, Therates sausai.

Knobby Tiger

Species Accounts

Brown-vented Knobby Tiger Beetle, Therates dohertyi Horn (Plate 10) [Map 81] Identification: Length 6.3 - 7.6 mm; head and prothorax shiny greenish-black; elytra shiny brownish-black with brownish-yellow

"C"-shaped front mark, short yellow central band, and light yellow rear patch; labrum yellowish; rear end of abdomen brownish.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India. Foothill forest (500 1500 m)of

Behaviour: Unknown.

Seasonality: May.

Map 81

Brown-vented Beetle, Therates dohertyi.

Knobby Tiger

Red-fronted Knobby Tiger Beetle, Therates waagenorum Horn (Piatelo) [Map 82] Identification: Length 5.7 - 6.2 mm; head shiny greenish-black with extensive reddish

forehead; pronotum shiny greenish-black; elytra shiny brownish-black with brownishyellow front mark sometimes separated into

three large dots or coalescing to connect to each other as well as with the yellow horizontal central dot, and pale yellow rear

patch; labrum yellowish with brownish sides.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India. On moist forest floor above 600 m.

Map 82 Red-fronted Knobby Tiger Beetle, Therates waagenorum.

Behaviour: Forages on forest floor but to escape danger flies up to land on leaves of undergrowth 50 - 80 cm high.

Seasonality: Unknown.

85

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Wrinkled Knobby Tiger Beetle, Therates westbengalensisWiesner (Piatelo) [Map 83] Identification: Length 6.2 mm; head and pronotum greenish-black; elytra yellowbrown with brighter yellow patches at the rear end and a long narrow stripe running

along the side from the shoulder two thirds the length of the elytra.

Subspecies and morphological variants: No subspecies of this rare species have been

described.

Distribution and habitats: Known only from Darjeeling in northeastern India. Forest

floor (1500 m).

Behaviour: Unknown. Seasonality: Unknown.

Map 83 Wrinkled Knobby Tiger Beetle, Therates westbengalensis.

Hennig's Knobby Tiger Beetle, Therates hennigi Horn (Plate 11) [Map 84] Identification: Length 11.0 - 12.1 mm; head and pronotum shiny black with blue-green or violet reflections; almost the entire front

half of the elytra orange, rear half of the elytra with two large patches of shiny black

in the middle and rear separated by a broad bright yellow middle band extending the entire width of the elytra and a large yellow area at the rear end of the elytra.

Subspecies and morphological variants: Two subspecies have been described:

T. h. hennigi - orange front half of elytra connected to the yellow central band by a narrow bright strip.

Map 84 Hennigs Knobby Tiger Beede, Therates hennigi', A, T. h. hennigi, B, T. h. dormeri.

T. h. dormeri - orange front half of elytra separate from central band.

Distribution and habitats: Endemic to low to mid-elevation forests of Northeastern India (350-1200 m):

T. h. hennigi - western part of northeastern India (Meghalaya, Assam). T. h. dormeri - eastern part of northeastern India (Arunachal Pradesh, Nagaland, Manipur).

Behaviour: Often forages on open substrate and undergrowth in forests of large trees near tops of ridges.

86

Seasonality: April to July.

Species Accounts

Wrinkled Tiger Beetles/ genus Rhytidophaena This peculiar genus has only four species. Two of them are found in India, and one is endemic. They are usually collected at night lights and little is known of their biology. They are spectacularly coloured with bright metallic blues, greens and reds.

Key VIII: Wrinkled Tiger Beetles, genus Rhytidophaena Identification Key to Rhytidophaena Species 1 a.

Ornamental Wrinkled Tiger

Centre of elytra shiny, red-coppery

Beetle, Rhytidophaena limbata 1 b.

Centre of elytra dull, blue-violet to red-

Plain Wrinkled Tiger Beetle, Rhytidophaena inornata

violet

Ornamental Wrinkled Tiger Beetle, Rhytidophaena limbata Wiedemann (Plate 11) [Map 85] Identification: Length 9-10 mm; bright and shiny crimson, occasionally with violet tinge on shoulders; pronotum squared and green or bright blue on its sides; central line

between the elytra and sides green with two, distinct white spots, one at mid-margin and another just before the rear end; underside shiny, bright blue or violaceous; legs metallic green or black; labrum white in male and dark metallic in the female.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and

habitats:

Northern

and northeastern India (800 m), Nepal,

Map 85

Ornamental Wrinkled Tiger Beetle, Rhytidophaena limbata.

Bangladesh, and Pakistan.

Behaviour: Unknown.

Seasonality: July.

87

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Plain Wrinkled Tiger Beetle, Rhytidophaena inornata Horn (Plate 11) [Map 86] Identification:

Length

10

mm;

head

dark coppery-bronze with purple irons;

pronotum

bronze-green,

centre

purple,

dull blue to red-violet, margins greenish with two, distinct white spots at mid-margin and another near the rear end; underside of body bluish and legs bronze. elytra

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northern India. Floor of open forest and adjacent fields (200 m).

Behaviour: Often attracted to lights at night.

Map 86 Plain Wrinkled Tiger Beetle,

Seasonality: August.

Rhytidophaena inornata.

Apothecary Tiger Beetles, genus Prothyma This is a somewhat ambiguous genus whose tentative species members share only a few characters, such as, a lack of white, hair-like setae on the underside of the body. This genus presently consists of a mix of species that requires further studies to reveal their actual evolutignary relationships. Most are active during the day on open ground and fly short distances to escape danger. However, they are regularly attracted to lights at night. They are found in a wide range of habitats from open savanna to forest floor. Species representing this genus are found from central Africa to Taiwan and the Philippines. As presently defined, there are 40 species in this genus. Only two species occur in India, neither of which is endemic.

Key VII: Apothecary Tiger Beetles, genus Prothyma Identification Key to Prothyma Species la.

1b.

88

Elytra

with

three

white

spots

--

Assam Tiger Beetle,

shoulder, mid-margin and near the rear

Prothyma assamensis

Elytra with two small white spots - mid­

Azure Tiger Beetle,

margin and near the rear

Prothyma proximo

Species Accounts

Azure Tiger Beetle, Prothyma proximo Chaudoir (Plate 11) [Map 87] Identification: Length 10.5-12 mm; head and

pronotum with

distinct transverse

grooves, dark green to black; elytra dark blue; two, small white spots along outer elytra margin; undersides and femora metallic blue or green.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and

habitats:

Widely

in India from Gujarat to Darjeeling south throughout peninsular India (1000 m) and Sri Lanka.

distributed

Behaviour: Unknown.

Map 87 Azure Tiger Beetle, Prothymapróxima.

Seasonality: July to November.

Assam Tiger Beetle, Prothyma assamensis Rivalier (Plate 11) [Map 88] Identification: Length 13 mm; upper surface of head, pronotum and elytra shiny olive with blue-green reflections; pronotum very shiny with fine transverse grooves; three, pale whitish spots on outer edges of elytra, one each at the shoulder, mid­ margin, and near the rear; undersides dark blue-green.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India and Nepal. On or near forest floor (120 -1150 m)

Behaviour: Unknown.

Seasonality: April to October.

Map 88 Assam Tiger Beetle, Prothyma assamensis.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Jungle Tiger Beetles, genus Heptodonta Most species of this genus are greenish-olive and lack any white hair-like setae on the underside of their bodies except as a fringe at the base of the hind legs. They have a characteristic labrum with seven pointed tooth-like projections. Active during the day, they feed on the ground of moist open forest but fly to leaves of undergrowth bushes to escape danger. The 10 species in this genus occur from northeastern India to the Philippines. Only a single species is found in northeastern India, and it is endemic.

Beautiful Jungle Tiger Beetle, Heptodonta pulchella Hope (Plate 11) [Map 89] Identification: Length 15-17 mm; dull olive-green; elytra dull with fine sculpturing; upper leg parts reddish, lower leg parts black.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and

habitats:

Northern

and northeastern India, Nepal, Myanmar,

Vietnam, Laos, and China. Bare soil, agricultural fields, and forest clearings near streams (200 - 2500 m).

Behaviour: Unknown. Seasonality: April to August.

Map 89 Beautiful Jungle Tiger Beetle, Heptodonta pulchella.

Species Accounts

Glossy Tiger Beetles, genus Pronyssa This genus has eight species. Five of them occur within India and four are endemic to northeastern India. It is closely related to the Jungle Tiger Beetles (Heptodonta) but separated by the extremely bright metallic reflections on its elytra and the spheroid shape of its pronotum.

Key X: Glossy Tiger Beetles, genus Pronyssa Identification Key to Pronyssa Species 1a.

Larger (body length > 14 mm)

Hennig's Glossy Tiger Beetle, Pronyssa hennigi

1b. 2a. (1b.)

Smaller (body length < 14 mm)

2

Elytra colour pure green, with blackish

green reflections 2b.

Green Glossy Tiger Beetle, Pronyssa nodicollis

Elytra colour not pure green, lateral margins

3

blue-violet 3a. (2b.)

Central dot on elytra forming a dash

Montane Glossy Tiger Beetle, Pronyssa montanea

3b.

Central dot on elytra roundish

4

4a. (3b.)

Sides of female labrum yellow

Assam Glossy Tiger Beetle, Pronyssa assamensis

4b.

Sides of female labrum dark metallic

Kraatz's Glossy Tiger Beetle,

Pronyssa kraatzi

Hennig's Glossy Tiger Beetle, Pronyssa hennigi (Horn) (Plate 11) [Map 90] Identification: Length 14.8 mm; head and pronotum dark, coppery green; elytra shiny coppery-green with three white spots along

margin; legs long, red to dark. Labrum shiny blue-violet in the centre.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India.

Behaviour: Unknown.

Seasonality: Unknown.

Map 90 Hennig’s Glossy Tiger Beetle, Pronyssa hennigi. 91

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Green Glossy Tiger Beetle, Pronyssa nodicollis Bates (Plate 11) [Map 91] Identification: Length 11.3 - 13.5 mm; clear green with black-green reflections; pronotum short and rounded on sides; elytra parallel-sided, tapering at the rear to a

point, three white spots along each margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India, Nepal, Myanmar, Vietnam, and China.

Dirt roads and open forest floor (500 - 2600 m).

Behaviour: When disturbed on flat ground often flies to adjacent vertical slopes. Seasonality: April to July.

Map 91

Green Pronyssa nodicollis.

Glossy Tiger

Beetle,

Kraatz's Glossy Tiger Beetle, Pronyssd kraatzi Horn (Plate 12) [Map 92] Identification: Length 10.2 - 12.5 mm; bluish green; pronotum rounded on sides; elytra with blue sides and three (male) or

two (female) white spots.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India. Fields and forest edge (500 - 3000 m).

Behaviour: Unknown.

Seasonality: June to September. Map 92 Kraatz’s Glossy Tiger Beetle, Pronyssa kraatzi.

92

Species Accounts

Assam Glossy Tiger Beetle, Pronyssa assamensis Sawada and Wiesner (Plate 12) [Map 93] Identification: Length 12.2 - 12.3 mm; copper brown with blue-violet reflections;

pronotum narrower at the front than the back; elytra sides blue with two, distinct white spots along margins, one at the middle one at the rear.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India (500 - 950 m).

Behaviour: Unknown. Seasonality: April to May. Map 93 Assam

Glossy Tiger

Beetle,

Pronyssa assamensis.

Montane Glossy Tiger Beetle, Pronyssa montanea Sawada and Wiesner (Plate 12) [Map 94] Identification: Length 11-13.3 mm; blue­ green; elytra green with blue-violet sides and three white spots along margin, the middle one a horizontal dash.

Subspecies and morphological variants: No subs’pecies have been described.

Distribution and habitats: Endemic to northeastern India (500 - 1150 m).

Behaviour: Flies up from forest floor to land on leaves of undergrowth bushes when in

danger (Plate 32.1).

Seasonality: April to July. Map 94 Montane Glossy Tiger Beetle, Pronyssa montanea.

93

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Subfamily Cicindelinae, Tribe Cicindelini, Subtribe Cicindelina This group includes almost 900 of the 2900 known species of tiger beetles

worldwide. Adults range tremendously in size (5 -25 mm) and colour and live in habitats from forest and alpine to desert grassland and ocean beaches. They occur on every continent except Antarctica and on all but the most isolated oceanic islands. A great variety of soil surface types are used as foraging habitat by adults and sites for burrowing larvae. Adults of most species in this group forage for food and mate in open areas with little vegetation, but some live on the floor of tropical rain forests. Others are so specialized they occur only on cliff edges, large boulder tops

or on fallen logs in moist forests. They quickly respond to predators and danger

by flying short distances, although a few flightless species scurry under whatever cover they can find to escape. All the species in this group fall within a classification system, based primarily on characters found within the male genitalia, that was devised by Rivalier. He

organized the species of this group into 65 sub-groups that have been variously

considered genera or subgenera.This field guide follows recent molecular analyses which support most of Rivalier's groups as genera, although these molecular analyses also redefined several of Rivalier's groups. Many species in this group have geographical populations that have been

described as subspecies. These sub-populations usually show intermediate forms where their ranges meet. The adult characters that most often are used to distinguish the subspecies include: body colour, elytral pattern of markings, and body size. However, as discussed in Chapter 3, these characteristics may not accurately define evolutionarily distinct groups. Within some populations individuals with distinctly different forms, such

as body colour, occur together (polymorphism), and many populations have

occasional individuals that exhibit rare colour forms (melanism) or unusual elytral patterns. Many species show considerable size difference between the sexes (females generally larger than males), and in a few species, the sexes exhibit noticeable differences in pronotum, elytra and mandible shape (sexual dimorphism).

Beautiful Tiger Beetles, genus Calochroa This genus is represented by some of the largest species of tiger beetles found in India. They share a colourful body that displays striking patterns of large stripes and spots on the elytra. However, the diversity of many other features exhibited by the species in this genus suggests that more detailed studies are needed to reliably determine their relationships. As presently defined, 11 species of Beautiful Tiger Beetles occur in India, and four of them are endemic.

94

Species Accounts

Key XI: Beautiful Tiger Beetles, genus Calochroa Identification Key to Calochroa Species la.

Very large brilliant, multi-coloured body;

Greater Spotted Tiger

elytra marked with a spot at shoulders and

Beetle,

several large yellow irregular spots; second

Calochroa octonotata

small segment of front and middle legs each with a single (on a few specimens double) subapical seta

lb.

Medium to large uniformly coloured darker

2

body; elytra entirely unmarked or marked

with a dot at shoulders or not and several small yellow nearly oval spots or a single

band at middle; second small segment of front and middle legs without a subapical

seta 2a. (1b.)

Narrow lateral strips along sides of elytra mostly

pale

tawny

and

nonmetallic,

Blackish Tiger Beetle,

Calochroa tritoma

especially toward front 2b.

Narrow lateral strips along sides of elytra

3

entirely dark, often 3a. (2b.)

Labrum

short,

transverse,

longitudinal

4

dimension not exceeding half transverse dimension; three labral teeth, small to

minute, projecting toward the front 3b.

Labrum elongate, longitudinal dimension

6

exceeding half transverse dimension; three to five labral teeth, moderate to large, projecting toward the front

4a. (3a.)

4b. 5a. (4b.)

Pronotum hairy at lateral margin, remaining

Six-dotted Tiger Beetle,

surface bare

Calochroa sexpunctata

Pronotum entirely bare

5

Elytra almost entirely green except for a

Green Tiger Beetle,

single small dot along lateral margin at

Calochroa whithillii

centre, first antennal segment with one hair

near it tip

95

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

5b.

Elytra black with three large spots in a row;

Yellow-spotted Tiger

first antennal segment with two hairs near

Beetle, Calochroa

its tip

6a. (3b.)

Second small segment of legs unpigmented,

flavomaculata 7

pale tawny 6b.

Second small segment of legs pigmented,

8

dark metallic 7a. (6a.)

7b.

Labrum completely shiny purplish black;

Calochroa assamensis

Labrum partly shiny purplish black, two pale

Orange-striped Tiger

areas at middle on most specimens; elytra

8a. (6b.)

Four-spotted Tiger Beetle,

elytra with spots at shoulder and in middle

Beetle,

with continuous stripe down its middle

Calochroa hamiltoniana

Last abdominal segments reddish orange

Bicoloured Tiger Beetle,

Calochroa bicolor 8b.

Last abdominal segments metallic blue,

9

purple, or black 9a. (8b.)

Elytra without a large spot at shoulder

Black-headed Tiger

(minute dot present on a few specimens),

Beetle,

but with three large orange spots in a line

Calochroa fabriciana

down the centre always separated, labrum

only slightly raised at middle

9b.

Elytra with a large spot at shoulder and row

10

of three large yellow-orange spots down the

centre joined on some specimens, labrum

distinctly ridged in middle 10a. (9b.)

Elytra apex short, distance between apical spot and elytra tip shorter or the same as the

10b.

Beetle,

diameter of apical spot

Calochroa octogramma

Elytra apex elongated, distance between

Safranek's Tiger Beetle,

apical spot and elytra tip much longer than

Calochroa safraneki

the diameter of apical spot

96

Lesser Spotted Tiger

Species Accounts

Yellow-spotted Tiger Beetle, Calochroa flavomaculata (Hope)

(Plate 12) [Map 95]

Identification: Length 13.5 - 16 mm; velvety dark

black

to

greenish

or

bluish-black;

pronotum squarish with no hair-like setae; elytra each with three pale yellow spots down

the length of each elytron half way between the mid-line and the margin. Easily confused with the Six-dotted Tiger Beetle (C. sexpunctata) which has white hair-like setae on the upper edges of the thorax. The ranges of the two species apparently overlap little within India.

Subspecies and morphological variants: Two other subspecies have been described, one from Indonesia and the other from Africa.

Distribution and habitats: Throughout

Map 95

Yellow-spotted Calochroa flavomaculata.

Tiger

Beetle,

India (except desert northwest and most of the east coast), west tp Pakistan and east to Taiwan, and Philippines. Found on grassy meadows, rice paddies, open forest floor and sandy banks of rivers (520 - 2400 m).

Behaviour Predator on rice paddy pests. Often active during rainstorms and readily attracted to lights at night. Climbs grass blades and stems of small bushes to pass the night.

Seasonality: July to January.

Six-dotted Tiger Beetle, Calochroa sexpunctata (Fabricius) (Plate 12) [Map 96] Identification: Length 12-14 mm; head shiny bronze above and metallic green on the sides; pronotum squarish with hair-like setae on the upper edges; elytra dull black with green sides and three oval spots in a row along the mid-line. Easily confused with the Yellowspotted Tiger Beetle (C. flavomaculata), which has no hairs on the side of the thorax.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: East and northwest coast of India, Andaman Islands and Nicobar

Islands, Sri Lanka, Nepal, west to Afghanistan and east to Philippines. Mud along ponds, rice paddies and moist alkali soils (500 -1000 m).

Map 96 Six-dotted Tiger Beetle, Calochroa sexpunctata.

Behaviour Attracted to lights at night. Also regularly climbs grass stems to pass the night or cool, rainy weather. Feeds on rice pests such as the coreid bug, Leptocorisa acuta.

Seasonality: May to October. 97

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Green Tiger Beetle, Calochroa whithillii Hope (Plate 12) [Map 97] Identification: Length 13.5-18 mm; head and pronotum green, blue-green to blue; elytra dull green, blue-green or blackish-

green with shiny green sides and a single

small white dot in the middle near the margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Forested mountain roads, sandy paths, rice paddies, and beaches along rivers (700 - 1900 m) in

Behaviour: Often occurs in rice paddies

Map 97 Green Tiger Beetle, Calochroa whithillii.

the Yellow-spotted Tiger Beetle (C. flavomaculata). Adults occasionally gather in large numbers close together in patches along rivers.

with

Seasonality: May to October.

Greater Spotted Tiger Beetle, Calochroa octonotata (Wiedemann) (Plate 12) [Map 98] Identification: Length 20 - 25 mm; head and pronotum deep purple-blue and metallic green with bright copper-red reflections; elytra blue and green with copper reflections and three large yellow

markings on each elytron.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern and northeastern India, Bhutan, Nepal, Bangladesh,

Myanmar, and China. Gravel and sand banks

of rivers, but moves to floor of adjacent open forest during floods (120 - 520 m).

Behaviour: Noisy fliers that often move

Map 98 Greater Spotted Tiger Beetle, Calochroa octonotata.

long distances. Its bright pattern of colours becomes camouflage among the leaf debris lying on the ground it inhabits.

Seasonality: May to October. 98

Species Accounts

Blackish Tiger Beetle, Calochroa tritoma (Schmidt-Goebel) (Plate 13) [Map 99] Identification: Length 14-17 mm; head and pronotum black on top and blue-green and coppery to purple on the sides; elytra dull black with extensive yellow orange bands running down the elytron from the shoulder and two large yellow dots in the middle and rear of each elytron.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India, Bangladesh, Myanmar, and Thailand. Forest floor openings of foothills (370 m).

Behaviour: Unknown.

Map 99 Blackish Tiger Beetle, Calochroa

Seasonality: April to July.

tritoma.

Lesser Spotted Tiger Beetle, Calochroa octogramma (Chaudoir) (Plate 13) [Map 100] Identification: Length 12-15 mm; head shiny black with bright copper and green reflections above and green and purple on the sides; pronotum black with green

and copper reflections; elytra dull black each with four pale yellow spots, one often hidden below the shoulder, and oval-shaped spot at the front and two more round ones

lined up along the length of each elytron at the middle and rear.

Subspecies and morphological variants: C. o. labionigra has been described from central valleys of Nepal.

Distribution and habitats: Northern and

Map 100 Lesser Spotted Tiger Beetle, Calochroa octogramma.

northeastern India, Nepal, Myanmar, and China. Along streams but moves to moist cultivated fields, grassy meadows, pastures,

and open forests during floods (100 - 3500 m) in

Behaviour: Runs along rocky trails and gravelly areas often on steep slopes. Seasonality: May to July.

99

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Safranek's Tiger Beetle, Calochroa safraneki Werner and Wiesner (Plate 13) [Map 101] Identification: Length 13-16 mm; head coppery;

pronotum

trapezoidal

and

coppery; elytra dark greenish-brown with

one shoulder spot and three larger oval or round spots in a row along the length of each elytron. Similar to the Black­ headed Tiger Beetle (C. fabriciana) which is

distinguished by the absence of the shoulder

spot on the elytra. Very similar to the Lesser Spotted Tiger Beetle (C. octogramma), but it is distinguished by the longer distance between the rear spot and the end point of the elytra, and their ranges do not overlap.

Subspecies and morphological variants: No subspecies have been described.

Map 101 Safranek’s Tiger Beetle, Calochroa safraneki.

Distribution and habitats: Known only from a small area of western India (20 - 600 m) in Maharashtra. Along paths in partially deciduous but shady forest.

Behaviour: Unknown. Seasonality: June to July.

Black-headed Tiger Beetle, Calochroa fabriciana (Horn)

(Plate 13) [Map 102]

Identification: Length 12-18 mm; head and pronotum shiny black to bronze above and metallic green to blue on the sides; elytra dull velvety black with narrow green sides and three large orange oval spots along the length of each elytron (Plate 32.2). The similar Safranek'sTiger Beetle (C. safraneki) and Lesser-

spotted Tiger Beetle (C. octogramma) both have an additional light spot on the shoulder.

Subspecies and morphological variants: Individuals from the northern part of its range

have smaller, yellow spots on the elytra.

Distribution and habitats: Western Ghats of southern India. Partially shaded road cuts, secondary forest, open primary forest, and coffee plantations (100 - 900 m).

Behaviour: Unknown.

Seasonality: May to July. 100

Map

102 Black-headed Calochroa fabriciana.

Tiger

Beetle,

Four-spotted Tiger Beetle, Calochroa assamensis (Parry) (Plate 13) [Map 103] Identification: Length 18-20 mm; head and pronotum greenish-black above and purple

on the sides; elytra dull, dark greenish-black and purple with a large yellow shoulder spot and an even larger yellow spot at mid­ margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India,

Nepal,

Bhutan,

Bangladesh,

and

Myanmar. Floor of shaded forest with dense undergrowth and along sandy forest streams (150-1000 m) in

Behaviour: When disturbed they fly up

Map

103 Four-spotted Calochroa assamensis.

Tiger

Beetle,

from the forest floor to land on leaves of undergrowth vegetation or into dense vegetation on the ground.

Seasonality: April to October.

Bicoloured Tiger Beetle, Calochroa bicolor (Fabricius) (Plate 13) [Map 104] Identification: Length 15-17 mm; head and pronotum metallic green to copper; elytra dull black, dark indigo, or blue-green with or without three small spots arranged in a row along the length of the elytron. End of the abdomen under elytra reddishorange.

Subspecies and morphological variants: There are four subspecies described from India:

C. b. bicolor - metallic green head and pronotum; dull dark blue to green elytra

with no light markings. C. b. atavus - similar to C. b. bicolor but with three large yellow spots on elytra.

Map 104 Bicoloured Tiger Beetle, Calochroa bicolor; A, C. b. bicolor, B, C. b. atavus, C, C. b. haemorrhoidalis, D, C. b. xanthospilota.

C. b. haemorrhoidalis - copper head and

pronotum; elytra black with small yellow shoulder spot and two large yellow spots

along the elytron. 101

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

C. b. xanthospilota - similar to C. b. haemorrhoidialis, but spots on elytra much smaller.

Distribution and habitats: Usually floor of mature to secondary forests but also wooded fields, tall grass, cropland, and rice paddies (50 - 2400 m).

C b. bicolor - Northeastern India (Bihar, Assam, West Bengal, Meghalaya, Sikkim), Nepal, Bangladesh, Myanmar, Thailand, and China.

C. b, haemorrhoidialis - East central and southern India and Pakistan. C. b. xanthospilota - Southwestern India and Sri Lanka.

C. b. atavus - Northwestern India and Pakistan.

Behaviour: Attracted to lights at night.

Seasonality: May to July.

Orange-striped Tiger Beetle, Calochroa hamiltoniana (Thomson) (Plate 14) [Map 105] Identification: Length 14.5-17 mm; body slender; head and pronotum green or bronze; elytra green-bronze with narrow orange and black stripes the length of each elytron.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Forest floor of deeply shaded and moist, mature forest in Western Ghats (80 - 350 m).

Behaviour: Flies from forest floor to leaves

102

of undergrowth plants when disturbed.

Map 105 Orange-striped Tiger Beetle,

Seasonality: April to July.

Calochroa hamiltoniana.

Species Accounts

Temperate Tiger Beetles, genus Cicindela Although there are few external characters that reliably separate this large group of Temperate Tiger Beetles from other groups, Rivaller, found male genitalia provided relatively consistent distinguishing characters. The nearly eighty species in the genus are the most common tiger beetle species at higher latitudes in North America south to northern Mexico, in Eurasia and the Middle East, but several distinctive species occur as far south as India and South Africa. As presently defined, ten species of this Holarctic genus are found in India, and seven of them are endemic. Three subgenera are found in India (Cidndela, Pancallia, and Ancylia). All species in these groups have medium to large bodies, bulky heads with eyes less prominent, pronotum blocky and deeply incised with moderately rough to pebbled surfaces and few hairs. Their elytra are dark and boldly marked with broad yellow or white bands or spots.

Key XII: Temperate Tiger Beetles, genus Cicindela Identification Key to Cicindela Species la.

Front and top of head between eyes

Granulateci Tiger Beetle,

hairy with numerous, long, erect hairs in

Cidndela granulata

addition to two pairs of setae close to eyes

1b.

Front and top of head mostly bare except for

2

two pairs of single or multiple hairs close to each eye

2a. (1b.)

Elytra markings absent, reduced to a spot

Cobalt Tiger Beetle,

at elytra tips, or forming a wide longitudinal

Cidndela cyanea

stripe down the middle

2b.

Elytra markings forming separate or fused

3

bands, one or more band slanting and touching outer and/or inner margins at front

and rear outer angles

3. (2b.)

Elytra markings two or three transverse

4

bands fused into broad patterns touching both margins, head with two or more paired

groups of hairs near inside of each eye 3b.

Elytra markings one or more slanting bands

8

at centre touching only outer margins, head with only two pairs of single hairs near inside of each eye

103

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

4a. (3a.)

Cheeks on head below eyes with hairs

Shivah Tiger Beetle, Cicindela shivah

4b.

5a. (4b.)

Cheeks on head below eyes without hairs

Elytra

shoulder bands long, extending

5 6

obliquely backwards onto centre, elytra

markings either fused or almost touching 5b.

Elytra shoulder bands short, not extending

7

onto centre, other elytra markings not joining each other

6a. (5a.)

Elytra bands broad and fused at middle to

6b.

Elytra bands narrower and almost touching

Golden-banded Tiger

Beetle, Cicindela aurofasdata

at middle 7a. (5b.)

Lowland Tiger Beetle,

Cicindela goryi

form a cross

Elytra with middle band only on posterior

Yellow-shouldered Tiger

halves, but not touching sutural ridge, elytra

Beetle, Cicindela angulicollis

surface smooth with no punctures evident

7b.

Elytra middle band on both halves and

touching

sutural

ridge,

elytra

surface

Broad-striped Tiger Beetle, Cicindela princeps

shallowly punctured

8a. (3b.)

8b.

Pronotum longer than wide, head with

Ghats Tiger Beetle,

cheeks below eyes bare

Cicindela andrewesi

Pronotum wider than long, head with

9

cheeks below eyes on most specimens hairy but some specimens with bare cheeks 9a. (8b.)

Elytra middle band transverse and on most

Transverse-banded Tiger

specimens curved toward tip at centre,

Beetle,

pronotum at sides nearly covered by sparse

Cicindela calligramma

flattened and semi erect hairs arising out of

large punctures 9b.

Elytra middle band oblique or only an oval or

Dappled Tiger Beetle,

large circular spot fused with other maculae

Cicindela guttata

or separate, pronotum at sides mostly bare or with flattened hairs only on bottom half

arising out of minute punctures

104

Species Accounts

Granulated Tiger Beetle, Cicindela granulata Gebier (Plate 14) [Map 106] Identification: 15-18 mm; head and pronotum shiny black; elytra dull black to purple-black with highly variable yellow markings. Subspecies and morphological variants: Five subspecies of this widespread species have been described but only C g. stoliczkana enters India.

Distribution

and habitats: Extreme northern India in Kashmir, Pakistan, Kyrgyzstan, Kazakhstan, and China. Subalpine mountains on bare patches or sparsely vegetated areas.

Behaviour: Unknown.

Map 106 Granulated Tiger Beetle, Cicindela

Seasonality: July.

granulata.

Cobalt Tiger Beetle, Cicindela cyanea Fabricius (Plate 14) [Map 107] Identification: Length 19.5 - 23 mm; head and pronotum dull black on top and shiny blue or green on the sides; elytra dull blue or blackish green

usually without any

markings, but some specimens with variable markings ranging from a spot at elytra tips to a wide lengthwise orange stripe down

the side border of each elytron. Individuals without elytra markings could be confused with the Bicoloured Tiger Beetle (C. bicolor), which is much smaller.

Subspecies and morphological variants: Although individuals

the

have

striped been

and given

unstriped different

Map 107 Cobalt Tiger Beetle, Cicindela cyanea.

subspecies names, because they both occur

together, most experts consider them morphological variants and not subspecies.

Distribution and habitats: Northeastern India, Nepal, and Bangladesh. Habitat unrecorded.

Behaviour: Rarely observed and ecology/behaviour unrecorded. Seasonality: May to July. 105

K Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Yellow-shouldered Tiger Beetle, Cicindela angulicollis Horn (Plate 14) [Map 108] Identification: Length 19-22 mm; head and pronotum black; elytra dull black with three large yellow to white bands, front

and rear bands short, middle band longer; labrum with large yellow spot in middle. Intermediate between the Shivah Tiger beetle (C. shivah) and the Broad-striped Tiger beetle (C princeps). The Shivah Tiger Beetle has an all-black labrum, and the Broadstriped Tiger Beetle is shorter, more robust, and has a much larger front yellow marking on the elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to

Map 108 Yellow-shouldered Tiger Beetle, Cicindela angulicollis.

southern India. Floor of secondary forest, teak plantations, openings in mature forest, occasionally in tall grass adjacent to forest (above 500 m) in

Behaviour: Moderately strong flier that flies long distances when disturbed. Seasonality: June to October.

Shivah Tiger Beetle, Cicindela shivah Parry (Plate 14) [Map 109] Identification: Length 17-18 mm; long and narrow body form; dull black above

and below; cheeks with hair-like setae; pronotum covered with dense dark hair­ like setae; elytra with shiny purple sides and three bold yellow spots, anterior spot covers most of shoulder, middle spot widened at lateral margin but narrowed and broadly rounded at middle without touching suture, apical spot and middle spot either separate or narrowly joining laterally; labrum all black. Similar to the Yellow-shouldered Tiger

beetle (C. angulicollis) that has a yellow spot on the labrum.

106

Map 109 Shivah Tiger Beede, Cicindela shivah.

Species Accounts

Subspecies and morphological variants: No subspecies have been described for this rare species.

Distribution and habitats: Known only from Pune, Maharashtra and the North Kanara District of Karnataka in southern India. No habitats have been described for

this highly localized species.

Behaviour: Unknown.

Seasonality: July.

Broad-striped Tiger Beetle, Cicindela princeps Vigors (Plate 15) [Map 110] Identification: Length 18.5 - 20 mm; head and pronotum shiny black; elytra dull black with bold yellow front band, broad yellow band across entire middle, and narrow yellow margin on rear. Similar to the Golden-

banded Tiger Beetle (C. aurofasciata), but which has distinctly different markings on the elytra.

Subspecies and morphological variants: Two subspecies have been described from India and the status of a third form is uncertain:

C. p. princeps - dull elytral surface with non­ contrasting punctuation or no punctation on unbanded areas; yellow bands on elytra

Map

110 Broad-striped Tiger Beetle, Cicindela princeps-, A, C. p. princeps, B, C. p. ducalis, C, C. p. pochoni.

broad.

C. p. ducalis - shallow bluish green to coppery punctuation across entire elytra surface that is unbanded; yellow bands narrow. C. p. pochoni - yellow shoulder band and central band connected, but this character is not consistent.

Distribution and habitats: C. p. princeps - mountainsides (660 m) in tall grass and scrub forests in southern India.

C. p. ducalis - lowland forest paths and roads through forests in central and northeastern India. C. p. pochoni - known only from Kotagri in Nilgiri Hills of Tamil Nadu.

Behaviour: Flies short distances and is extremely sluggish on cool mornings. Seasonality: June to July.

107

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Golden-banded Tiger Beetle, Cicindela aurofasciata Dejean (Plate 15) [Map 111] Identification: Length 15-23 mm; head and pronotum shiny black or dark bronze; elytra black with bluish-green or copper

reflections and front yellow band points obliquely back to almost meet middle yellow band that crosses both elytra. Yellow bands not connected and narrower than in

Lowland Tiger Beetle (C. goryi).

Subspecies and morphological variants: Formerly two subspecies (C. a. aurofasciata and C. a. goryi) were recognized for India, but the distinctive anatomical, habitat, and behavioural differences between these two

populations are now considered sufficient to distinguish them as two separate species.

Map 111 Golden-banded Tiger Beede, Cicindela aurofasciata.

Distribution and habitats: Endemic to southern India. Higher elevation (1500 2100 m) in open grassy areas.

Behaviour: Flies only short distances of less than a meter when disturbed. Larvae construct angled turrets over the tunnel entrance. Seasonality: May to July.

Lowland Tiger Beetle, Cicindela goryi Chaudoir (Plate 15) [Map 112] Identification: Length 15-23 mm; head and pronotum shiny black or dark bronze; elytra black with bluish-green or copper

reflections, broad yellow bands across front

and middle fused to form an "X".

Subspecies and morphological variants: Formerly two subspecies (C. a. aurofasciata and C. a. goryi) were recognized for India,

but the distinctive anatomical, habitat, and behavioural differences between these two

populations are now considered sufficient to distinguish them as two separate species.

Distribution and habitats: Endemic to southern India. Lower elevation (600 -1500

m) on floor of secondary and primary forests

and brushy areas.

Map 112 Lowland Tiger Beetle, Cicindela goryi.

Species Accounts

Behaviour: Flies 2 to 5 m when disturbed. The larvae construct peculiar curved turrets above their tunnels.

Seasonality: May to July.

Dappled Tiger Beetle, Cicindela guttata Wiedemann (Plate 14) [Map 113] Identification: 13.5 - 16 mm; head and pronotum shiny dark green; elytra dull dark green or black-green with yellowish

markings - shoulder a long crescent, middle mark small and round, rear mark a small crescent along the curved edge of the rear

of the elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northern India. Scrubby brush areas, along

roads, and open area inside of forest (600 m).

Behaviour: Unknown.

Map 113 Dappled Tiger Beetle, Cicindela

Seasonality: June.

guttata.

Ghats Tiger Beetle, Cicindela andrewesi Horn (Plate 14) [Map 114] Identification: Length 13.5 -16 mm; slender body form; head and pronotum shiny black­ elytra dull black with three separate yellow-

orange oblong spots or connected to form two longitudinal curved lines running the length of each elytron; cheeks without hair­ like setae.

Subspecies and morphological variants: two subspecies have been described, but

some experts consider they should be treated as two separate species.

C. a. andre wesi - orange elytra markings in form of three oblong spots arranged in a row along length of each elytron.

Map 114 Ghats Tiger Beetle, Cicindela andrewesi; A, C. a. andrewesi, B, C. a. zingaroana.

109

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

C. a. zingaroana - orange elytra markings coalesced into two sinuate lines running the length of each elytron.

Distribution and habitats: C a. andrewsi - forested paths, coffee plantations and road cuts (600 to 900 m) of southern Western Ghats in India.

C. a. zingaroana - along shady paths and openings in partially deciduous forest of northern Western Ghats (600 m) in India.

Behaviour: Flies from disturbance into forest and hides among fallen leaves. Often active during heavy rainfall.

Seasonality: May to July.

Transverse-banded Tiger Beetle, Cicindela calligramma Schaum (Plate 15) [Map 115] Identification: Length 13.5-16 mm; head pronotum shiny copper, greenish copper, or green; elytra dull green or

and

brown with three separate broad or fused light yellow markings; if separate the front marking runs along the front edge of the elytra and shoulder to then turn back and

inward toward the central line of the elytron, the middle marking is a crescent running across the elytron's width with the open half facing back, the rear marking runs along the edge of the rounded rear end of each

elytron; cheeks with long, sparse, hair-like setae.

Subspecies and morphological variants:

Map 115 Transverse-banded Tiger Beetle, Cicindela calligramma.

There is considerable variability within populations in ground colour and markings

on the elytra, but the population in the Eastern Ghats tends to be dark green with

reduced elytra markings, and superficially resembles the Dappled Tiger Beetle (C. guttata), which has hairs on only the lower parts of the cheeks.

Distribution and habitats: Southern Western Ghats and Eastern Ghats (350 - 430 m) in India and Sri Lanka. Secondary forest floor and shrubby areas.

Behaviour: Strong fliers. Extreme variation in adult body size within each sex may be due to differences in amount of food received as larvae.

Seasonality: May to November.

110

Species Accounts

Margined Tiger Beetles, genus Calomera Another somewhat ambiguous and poorly defined genus, the 36 species of Margined Tiger Beetles occur over wide swaths of Africa, Europe and Asia. Because of this extremely wide geographical distribution across continents, different lineages are likely to be recognized with additional taxonomic studies. Most of these species were formerly grouped under the genus Lophyridia. Many share some characters of male genitalia and extensive areas of long, flattened, hairs (setae) densely covering various parts of the body. Typically the pattern of light lines and spots on the elytra include a continuous line along the outer edge (margin) of the elytra. Two of the nine species of this genus found in India are endemic.

Key XIII: Margined Tiger Beetles, genus Calomera Identification Key to Calomera Species 1a.

Head with area above upper lip (clypeus)

2

with hairs

1b.

Head with area above upper lip (clypeus)

6

bare of hairs 2a. (la.)

First segment of antenna with many erect sensory bristles

Brackish Tiger Beetle, Calomera littoralis conjunctaepustulata

2b.

First segment of antenna with only one

3

erect sensory bristles

3a. (2b.)

Front of head with cluster of hairs above

4

antennae

3b.

Front of head without cluster of hairs

5

above antennae 4a. (3a.)

Apical elytra band divided, no marking at

Waterfront Tiger Beetle,

outer shoulder on all specimens, front of

Calomera chloris

head with hairs near middle, female elytra expanded laterally near centre 4b.

Apical elytra band unbroken, dot at outer

Forest Stream Tiger Beetle,

shoulder missing on only a few specimens,

Calomera funerea

front of head bare near middle, female elytra not expanded laterally anywhere

5a. (3b.)

Elytra with band at shoulder

Beach Tiger Beetle, Calomera aulica

5b.

Elytra missing band at shoulder

Anamalai Tiger Beetle,

Calomera quadrlpunctulata 111

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

6a. (1b.)

Elytra band at shoulder unbroken although

7

it may be short, long or connected to lateral

expansion of middle band

6b.

Elytra band at shoulder divided, each portion

Riverine Tiger Beetle,

widely separated and not connected to

Calomera fowleri

lateral expansion of middle band 7a. (6a.)

Elytra

middle

band

with

transverse

8

portion concave along its front margin, terminal portion broadly connected

7b.

Elytra

middle

band

with

transverse

portion not concave along its front

Riverbank Tiger Beetle,

Calomera cardoni

margin, terminal portion separated or

only narrowly connected 8a. (7a.)

Lateral margins of female elytra expanded

Sand Tiger Beetle,

outward; fourth male antennal segment

Calomera angulata

without a long comb of stiff bristles

8b.

Lateral margins of female elytra uniformly

Combed Tiger Beetle,

parallel; fourth male antennal segment

Calomera plumigera

with a long comb of stiff bristles

Brackish Tiger Beetle, Calomera littoralis (Fabricius) (Plate 15) [Map 116] Identification: Length 10-15 mm; head and pronotum dark brown, black or greenish brown; elytra dark brown with ivory-coloured markings that are variously connected or separated into as many as six large spots.

Subspecies and morphological variants: Ten subspecies of this widespread species

have been described. Only the subspecies

C. I. conjunctaepustulata is present as an isolated population in India.

Distribution northwestern

and India,

habitats: Iran,

Extreme Azerbaijan,

Georgia, Ukraine, Russia, Afghanistan, Kyrgyzstan, Pakistan (Baluchistan, Sind), NE Tibet, Tadzhikistan, Uzbekistan, Kazakhstan,

Map 116 Brackish Tiger Beetle, Calomera littoralis.

Mongolia, and China. Brackish swamps and saline lake edges from the coast to mountain plateaus.

Behaviour: Unknown. Seasonality: April to July. 112

Species Accounts

Beach Tiger Beetle, Calomera aulica (Dejean) (Plate 15) [Map 117] Identification: Length 10-12 mm; head and pronotum brown, green, dark copper-

green or nearly black; elytra dark brown with

copper reflections and bold ivory-coloured markings that include a "C"-shaped shoulder mark, "L"-shaped middle mark extending

from each mid-margin toward the mid-line, one or two small dots, and a "J"-shaped mark that follows the. curving rear edge of the elytra.

Subspecies and morphological variants: Five subspecies have been described for this wide-spread species. Only the nominate Map 117 Beach Tiger Beetle, Calomera aulica.

form occurs in India.

Distribution

and

habitats:

Extreme

northwestern India west to Greece and throughout Africa. Coastal beaches and estuaries.

Behaviour: Unknown.

Seasonality: July to October.

Sand Tiger Beetle, Calomera angulata (Fabricius) (Plate 15) [Map 118] Identification: Length

10.5

-

14

mm;

female significantly expanded at mid-body; bronze to dark copper, occasionally green or nearly black; male

slender

and

complete ivory-coloured markings on elytra

with "C"-shaped shoulder mark connected along margin to right-angled middle mark, and rear mark running along margin of

the pointed end of elytra with a club-like extension toward the middle of the elytron; mandibles relatively long. Similar to the Riverine Tiger Beetle (C. fowleri), but its thin and disrupted elytra pattern of markings, shorter mandibles, and its habitat of forest river edges distinguish it. The similar

Map 118 Sand Tiger Beetle, Calomera angulata.

Combed Tiger Beetle (C. plumigera) is larger with bolder markings on the elytra and 113

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

the male has a comb of bristles on the fourth segment of the antennae. Many earlier

authors referred to the Sand Tiger Beetle as C sumatrensis, but this name is no longer valid.

Subspecies and morphological variants: Two subspecies have been described, one in Japan. The nominate subspecies occurs throughout the Indian subcontinent. A blue-green morph occasionally occurs within populations of northeastern India.

Distribution and habitats: Throughout India, Nepal and Sri Lanka west to Afghanistan and east to Philippines and Taiwan. Sandy river banks, inner ocean bay beaches, and edges of lakes and reservoirs (0 - 2500 m).

Behaviour: Often occurs on the same sandy banks together with the Riverbank Beetle (C. cardoni) and the Combed Tiger Beetle (C. plumigera). In pre and post monsoon it forms nocturnal and, during mid monsoon, diurnal roosts of up to 5000 individuals on a single riverside bush together with the Combed Tiger Beetle (C. plumigera) and the Waterfront Tiger Beetle (C. chloris) (Plate 32.3).

Seasonality: April to November.

Riverine Tiger Beetle, Calomera fowleri (Heynes-Wood and Dover) (Plate 16) [Map 119] Identification: Length 10.5-12 mm; male body slender and female considerably stouter; head and pronotum dark broyvn to bronze with copper-green reflections;

elytra brown to copper with a disrupted pattern of thin markings often separated into spots and short lines; mandible relatively short. Similar to the Sand Tiger

Beetle (C. angulata), but its bold and complete elytra pattern of markings, longer mandibles, and its habitat of open sandy

beaches distinguish it.

Subspecies and morphological variants: No subspecies have been described.

Map 119 Riverine Tiger Beetle, Calomera

Distribution and habitats: Known only fowleri. from peninsular India. Moist stream edges in forests and occasionally adjacent grasslands (50 - 1000 m).

Behaviour: Forages in wet sand and gravel areas. Seasonality: May to November.

114

Species Accounts

Combed Tiger Beetle, Calomera plumigera (Horn) (Plate 16) [Map 120] Identification: Length 12.5-16 mm; head and pronotum bronze with copper-green reflections; elytra dark brown to olive green or copper with standard elytral pattern of

light yellowish markings, but with shoulder, middle and rear markings usually connected along the margin; middle band sinuate; male with large comb-shaped set of stiff bristles.

Similar to Sand Tiger Beetle (C. angulata), which is smaller, has thinner markings on the elytra, middle elytral band right-angled, and the male lacks the comb of bristles on

the antennae.

Subspecies and morphological variants: Five subspecies have been described for this wide-spread species, but only two occur in India:

Map 120 Combed Tiger Beetle, Calomera plumigera; A, C. p. plumigera, B, C. p. macrograptina.

C. p. plumigera - smaller, body colour brighter copper, elytra with thinner markings on elytra. C. p. macrograptina - larger, body colour dark brown to olive-green, elytra with

bolder markings.

Distribution and habitats: along wet sandy and gravelly banks of large rivers (300 -2100 m).

C. p. plumigera - known only from southern India. C. p. macrograptina - northern India, Pakistan, Nepal, Bangladesh, and Myanmar.

Behaviour: Often occurs on the same sandy banks together with the Sand Tiger Beetle (C. angulata) and the Riverbank Tiger Beetle (C. cardoni). Attracted to lights at night. In pre- and post-monsoon forms nocturnal roosts and during mid monsoon

forms diurnal roosts of up to 5000 individuals on a single riverside bush together with the Sand Tiger Beetle (C. angulata) and the Waterfront Tiger Beetle (C. chloris)

(Plate 32.3).

Seasonality: May to September.

Riverbank Tiger Beetle, Calomera cardoni (Fleutiaux) (Plate 16) [Map 121] Identification: Length 12-13.5 mm; brown to blackish brown; standard elytral pattern, but front marking restricted to lower shoulder, middle marking usually separated into two parts or connected by fine line. 115

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Subspecies and morphological variants: No subspecies have been described, but individuals from northwestern India are often uniformly blackish-brown with elytra

pattern of light markings greatly reduced

and separated.

Distribution and habitats: Widespread throughout India and Sri Lanka. Wet sandy banks of small to large rivers (350 - 700 m).

Behaviour: Often occurs on the same sandy banks together with the Sand Tiger Beetle (C. angulata) and the Combed Tiger Beetle (C. plumigera). Map 121 Riverbank Tiger Beetle, Calomera

Seasonality: May to November.

cardoni.

Waterfront Tiger Beetle, Calomera chloris (Hope) (Plate 16) [Map 122] Identification: Length

11.5 - 12 mm; bright green, copper green, black green

or blue; elytra with three bold white spots

along margin of rear half and surface distinctly

granulate

with

punctures.

Similar to the Forest Stream Tiger Beetle (C. funerea) but elytra surface different, and

white spots along rear margin wider and often connected in Waterfront Tiger Beetle.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern and northeastern India, Afghanistan, Pakistan, Nepal, Bhutan, Laos, and possibly Myanmar. Map 122 Waterfront Tiger Beetle, Calomera chloris. Pebbly mountain streams (120 - 1800 m) and wet sandy and gravel areas along open rivers and occasionally ponds away from

forests (below 300 m).

Behaviour: In pre- and post-monsoon forms nocturnal and, during mid monsoon, diurnal roosts of up to 5000 individuals on a single riverside bush together with the Combed Tiger Beetle (C. plumigera) and the Sand Tiger Beetle (C. angulata) (Plate

32.3).

Seasonality: February to November. 116

Species Accounts

Forest Stream Tiger Beetle, Calomera funerea (Madeay)

(Plate 16) [Map 123]

Identification: Length 12-14 mm; dull greenish black, green bronze or black; elytra

with four or three thin white spots along the margin and surface granulate with patches not punctate. Green forms similar to the Waterfront Tiger Beetle (C. ch loris) but elytra I

surface different.

Subspecies and morphological variants: Four subspecies have been described, of which two occur in India: C. f. funerea - copper-brown with four spots

along the elytra I margin including one at the shoulder.

C. f assimilis - greenish-black to green with three spots along the elytral margin.

Map 123 Forest Stream Tiger Beetle, Calomera funerea; A, C. f. funerea, B, C. f assimilis.

Distribution and habitats: Sandy banks of forested streams and occasionally lakes (150 - 2000 m): C. f. funerea - northern India, Myanmar,Thailand, Malaysia, Laos, Cambodia, Vietnam,

Indonesia, and China. C. f. assimilis - central India, Pakistan, Nepal, Bhutan, Bangladesh, Myanmar,Thailand,

Laos, and China.

Behaviour: Apparently replaced along forested river banks by the Riverine Tiger Beetle (C. fowleri) in the Western Ghats and Deccan Plateau.

Seasonality: April to November.

AnamalaiTiger Beetle, Calomera quadripunctulata (Mandi) (Plate 16) [Map 124] Identification: Length 15-16 mm; top of head and pronotum copper, sides violet­ green; elytra bronze or bronze green with two white-spots along back half on each margin.

Subspecies and morphological variants: No subspecies have been described for this highly localized species.

Distribution and habitats: Known only from South India in the Anamalai Hills of western Tamil Nadu (1075 m).

Behaviour: Its localized habitat may make this species vulnerable to development and dam building.

Map 124 Anamalai Tiger Beetle, Calomera

Seasonality: April to November.

quadripunctulata.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Ornamented Tiger Beetles, genus Cosmodela These relatively large species present a striking external appearance. They possess brilliant multi-coloured bodies with iridescence on the head, in furrows on the pronotum, and around the edges of their elytra marked with bold spots. Their finely ridged heads and pronotum are delicately sculptured and devoid of hairs, creating an almost polished appearance.The 12 species in this genus are restricted to southern and southeastern Asia. None of the seven species reported from India is endemic.

Key XIV: Ornamented Tiger Beetles, genus Cosmodela Identification Key to Cosmodela Species la.

Elytra marked with four or five spots,

2

central spot either separate or narrowly

connected to a marginal spot, shoulder spot large, one or more spots narrowly

connected along lateral margins on

some specimens

1b.

Elytra marked with three spots always

3

separate and all aligned centrally with at most a small shoulder spot

2a. (la.)

Elytra central spot enlarged and isolated

Twin-spotted Tiger Beetle,

on most specimens, rarely connected

Cosmodela didyma

to a marginal spot, elytra posterior end broadly tapered to an elongated point

with a distinctively long spine at its tip 2b.

Elytra central and marginal spots enlarged

Goblet-marked Tiger Beetle,

and always narrowly connected, elytra

Cosmodela diehli

posterior end

broadly rounded, not

elongated nor ending in a distinctively long spine at its tip 3a. (1b.)

Head at sides below eyes with distinct

4

flattened hairs over most of its surface

3b.

Head at sides below eyes appearing bare,

6

but actually with very fine, indistinct hairs 4a. (3a.)

4b.

Pronotum with hairs along lateral edges

Pronotum without lateral hairs

5

Iridescent Tiger Beetle, Cosmodela duponti

118

Species Accounts

5a. (4a.)

Head and pronotum with metallic copper

Intermediate Tiger Beetle,

reflections on top; elytra middle band

Cosmodela intermedia

comprised of two spots, separate or

narrowly joined to each other

5b.

Head and pronotum with greenish purple

White-spotted Tiger Beetle,

reflections dorsally; elytral middle band

Cosmodela fleutiauxi

comprised of one spot with a short,

oblique projection posteriorly

6a. (3b.)

Male

a

labrum

darkened

abruptly

raised

ridge down

along

its middle

Arched Tiger Beetle, Cosmodela virgula

contrasting with paler lateral areas, female

thorax below elytra shoulder broadly concave on most specimens, shallowly

grooved only at middle on others

6b.

Male labrum lacking an abrupt darkened

Coppery-green Tiger Beetle,

ridge down its middle, no more than

Cosmodela juxtata

darkened margins contrasting with paler central area, female thorax below elytra

shoulder forming a small shallow cavity

above middle

Intermediate Tiger Beetle, Cosmodela intermedia (Chaudoir) (Plate 17) [Map 125] Identification: Length 14-15 mm; head and pronotum iridescent copper on top and blue-green on the sides; elytra olive­ green to purple-green with five separate white spots or sometimes the middle two spots are joined together by a thin line. The C /. chitwanae subspecies superficially resembles the Arched Tiger Beetle (C. virgula\, but the latter is darker with bolder markings on the elytra including a complete connection between the two middle dots.

Subspecies and morphological variants: Two subspecies have been described: C /. intermedia - olive-green elytra with small dots for markings (northwestern India).

Map

125 Intermediate Cosmodela intermedia.

Tiger

Beetle,

C. /. chitwanae - dark purple-green elytra and thicker white markings in which the

middle ones are joined together to form a broad band.

119

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Distribution and habitats: Rocky shores and large exposed boulders of mountain streams and adjacent forests during floods (300 - 3600 m).

C. /. intermedia - Northwestern India, Afghanistan, Pakistan, and Nepal. C /. chitwanae - Nepal (Chitwan National Park and likely adjacent northeastern India).

Behaviour: Commonly occurs in the same gravel and rocky mountain stream habitat as the Stream Tiger Beetle (Cylindera subtilesignata), the Mountain Stream Tiger Beetle (Cylindera albopunctata), and occasionally with the Arched Tiger Beetle (C. virgula), but the latter is usually found in more shaded areas.

Seasonality: May to August.

White-spotted Tiger Beetle, Cosmodela fleutiauxi (Horn) (Plate 17) [Map 126] Identification: Length 13 - 15 mm; head and pronotum greenish-purple; elytra dull dark blue, black with green and purple

iridescence on the sides and along the

middle line separating the elytra, four white spots and middle one with backward projection portion.

Subspecies and morphological variants: One other subspecies has been described from Myanmar to China.

Distribution and habitats: Northeastern India, Bhutan, Nepal, and Bangladesh. Moist forest river and stream sides; adjacent agricultural fields and clay soil when streams flood (180-1500 m).

Map 126 White-spotted Tiger Beetle, Cosmodela fleutiauxi.

Behaviour: Flies to undergrowth leaves from forest floor when disturbed.

Seasonality: May to October.

Iridescent Tiger Beetle, Cosmodela duponti (Dejean) (Plate 16) [Map 127] Identification: Length 15-18 mm; head and pronotum shiny blue-green oi copper; elytra shiny blue-green or copper around edges and across middle dividing dark area into two parts, dull blackish purple in the centre with white markings consisting of three small spots and middle one long and narrow acrosi

the elytron (Plate 31.1).

120

Species Accounts

Subspecies and morphological variants: Considerable individual variation in pronotum

and elytra colours within populations, but only one additional subspecies has been described from Thailand.

Distribution and habitats: Peninsular and northeastern India, Bangladesh, Myanmar, Thailand, Laos, Malaysia, Vietnam, and China.

Near small forest streams or in stream beds on large rocks and boulders (300 - 1150 m).

Behaviour: Often flies from forest floor to leaves of undergrowth bushes to escape danger.

Seasonality: May to October.

Map 127 Iridescent Tiger Beetle, Cosmodela duponti.

Twin-spotted Tiger Beetle, Cosmodela didyma (Dejean) (Plate 17) Identification: Length 13-16 mm; head dark greenish to black olive; pronotum blue-green to brassy reddish black; elytra dull black with shoulders and all margins usually bright blue-green, marked with five yellow spots usually separate, but narrowlyjoined along lateral margin on some specimens, shoulder spot divided in two, marginal and central spots and apical spots evident; labrum bright, outlined

in black, occasionally individuals lack green, red or other coloured reflections and exhibit only a pure mat black finish dorsally. Closely resembles the Gobletmarked Tiger Beetle (C. diehli) except for its more elongated elytra with a spine at its tip and two central spots which are separated on most specimens. Other less superficial differences exist in male genitalia shape and extent to which the groove on the side of the female thorax is depressed to receive the male

mandibles in copulation.

Subspecies and morphological variants: No subspecies have been described. Some specimens are known from Indonesia with joined shoulder spots and a narrow marginal connection between the middle and apical spots.

Distribution and habitats: Regular on nearby Indonesia, but because published records of this species from Nicobar Islands may be the result of misidentification, we consider it HYPOTHETICAL for India.

Behaviour: Unknown.

Seasonality: Unknown.

121

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Goblet-marked Tiger Beetle, Cosmodela diehli Wiesner (Plate 17) [Map 129] Identification: Length 13-15 mm; head greenish-red, finely ridged, without hairs at side; labrum with six bristles, yellow, laterally

black-lined, the black border occasionally widened towards the base in the middle;

pronotum wider than long, centrally dull granulated, glossy, bare, greenish coppery with distinct blue violet in transverse and middle furrows; elytra dull black, laterally

with narrow green-copper margins, inner

edge and posterior ends fringed with bright green, tips not elongated towards their ends and with short sutural spine; elytra markings consist of five spots, a shoulder spot divided in two, marginal and middle spots large

Map 129 Goblet-marked Tiger Bee Cosmodela diehli.

and usually narrowly joined, and a large

oval apical spot. Twin-spotted Tiger Beetle (C didyma) looks similar but separat externally by the shape of the elytra and the extent the middle elytra markir are joined. Their reproductive isolation is assured based on male internal geniti characters and differences in the form of the groove on the side of the female thoi that accepts the male mandibles while mating.

Subspecies and morphological variants: No subspecies have been described.! variation noted in elytra markings for the few specimens known from Indonesia a India is limited to the degree to which the marginal and middle spots are connect!

Distribution and habitats: India (Great Nicobar Island) and Indonesia (Sima Island along west coast of Sumatra).

Behaviour: Unknown. Seasonality: July on Great Nicobar Island, India; February on Simalur Island, Sumal Indonesia.

Coppery-green Tiger Beetle, Cosmodela juxtata (Acciavatti and Pearson) (Plate 17) [Map 130] Identification: Length 15-18 mm; head and pronotum metallic copper edged w green; elytra greenish black with small yellow shoulder dot and three large yelk circles in a row along the length. Very similar to Arched Tiger Beetle (C. virgula) t differs in male genitalia and form of the groove on the side of the female thorax tl accepts the male mandibles in copulation; also, the middle mark on the elytra oft

122

Species Accounts

Arched Tiger Beetle tends to be crescent­ shaped rather than more oval or round as in the Coppery-green Tiger Beetle.

Subspecies and morphological variants: Formerly considered a subspecies of C. aurulenta. Individuals from Jammu and Kashmir are considerably larger (22 mm).

Distribution and habitats: Northern and northeastern India, Myanmar,Thailand, Laos, Cambodia, Vietnam, and China. Forested stream sides (300 - 500 m).

Behaviour: Near streams lakes, and rivers on sand in among large boulders and rocks; during floods often moves into adjacent

Map 130 Coppery-green Tiger Beetle, Cosmodela juxtata.

forests and fields.

Seasonality: May to November.

Arched Tiger Beetle, Cosmodela virgula (Fleutiaux) (Plate 17) [Map 131] Identification: Length 15-18 mm; very similar to the Coppery-green Tiger Beetle (C. juxtata) but differs in male genitalia and form of the groove on the side of the female thorax that accepts the male mandibles in copulation; also, the middle mark on the elytra of the Arched Tiger Beetle tends to be crescent-shaped rather than more oval or round as in the Copperygreen Tiger Beetle.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and

habitats:

Northern

and northeastern India, Nepal, Bhutan, Bangladesh, Myanmar, Thailand, Cambodia,

Map 131 Arched Tiger Beetle, Cosmodela virgula.

Laos, Vietnam, and China. Along moist forested rivers and streams; moves to adjacent agricultural fields, sandy and clay open areas during floods; also narrow

shaded paths in forest (150 - 2000 m).

Behaviour: Often flies to land on leaves of low bushes when disturbed.

Seasonality: May to October.

123

h Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Small-eyed Tiger Beetles, genus Plutacia This genus is endemic to India and contains only two rare species. Their bodies are medium to large and robust with eyes that do not distinctly protrude outward from the head. They also share broad markings that almost reach the side margins on the elytra.

Key XV: Small-eyed Tiger Beetles, genus Plutacia Identification Key to Plutacia Species 1a.

Elytra

marked

with

three

obliquely

transverse bands, thorax evenly rounded

Opulent Tiger Beetle,

Plutacia dives

along lateral edge, male antennae without

comb of stiff bristles 1b.

Elytra markings a small humeral spot and a

Ridged Tiger Beetle,

large, nearly trapezoidal spot at middle, thorax

Plutacia notopleuralis

with narrowly raised, reflexed and striated

ridge lateral edge, male antennae with comb of stiff bristles

Opulent Tiger Beetle, Plutacia dives (Gory) (Plate 17) [Map 132] Identification: Length 15-17 mm; head and pronotum shiny green and copper red; elytra velvety green with metallic copper edges and three wide yellow marks edged

with black, the shoulder mark is a band that runs from the shoulder at an angle back

toward the middle. Th% middle mark is a

band that runs almost completely straight across the two elytra, and the rear mark

is "C"-shaped following the outline of the elytral margin at the back.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: This very rare

Map 132 Opulent Tiger Beetle, Plutacia dives.

species is endemic to India and has been collected from widely separated sites in northeastern, central and southern India, but its habitat is unrecorded.

Behaviour: Unknown. Seasonality: Unknown. 124

Species Accounts

Ridged Tiger Beetle, Plutacia notopleuralis (Acciavatti and Cassola) (Plate 17) [Map 133] Identification: Length 20.5 mm; black with sides of head green and purple and sides

of pronotum shiny green with a unique narrow ridge along its sides; elytra dull black with two yellow orange markings, shoulder spot small, middle spot large and forming

a trapezoid. Males have a small comb of bristles on each antenna.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Extremely rare species known only from a single male

specimen collected in Odisha in eastern India.

Behaviour: Unknown.

Map 133 Ridged Tiger Beetle, Plutacia notopleuralis.

Seasonality: Unknown.

Reticulated Tiger Beetles, genus Lophyra. This genus is comprised of 81 species distributed from Africa to the Philippines. They all share similar characteristics of the male genitalia. Most are small to medium-sized and have intricate patterns of broadly reticulated pale markings across the elytra.The genus has been divided into eight subgenera.Two subgenera with 11 species are represented in India, five in the subgenus Lophyra, and six in the subgenus Spilodia. Each subgenus includes a single species endemic to India.

Key XVI: Reticulated Tiger Beetles, genus Lophyra Identification Key to Lophyra Species la.

Elytra with expanded markings broadly

2

connected along either lateral or inner

margins or both margins, head with patches of hairs behind or in front of

eyes, thorax at sides densely covered by flattened white hairs 1b.

Elytra with markings not expanded nor

6

connected along lateral margins and spots

widely separated near inner margins, head

without patches of hairs near eyes, thorax at sides only sparsely hairy 125

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

2a. (la.)

Elytra lunules entirely obscured by expansive lighter surface leaving only

Waxen Tiger Beetle,

Lophyra cerina

four small darker coloured areas at centre 2b.

Elytra lunules distinct either connected

3

or separate at lateral margins, darker

coloured area extends over entire surface 3a. (2b.)

Elytra lunules all connected or narrowly

4

separated along lateral margins, head with wide hairs clustered above antennal

/

insertion on head

3b.

Elytra lunules all widely separated at

Bald Tiger Beetle,

lateral margins, head without cluster of

Lophyra striatifrons

wide hairs above antennal insertion 4a. (3a.)

Elytra along front half of sutures with

all

spots

connected,

dark

surface

Harlequin Tiger Beetle,

Lophyra histrio

pigmentation joins suture at two places

only in rear half 4b.

Elytra along front half of sutures with two

5

separate spots, dark surface pigmentation joins suture at three places along entire length 5a. (4a.)

Elytra markings broad and varied in

Lattice Tiger Beetle,

width, ivory and outlined in blue or black,

Lophyra cancellata

surface colour varied from dull copper to

green to blue-green, lunules usually all connected laterally with expanded ends and middle band elbowed, head with

bare cheeks

5b.

Elytra

markings

wide,

yellow-white,

surface colour dull copper, lunules at

Braided Tiger Beetle,

Lophyra catena

least partially connected laterally without

expanded ends and middle band only slightly bent, head with hairy cheeks

6a. (1b.)

Elytra lunule at shoulder long, completely

7

formed and reaching almost middle band, remaining markings bandsoreiongated spots 6b.

Elytra lunule at shoulder short, complete or divided, but not reaching middle band,

remaining markings oval or small spots

126

10

Species Accounts

7a. (6a.)

Elytra lunule at shoulder wide, on most

Belted Tiger Beetle,

specimens extending to touch middle

Lophyrastriolata

spot, on other specimens fused with

irregularly shaped middle spot located

more

centrally

inward

from

lateral

margin 7b.

Elytra lunule at shoulder narrow, never

8

fused with a more regularly shaped middle spot extending from lateral to

sutural margins 8a. (7b.)

Head and prothorax brilliant copper-red,

Tangled Tiger Beetle,

elytra velvety black with white markings,

Lophyra lefroyi

middle band short without bulges at apical end 8b.

Head

and

pronotum

dark

copper,

9

elytra dull velvety green with whitish

yellow markings, middle band long and descending toward suture in rear half

with or without a bulge at apical end 9a. (8b.)

Elytra

lunule

at

shoulder

long,

terminating almost at middle band,

Plaited Tiger Beetle, Lophyra vittigera

middle band of uniform width at its

apical end 9b.

Elytra lunules at shoulder and middle

Many-spotted Tiger Beetle,

band completely formed, middle band

Lophyra multiguttata

very narrow at middle and enlarged at both ends

10a. (9b.)

Elytra lunule at shoulder unbroken and

Lined Tiger Beetle,

very narrow, but other markings only

Lophyra lineifrons

as extremities, cheeks bare on most specimens, but sparsely hairy on a few

others, fourth antennal segments on males without bristles 10b.

Elytra with all markings fragmented

Tiny-spotted Tiger Beetle,

into their extremities, cheeks bare on all

Lophyraparvimaculata

specimens, fourth antennal segment on

males with bundle of stiff bristles

127

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Braided Tiger Beetle, Lophyra catena (Fabricius)(Plate 18) [Map 134] Identification: Length 10-12 mm; head and pronotum copper on top and green or

blue on the sides; pronotum covered with

thick, white hairs; elytra dull copper with intricate pattern of thick yellowish-white lines that obscure most of the background

colour, light markings along inner edges of elytra broken up into two or occasionally four short spots in front third; cheeks covered with white hairs. Similar to the Lattice Tiger Beetle (L cancellata) and the

Map 134 Braided Tiger Beede, Lophyra catena.

Harlequin Tiger Beetle (L histrio), but both of them have bare cheeks.

Subspecies and morphological variants: Only one other subspecies has been described, and it is restricted to Sri Lanka.

Distribution and habitats: Throughout much of India, Pakistan, Myanmar, and Sri Lanka. Upland grasslands, old fields, dry white sandy upper beaches of rivers and

lagoons near the ocean and occasionally in large sunny forest clearings (0 - 245 m).

Behaviour: Unknown. Seasonality: April to November.

Waxen Tiger Beetle, Lophyra cerina Naviaux and Acciavatti (Plate 18) [Map 135] Identification: Length 9 - 10.5 mm; head and pronotum bright copper-red; elytra yellowish-orange with waxy lustre and five small blue spots running along the length near the midline; midline between the two elytra also blue.

Subspecies and morphological variants: No subspecies have been described for this extremely localized species.

and habitats: Presently known only from the southwest side of the Map 135 Waxen Tiger Beede, Lophyra cerina. Gulf of Mannar in extreme southern India (Turinelveli District, Tamil Nadu). Restricted to fine, yellow cream-coloured, dry sand well away from the ocean's edge. Distribution

Behaviour: Often occurs adjacent to the Braided Tiger Beetle (L catena), but does not enter onto its white sand habitat.

128 Seasonality: October.

Species Accounts

Bald Tiger Beetle, Lophyra striatrfrons (Chaudoir) (Plate 18) [Map 136] Identification: Length 9 -10.5 mm; head and pronotum copper; elytra contrastingly blackish-green with three standard whitish markings supplemented by oval spots and stripes near the middle-front part of the elytra. Cheeks with white hairs. Similar to

the Braided Tiger Beetle (L catena) and the Lattice Tiger Beetle (L cancellata) but much narrower body shape.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Widely distributed in India and Nepal. Open sandy areas among • tall grasses and muddy areas near grass

* Map 136 Bald Tiger Beetle, Lophyra striatifrons.

margins along forested rivers (500 - 1000 m).

Behaviour: Very wary and difficult to approach. Spends night in muddy grassy areas near forest edge.

Seasonality: June to August.

Harlequin Tiger Beetle, Lophyra histrio (Tschitscherine) (Plate 18) [Map 137] Identification: Length 10.5 - 12.5 mm; Head and pronotum copper; elytra dull, coppergreen with standard ivory-coloured markings

plus "L"-shaped markings at the front, light

markings along inner edges of elytra extend down the first half; cheeks bare. Similar to the

Lattice Tiger Beetle (L cancellata), which is limited to south and northeastern India, and the Braided Tiger Beetle (L catena), which has white hairs on its cheeks.

Subspecies and morphological variants: No subspecies have been described

Distribution and habitats: Northwestern India, Iran, Pakistan, Afghanistan, Saudi

Map 137 Harlequin Tiger Beetle, Lophyra histrio.

Arabia, United Arab Emirates, and Oman.

Sandy areas along rivers, irrigation canals, and flooded fields.

Behaviour: Unknown.

Seasonality: April to October. 129

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Lattice Tiger Beetle, Lophyra cancellata (Dejean) (Plate 18) [Map 138] Identification: Length 10-11 mm; head and pronotum copper to copper-green; elytra dull, dark copper-green, green or blue

with standard ivory-coloured markings, light spots along inner edges of elytra broken up into two or four short lines; cheeks bare. Similar to the Braided Tiger Beetle (L catena), but it has white hairs on the cheeks; the similar Harlequin Tiger Beetle (L histrio) is restricted to the deserts of northwestern

India.

Subspecies and morphological variants: Five subspecies have been described, but within India only two subspecies have been described. However, even within these

Map 138 Lattice Tiger Beetle, Lophyra cancellata-, A, L. c. cancellata, B, L. c. intemperata.

subspecies, colour and elytral pattern are

highly variable and intergrade forms predominate across central India:

L. c. cancellata - copper head and pronotum and dull dark copper-green elytra with broad markings predominate. L. c. intemperata - copper green to blue head pronotum, and elytra with narrow

markings predominate.

Distribution and habitats: Dry upper sandy beaches along medium-sized to large rivers (50-1400 m): L c. cancellata - southern India and Sri Lanka. L c. intemperata - northern India, Bangladesh, Myanmar, Thailand, Malaysia,

Cambodia, Laos, Vietnam, and China.

Behaviour: Unknown.

Seasonality: April to November.

Belted Tiger Beetle, Lophyra striolata (liliger) (Plate 18) [Map 139] Identification: Length 10-15 mm; head and pronotum copper above and right copper-green, blue-green and purple on the sides; elytra velvety black or greenish black with white or yellow markings forming variable pattern of separated longitudinal stripes and bands that do not reach the outer edge. Hairs on the cheeks separate it from the similar Tiny-spotted Tiger Beetle (L parvimaculata).

130

Species Accounts

Subspecies and morphological variants: Nine subspecies have been described but only the nominate form occurs in India. Within

India

the

southern

populations

tend to have smaller and more separated

markings on the elytra.

and habitats: Widely distributed across India, Nepal, Bangladesh, Myanmar, Thailand, Malaysia, Laos, Cambodia, Vietnam, China, Indonesia, and Philippines. Secondary forest floor and Distribution

(learings in mature forest but moves to road cuts and shrubby fields during the height of the monsoons (120-1600 m).

Behaviour: Unlike most other forest species

Map 139 Belted Tiger Beetle, Lophyra striolata.

when disturbed, this one does not fly up to land on leaves of undergrowth bushes but lands again on the ground or muddy banks. Occurs together with the Many-spotted liger Beetle (L multiguttata) and the Lesser Spotted Tiger Beetle (C. octogramma).

Seasonality: April to October.

Ml Tiger Beetle, Lophyra lineifrons (Chaudoir) te19) [Map 140] Identification: Length 10-13 mm; head and pronotum bright copper; elytra velvety black with white, narrow markings at the

shoulder and small spots at the middle •nd rear. Similar to southern populations

of the Belted Tiger Beetle (L striolata), but leparated by range. Hairs on the cheeks

leparate it from the similar Tiny-spotted Tiger Beetle (L parvimaculata).

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern India, Nepal, Bangladesh, Myanmar, Thailand, Malaysia, Cambodia, Laos, Vietnam, and China. Paths and forest floor in mature forest (200 - 750 m).

Map 140 Lined Tiger Beetle, Lophyra lineifrons.

Behaviour: Unknown. Seasonality: May to July. 131

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Tangled Tiger Beetle, Lophyra lefroyi (Horn) (Plate 19) [Map 141] Identification: Length 13.5-16 mm; head and pronotum bright copper-red above and shiny blue-green on the sides; contrasting elytra velvety black with fourteen white lines

and spots; cheeks without hairs.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northern India. Open areas in flood-plain high grasses (500 - 800 m).

Behaviour: Unknown.

Seasonality: July. Map 141 Tangled Tiger Beetle, Lophyra lefroyi.

Tiny-spotted Tiger Beetle, Lophyra parvimaculata (Fowler) (Plate 19) [Map 142] Identification: Length 14-15 rhm; head and pronotum copper-brown above and shiny blue green on the sides; elytra velvety black or greenish-black with white or yellow

markings reduced to circles and elongated spots. Cheeks without hairs and male with bristle comb on antenna.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern India and Nepal. Habitat unrecorded, but collected

over a wide elevational range (200 - 2500 m).

Behaviour: Unknown. Seasonality: May to June.

132

Map

142 Tiny-spotted Lophyra parvimaculata.

Tiger

Beetle,

Species Accounts

Plaited Tiger Beetle, Lophyra vittigera (Dejean) (Plate 19) [Map 143] Identification: Length 12 - 13 mm; head and pronotum dark copper to greenish

copper; elytra dull velvety green with ivory markings that include a longitudinal line from the shoulder, a curved mid-line, six small lines in between and two spots toward the rear; cheeks with a few hairs.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern India, Pakistan, Nepal, and Bangladesh. Clearings In tall grass habitats and agricultural fields of flood plains (500 - 800 m).

Behaviour: Unknown.

Map 143 Plaited Tiger Beetle, Lophyra vittigera.

Seasonality: June to July.

Many-spotted Tiger Beetle, Lophyra multiguttata (Dejean) (Plate 19) [Map 144] Identification: Length 12-13 mm; head and pronotum dark metallic copper above

and green and blue-copper on the sides; elytra greenish black with large yellow

marking at the shoulder and five or six smaller spots that often run together, the

rear marking is crescent-shaped; cheeks

distinctly but sparsely covered with hairs (setae). Superficially similar to the Twentyspotted Tiger Beetle (C. vigintiguttata) but smaller with different markings on the elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and

habitats:

Northern

Map

144 Many-spotted Tiger Lophyra multiguttata.

Beetle,

and northeastern India, Nepal, Bhutan, Bangladesh, and Myanmar. Moister grassy parts of river flood plains and savannas (250 - 2700 m).

Behaviour: Occurs together with the Belted Tiger Beetle (L striolata).

Seasonality: April to August. 133

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Woolly Tiger Beetles, genus Chaetodera

J

Seven species of this genus are found across Africa and central Asia. Two spi enter into northern parts of India, but neither is endemic to India. They $ complex patterns of lines on the elytra and, except for elytra, their bodiei covered both above and below with dense white, flattened hairs (setae). }

Key XVII: Woolly Tiger Beetles, genus Chaetodera Identification Key to Chaetodera Species 1a.

Head on cheeks and between eyes

White Tiger Beetle,

densely covered by white flattened hairs,

Chaetodera albino

elytra each with pattern of fused bands nearly obscuring the surface

1b.

Head bare on cheeks and between eyes,

Twenty-spotted Tigi

elytra each with pattern of ten oval or

Beetle, Chaetodera vigintiguttata

elongated spots distributed over the entire dark surface

White Tiger Beetle, Chaetodera albino (Wiedemann) (Plate 19) [Map 145] Identification: Length 13-16 mm; head and pronotum greenish-copper and covered by

dense white, flat hairs (setae); elytra almost completely covered by four broad whitish stripes with narrow intermittent dark copper

areas.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern India, Pakistan, Bangladesh, and Nepal. White

sandy areas along small and large rivers,

sand bars, and sand dunes (120 - 450 m).

134

Behaviour: Unknown.

Map 145 White Tiger Beetle, Chael

Seasonality: March to December.

albina.

Species Accounts

Twenty-spotted Tiger Beetle, Chaetodera vigintiguttata (Herbst) (Plate 19) [Map 146] Identification: Length 14.5 - 16 mm; head and pronotum copper above and violet on the sides; elytra black-green with ten oval and round yellowish spots on each elytron.

Subspecies and morphological variants: No subspecies have been described.

and habitats: Northern and northeastern India, Pakistan, Nepal, Distribution

Bhutan, and Bangladesh. Clearings in tall grasses along river flood plains and adjacent

agricultural fields (350 - 2100 m).

Behaviour: Occasionally flies up to land on leaves of undergrowth vegetation. Attracted

to lights at night.

Seasonality: March to November.

Map 146 Twenty-spotted Tiger Beetle, Chaetodera vigintiguttata.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

J

Tethys Tiger Beetles, genus Jansenia

Members ofJansenia are distinguished by an enlarged, bulky and peculiarly tapered! tip to the male aedeagus (genitalia) with a long convoluted internal whip—likeI structure (flagellum), long labrum covering mandibles with asymmetrical teeth|

reduced in size and number, mouth with second enlarged segment of jointed J tasters (labial palpi) below mandibles, and head and pronotum lacking hair-like setae. Most species are found on the ground in open forest, but some are restricted] to faces of large boulders and rocky cliff sides. The 40 species in this genus are] confined to the general Indian Subcontinent area. Thirty-two are endemic to India, two occur only in Myanmar; four are endemic to Sri Lanka, and two Indian] species are shared with Bhutan, Nepal, and Bangladesh. > Although studies of Jansenia during the last few decades have discovered many new species, more new species are likely to be discovered with additional]

j

field work, especially in the southern and central states of India.

The origin of this unique genus is uncertain, but it most likely involved theiq ancestors radiating Into a rich diversity of species as they drifted on the ancient Indian tectonic plate across the Tethys Sea from Gondwana to eventually collide) with Laurasia millions of years ago.

The key to species and their accounts that follow, along with their images,: should allow identification of the numerous Jansenia tiger beetle species occurring in India provided specimens are imaged or collected and examined! carefully; some specimens though may require study of male aedeagus shape and] tip under magnification for a mpre definitive identification due to the superficial similarity of the many species concentrated in southern India. J

Key XVIII: Tethys Tiger Beetles, genus Jansenia Identification Key to Jansenia Species la.

Thorax cylinder-shaped, longer than wide,

2

narrower than head and elytra; anterior transverse furrow on pronotum deeply and

evenly impressed to form an anterior collar

1b.

Thorax square-shaped, wider than long; anterior

transverse

furrow

on

pronotum

4

shallowly

impressed or if deeply impressed then unevenly so as not form an obvious anterior collar

2a. (la.)

Elytra shoulder area metallic marked with

copper or bluish iridescence; first large oval

segments (coxae) of front and middle legs opaque, metallic 136

3

Species Accounts

2b.

Elytra shoulder area nonmetallic pale yellow;

Spindle-necked Tethys

first large oval segments (coxae) of front and

Tiger Beetle, Jansenia fusissima

middle legs pale yellow with only a slight metallic sheen

3a. (2a.)

Elytra shoulder area iridescent copper-green,

Blue-margined Tethys

contrasting with blue lateral iridescent band;

Tiger Beetle,

female thorax below elytra shoulder with a pit

Jansenia stuprata

at centre

3b.

Elytra shoulder area iridescent blue, merging

Blue-shouldered

with blue lateral iridescent band; female thorax

Tethys Tiger Beetle,

below elytra shoulder with a broad central

Jansenia venus

groove 4a. (1b.)

Elytra colours on margins and centres abruptly

5

contrasting; lateral margin iridescent blue, blue-green to purple, broadly occupying much

of shoulder angle; similarly* coloured, but narrower inner margin; centre bright purplish

red or polychromatic; elytra centre lacking large metallic dimples (foveae) contrasting

with surface colour 4b.

Elytra colours on margins and s gradually blending;

lateral

margin

diffuse

9

metallic

green to copper or shiny black to violet and

red, narrowly occupying shoulder angle; only

suture slightly metallic; elytra centre varied in colour, either shiny black or dull black, bronze, green or greenish brown, but not purplish red or multicoloured (polychromatic); elytra centre

on most specimens with large or small metallic

dimples (foveae) contrasting with dark surface 5a. (4a.)

Elytra lateral margins with iridescent colours

6

interrupted by copper-red or purplish red colour intruding from centre 5b.

Elytra lateral margins with iridescent colours

7

continuous 6a. (5a.)

Elytra centre copper with olive-bronze stripe; pronotum laterally bare except for a few

6b.

Crimson Tethyfc Tiger Beetle,

flattened hairs on some specimens

Jansenia rugoskeps

Elytra centre entirely purple-red; pronotum

Sandur Tethys Tiger

laterally with numerous flattened hairs

Beetle,

Jansenia sandurka 137

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

7a. (5b.)

!

Elytra iridescent colours of lateral margins wavy

Purple-sided Tethys

at middle; head with upper lip (labrum) lacking

Tiger Beetle,

abrupt middle ridge; longest leg segment

Jansenia azureocincta ।

(femora) pale and translucent 7b.

Elytra iridescent colours along lateral margins

(

8

straight at middle; head with upper lip (labrum)

8a. (7b.)

with abrupt middle ridge; longest leg segment (femora) opaque and metallic

: j

Elytra marked with two large oval spots and iridescence on outer and inner margins broadly

Twin-spotted Tethys ' Tiger Beetle, j

joined at apex; pronotum about as wide as

Jansenia chloropleura .

long, deep anterior and posterior transverse

furrows contrasting in colour with centre 8b.

j

Elytra marked with small faint dots varied in

Green-sided Tethys

!

presence and number (two or three dots or

Tiger Beetle,

)

none) with iridescence on outer and inner

Jansenia viridicincta

margins separated at apex; pronotum parallel sided, shallow anterior and posterior transverse

furrows not contrasting in colour with centre 9a. (4b.)

Front upper lip (labrum) surface roughened,

10

metallic, dark, bristles arising both behind front edge and at centre of middle ridge

9b.

Front upper lip (labrum) surface smooth,

13

nonmetallic, partially pale yellow or dull blackish brown, most bristles arising behind front edge, only one or two on front edge of

middle ridge 10a. (9a.)

Elytra without any markings or wide metallic

11

dimples (foveae); pronotum bare at sides 10b.

Elytra with two large spots at centre and wide

12

metallic dimples (foveae) at their centres;

pronotum hairy at sides 11a. (10a.)

Ridges on top of head between eyes mostly

Tiger Beetle,

large smooth areas free of punctures on centre

Jansenia indica

and laterally; elytra surface intermixed with small

shallow punctures, separated not coalesced, surrounded by a shallow iridescent blue-green

and purple depression; thorax and abdomen on

both sexes sparsely haired

138

Metal-lipped Tethys

straight to slightly wavy; elytra dull with many

Species Accounts

11b.

Ridges on top of head between eyes forming

Carpet-backed Tethys

irregular network; elytra shiny and polished with

Tiger Beetle, Jansenia reticulella

one or two small areas free of punctures only on centre, not laterally; elytra surface intermixed

with large deep punctures on centre often coalesced, not surrounded by iridescence; thorax

and abdomen on most females bare and only sparsely haired on male

12a. (10b.)

Head with cheeks on most specimens sparsely

Bronze-spotted Tethys

hairy near lower margin or bare; top of head

Tiger Beetle, Jansenia

concave with eyes large and prominent; female

tetragrammica

thorax below elytra shoulder with a shallow middle pit or small cavity 12b.

Head with cheeks bare, vertex of head flattened

Flat-headed Tethys

with eyes small and not prominent; female thorax

Tiger Beetle,

below elytra shoulder with a broad, indefinite

Jansenia appianata

groove or shallow dimple along posterior margin 13a. (9b.)

13b.

14a. (13a.)

14b.

Elytra centre with two small, irregular, and

Copper Tethys Tiger

poorly defined dull black areas contrasting with

Beetle, Jansenia

remainder of surface

motschulskyana

Elytra centre without irregular dull black areas

14

Elytra markings absent; labrum long, large abrupt

Purple-margined

middle ridge, dull and mostly pale yellow or black

Tethys Tiger Beetle,

with faint pale yellow areas on each side of ridge

Jansenia prothymoides

Elytra markings present; labrum short or long,

15

but middle ridge broadened or nonexistent and

most of its surface pale yellow with only edges or middle ridge darkened

15a. (14b.)

Elytra oval shaped with lateral sides convex with

16

shoulder angles nearly obtuse 15b.

Elytra rectangular shaped with lateral sides

17

parallel with shoulder angles nearly at a right angle

16a. (15a.)

Elytra with two lateral markings, anterior one

rounded and set inward from margin, posterior one largest, pear-shaped and touching margin 16b.

Flightless Tethys Tiger Beetle,

Jansenia westermanni

Elytra with three lateral markings, anterior and

Green-spotted Tethys

posterior ones forming thin narrow bands near

Tiger Beetle, Jansenia

margins, middle one rounded

pseudodromica

139

h Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

17a. (15b.)

17b.

Elytra with one or two lateral spots

Elytra with three lateral spots

18 Speckled Tethys Tiger Beetle, Jansenia crassipalpis

18a. (17a.)

Elytra transverse middle band present, extending

White-banded Tethys '

from lateral margin nearly to suture; elytra

Tiger Beetle,

surface abruptly changing from densely and

Jansenia grossula

'

deeply punctured in front of middle band to no

punctures behind middle band; metallic elytra dimples absent 18b.

Elytra transverse middle band absent, usually with

19

two spots laterally positioned, either separate or

joined, rarely one spot or a minute shoulder dot; elytra surface either gradually changing from deeply to shallowly punctured or punctures

of constant depth throughout; metallic elytra

dimples often present

19a. (18b.)

Elytra surface rough and unevenly contoured

20

either across all or part of entire width in front

half; elytra centre with one or more narrow

longitudinal furrows creating abrupt or shallow raised and irregular ridges on basal two-thirds, at

most only a few slightly raised areas apically 19b.

Elytra surface smooth, evenly contoured across

28

entire width in front half; elytra centre entirely lacking any surface raised areas or contrasting

colours throughout

20a. (19a.)

Elytra surface rough and unevenly contoured

21

across entire width on basal two-thirds creating

two narrow longitudinal ridges, either wavy

edged or straight edged with numerous small, irregular, coarsely raised areas toward the tip

20b.

Elytra surface contours slightly rough and uneven

25

only across inner half of width creating one broad longitudinal ridge, regular in width on

basal two-thirds becoming irregular with one to

a few large slightly raised areas toward the tip 21a. (20a.)

Elytra markings both aligned only along lateral margins in rear half, front markings as small

spots, rear markings as much larger ovals near outer angle 140

j j

22

Species Accounts

21b.

Elytra markings both not placed along lateral

Nathan's Tethys Tiger

margins in rear half, front markings displaced

Beetle, Jansenia nathanorum

inward as a very small dot at middle, rear markings a much larger oval spot near outer angle

22a. (21b.)

Elytra markings if present as two small

23

ragged edged spots in rear half close to

lateral margins, front spots smallest and slightly transverse, back spots largest often

with marginal extensions, spots reduced in size or missing on some specimens

22b.

Elytra markings always present as two large

Undulated Tethys Tiger

elongated spots in rear half close to lateral

Beetle,

margins, front spots slightly smaller and

Jansenia biundata

placed close to distinctly larger back spots with

marginal extensions 23a. (22a.)

Prothorax at sides with shallow parallel wrinkles

confined

to top

margin,

and

noticeably

shallower than moderately impressed parallel

Shaggy Tethys Tiger Beetle,

Jansenia dasiodes

ridges at lateral edge; middle of thorax at sides with sparse flattened hairs scattered on surface 23b.

Prothorax at sides with moderately impressed

24

wavy wrinkles from top edge extending downward to cover most of top half, and

wrinkled surface at top about as coarse as that at lateral edge; middle of thorax at sides almost

bare, no more than a few flattened hairs near

lower back margin 24a. (23b.)

24b.

Elytra dimples (fqveae) metallic blue-green in

Bristled Tethys Tiger

two sparse irregular rows, one on each side of

Beetle, Jansenia

middle ridge

semisetigerosa

Elytra dimples (foveae) metallic blue-green in one

Waved Tethys Tiger

sparse row on inner side of middle ridge, a few

Beetle, Jansenia

specimens with one fovea on lateral side 25a. (20b.)

Thorax at

sides

covered

with

moderately

vestiplicatica

Band-lipped Tethys

impressed wavy wrinkles about as coarse as

Tiger Beetle,

those at lateral edge on upper surface, side of

Jansenia legnotia

thorax mostly covered with hairs coming out of small punctures over most of its surface on male,

only on bottom half on female

141

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

25b.

Thorax at sides covered with shallow, parallel,

26

wrinkles confined to top margin and noticeably shallower than those at lateral edge of upper

surface, side of thorax mostly bare except for

hairs coming out of large punctures only on bottom half on male, only near bottom margin on female

26a. (25b.)

Elytra centre mostly shiny copper bronze

Bengaluru Tethys

with oval markings close to lateral edges,

Tiger Beetle, Jansenia bangalorensis f

front markings behind middle small and rear markings at outer angle only slightly larger

26b.

Elytra centre partially or completely black

27

with irregularly shaped markings only near lateral margins, front markings behind middle distinctly

smaller than

much

larger

rear

markings at outer angle

27a. (26b.)

Elytra centre shining partially black, interrupted

Streaked Tethys Tiger

by a narrow copper or copper-green streak

Beetle,

along line of dimples (foveae) between the

Jansenia plagatima

broad black longitudinal ridge and broad black

inner band, laterally with a broad copper-green margin flanked by a narrow purple edge, suture

copper; distal end of longest leg segments

(femora) with the boundary between pale yellow and unpigmented portions distinctly

defined on all surfaces 27b.

Elytra centre shining uniformly shiny black,

Purple-banded Tethys

laterally with a broad copper or copper-green

Tiger Beetle,

lateral band flanked by a broad purple margin,

Jansenia ostrina

suture black; distal end of longest leg segments

(femora) with the boundary between pale yellow

and unpigmented portions poorly defined on posterior surface

28a. (19b.)

Colours at lateral elytra margins and centre abruptly contrasting, marginal colour varied

either as narrow or wide iridescent multicoloured bands extending uniformly from shoulder at least to middle of lateral spots or as highly polished

bands with numerous punctures, contrasting abruptly in texture, luster and colour with

remainder of surface

142

29

Species Accounts

28b.

Colours at lateral elytra margins and centre

32

gradually merging, all specimens with no more than a narrow violet edge not reaching

margins

shoulder, shiny diffuse

gradually

merging in texture, luster and colour with

remainder of surface 29a. (28a.)

Elytra centre shiny black or copper-black, lateral

Red-headed Tethys

margins wide with violet, blue, green and red

Tiger Beetle, Jansenia tetrastacta

iridescence uniformly tapering anteriorly to shoulder angle and posteriorly to outer rear

angle, male front and middle legs with wide

pads on bottoms of second and third segments

29b.

Elytra centre dull black, bronze, or green to

30

greenish brown, lateral margins narrow with metallic copper, purple, blue or green, male

front legs only with wide pads bn bottoms of

second and third segments

30a. (29b.)

Elytra

centre

green

greenish

to

brown,

contrasting coloured dimples (foveae) absent-

lateral elytra margins broadly metallic copper

30b.

Elytra

centre

black,

bronze

or

copper,

Green-backed Tethys Tiger Beetle,

Jansenia chlorida

31

contrasting dimples (foveae) green, blue or purple in an irregular inner row, additional

dimples (foveae) in middle, especially on rear third and at inner shoulder angle; lateral elytra

margins broadly metallic purple to bluish and copper-green

31a. (30b.)

Longest

leg

segments

(femora)

opaque,

Green-legged Tethys

metallic copper-green except at extreme distal

Tiger Beetle,

end; labrum tawny except for wide black border

Jansenia psarodea

along entire anterior margin and extending

onto middle ridge for both sexes

31b.

Longest leg segments (femora) translucent,

Large-lipped Tethys

pale yellow throughout entire length; labrum

Tiger Beetle,

uniformly tawny over entire surface, narrowly

Jansenia rostrulla

darkened along entire anterior edge and barely

onto middle ridge for female 32a. (28b.)

Longest leg segments (femora) pale, entirely or

33

partially translucent often with slight metallic reflections blending with pale semi-translucent

portion at tip 143

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

32b.

Longest leg segments (femora) opaque, mostly

Burnished Tethys

metallic copper or greenish black except for

abrupt pale semi-translucent portion at tip

33a. (32a.)

Elytra maculae large, broadly touching lateral

Tiger Beetle,

f

Jansenia corrugatosa

Pitted Tethys Tiger

margin; anterior spot elongate and posterior spot

Beetle,

oblong with a long, rear extension; elytra typically

Jansenia cratera

with numerous metallic blue-green dimples

(foveae) at middle in addition to a row close to inner edges 33b.

Elytra maculae small, not touching lateral margin;

f Pale-legged Tethys

anterior and posterior spots oval, the latter with

Tiger Beetle,

a short rear extension only on a few specimens;

Jansenia choriodista

elytra typically with only a row of metallic blue­

green dimples (foveae) close to inner edges, on a few specimens with one or two middle dimples (foveae)

Flightless Tethys Tiger Beetle, Jansenia westermanni (Schaum) (Plate 19) [Map 147] Identification: Length 10-11 mm; head and pronotum copper; elytra oval-shaped, dull black edged with shiny copper, with

many green reflecting pits on surface, and medium yellow spot at mid-margin and large patch at rear.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Open grassy fields and scrub forest floor (430 - 930 m). Behaviour: Flightless and runs quickly to hide under forest floor leaf litter and bases of grass clumps when disturbed. Closely resembles females

of local

species

stinging Velvet Ants (Family Mutillidae).

Seasonality: July to September.

144

of

Map 147 Flightless Tethys Tiger Beetle, Jansenia westermanni.

Species Accounts

Green-spotted Tethys Tiger Beetle, Jansenia pseudodromica (Horn) (Plate 19) [Map 148] Identification: Length 11 mm; head and pronotum copper-brown; elytra oval­ shaped, dull black with copper margins, green reflecting surface pits, and yellow central spot and two short longitudinal bands near margin at front and rear.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known only from the type locality in Tiruchchirappalli

District, Mudukolam, India (Tamil Nadu).

Behaviour:

Flightless

and

likely

has

behaviour similar to that of the Flightless

Tiger Beetle (J. westermanni).

Map

148 Green-spotted Tethys Tiger Beetle, Jansenia pseudodromica.

Seasonality: Apparently adults are active during the Autumnal monsoon (November).

Nathan's Tethys Tiger Beetle, Jansenia nathanorum Cassola and Werner (Plate 20) [Map 149] Identification: Length 8 - 8.5 mm; Head copper with green reflections above and golden-green on the sides; pronotum copper-bronze; elytra dark bronze to shiny black with two dots on each elytron differently positioned than any other known Jansenia species - a small one in the middle half-way between the margin and the inner edge and a larger one near the rear margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern portion of the Western Ghats, Quilon District, Thenmala (900 - 1000 m) in southern India (Kerala).

Map 149 Nathans Tethys Tiger Beetle, Jansenia nathanorum.

Behaviour: Unknown. Seasonality: May to June.

145

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Shaggy Tethys Tiger Beetle, Jansenia dasiodes (Acciavatti and Pearson) (Plate 20) [Map 150] Identification: Length 8.5 - 10.5 mm; head and pronotum copper above and blue-green on the sides; elytra shiny black streaked with

copper or copper-green, surface rough and unevenly contoured, two longitudinal furrows on each elytron, usually with two ivory spots on rear half of each elytron. Similar to the Undulated Tethys Tiger Beetle

(J. biundata), which is dark brown rather than copper, and whose elytra spots are

larger and more oblong.

Subspecies and morphological variants: Individuals from northeastern Karnataka have elytra spots reduced in size or entirely missing.

Map 150 Shaggy Tethys Tiger Beetle, Jansenia dasiodes.

Distribution and habitats: Endemic to southern India. Scrub forest floor (700 1500 m).

Behaviour: Unknown. Seasonality: April to October.

Undulated Tethys Tiger Beetle, Jansenia biundata Naviaux (Plate 20) [Map 151] Identification: Length 8.8 mm; dark brown; elytral surface rough with two longitudinal furrows on each elytron, two light oblong marks along the rear margin of each elytron.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India (Karnataka). Scrub forest floor (900 m).

Behaviour: Unknown.

Seasonality: June. Map 151 Undulated Tethys Tiger Beetle, Jansenia biundata. 146

Species Accounts

Bristled Tethys Tiger Beetle, Jansenia semisetigerosa (Acciavatti and Pearson) (Plate 20) [Map 152] Identification: Length 8.5-10 mm; head and pronotum shiny copper above and blue­

green on the sides; elytra shiny black with copper streaks, rough and uneven surface

with single longitudinal furrow on each elytron, blue pits with green reflections, and two small light spots along rear margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Forest paths and old fields (770 - 900 m).

Behaviour: Unknown. Seasonality: May to June.

Map 152 Bristled Tethys Tiger Beetle, Jansenia semisetigerosa.

Waved Tethys Tiger Beetle, Jansenia vestiplicatica (Acciavatti and Pearson) (Plate 20) [Map 153] Identification: Length 8 - 9.5 mm; head and pronotum bright copper above and copper to copper-green on the sides; elytra shiny

copper-black with a wide copper band and a narrow bluish green band running along the edge, surface very rough and uneven with

indistinct longitudinal furrow on front half of each elytron, small transverse and oblong light marking at mid-margin and larger one at rear. .

Subspecies and morphological variants: No subspecies have been described but

some individuals lack the small spot on the mid-margin and others have the two spots connected along the margin in between them.

Map 153 Waved Tethys Tiger Beetle, Jansenia vestiplicatica.

Distribution and habitats: Endemic to southern India. Forest paths (760 m). Behaviour: Unknown.

Seasonality: May to October. 147

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cldndelldae

Band-lipped Tethys Tiger Beetle, Jansenia legnotia (Acciavatti and Pearson) (Plate 20) [Map 154] Identification: Length 8.8 mm; head and pronotum shiny copper above and copper-

green on the sides; elytra shiny black along raised

centre

contrasting with copper margin, surface

or copper-green along

relatively smooth with two light spots along the rear margin of each elytron, the middle one usually smaller than the rear one.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Floor of scrub forest (425 m).

Behaviour: A fall monsoon species.

Seasonality: September to October.

Map 154 Band-lipped Tethys Tiger Beetle, Jansenia legnotia.

Streaked Tethys Tiger Beetle, Jansenia plagatima (Acciavatti and Pearson) (Plate 20) [Map 155] Identification: Length 8.5 - 9.5 mm; head and pronotum shiny copper on top and

copper-green on the sides; elytra shiny black down the length and copper or copper-

green along the sides with a relatively smooth surface and two small white spots

along the rear half of the margin, usually the middle one is smaller than the rear one.

Similar to the Bengalaru Tethys Tiger Beetle (J. bangalorensis) but duller in colour and with a different-shaped male aedeagus.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and

habitats:

Endemic

to southern India. Floor of scrub forests

(660 m).

Behaviour: Unknown. Seasonality: May to September. 148

Map 155 Streaked Tethys Tiger Beetle, Jansenia plagatima.

Species Accounts

Bengaluru Tethys Tiger Beetle, Jansenia bangalorensis Cassola and Werner (Plate 20) [Map 156] Identification: Length 7.5 - 8.2 mm; head and pronotum shiny copper above and blue-green at sides; elytra shiny copperbronze, surface relatively smooth except for dense punctures on the front half, small whitish yellow spots on mid-margin and rear. Similar to the Streaked Tethys Tiger Beetle (J. plagatima), but brighter in colour and with a different-shaped male aedeagus.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Presently known only from floor of scrub forest in the vicinity of the city of Bengalaru in South India (Karnataka).

Map 156 Bengaluru Tethys Tiger Beetle, Jansenia bangalorensis.

Behaviour: Unknown.

Seasonality: July.

Purple-banded Tethys Tiger Beetle, Jansenia ostrina (Acciavatti and Pearson) (Plate 20) [Map 157] Identification: Length 8.5 - 9.5 mm; head and pronotum shiny copper above with blue green between eyes, and blue green on the

sides; elytra shiny black down the centre and broad copper-green and purple band down the sides, broad ridge on surface running the length of the elytra and two whitish spots - small one at mid-margin and larger one at rear. Similar to the Streaked Tethys Tiger Beetle (J. plagatima) from which it is distinguished by its uniformly shiny black elytral surface and broad purple band along the margins of the elytra.

Subspecies and morphological variants:

Map 157 Purple-banded Tethys Tiger Beetle, Jansenia ostrina.

No subspecies have been described.

Distribution and habitats: Endemic to central eastern India. Floor of scrub forests (100-550 m).

Behaviour: Above 500 m occurs with the Red-headed Tethys Tiger Beetle (J. tetrastacta).

Seasonality: March to July. 149

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Burnished Tethys Tiger Beetle, Jansenia corrugatosa (Acciavatti and Pearson) (Plate 21) [Map 158] Identification: Length 8.5-10 mm; head and pronotum shiny copper above with

spots of blue-green between eyes and blue­ green or bronze on the sides; elytra shiny black with an indistinct bronze longitudinal band and a narrow purple edge, surface

with no ridges or irregularities and two white spots along rear margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Floor of open scrub foresi (425 m).

Behaviour: Active during the fall monsoon.

Map 158 Burnished Tethys Tiger BeetL Jansenia corrugatosa.

Seasonality: September to December.

Pitted Tethys Tiger Beetle, Jansenia cratera (Acciavatti and Pearson) (Plate 21) [Map 159] Identification: Length 9 - 10.5 mm; head and pronotum shiny copper above with patches of bronze-green on the sides; elytra copper to bronze in centre grading into diffuse copper green and a narrow purple edge, smooth with large distinct punctures and two large white marks on the rear margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Floor of scrub forest. Behaviour: Active during fall monsoon.

Seasonality: October.

150

Species Accounts

Pale-legged Tethys Tiger Beetle, Jansenia choriodista (Acciavatti and Pearson) (Plate 21) [Map 160] Identification: Length 8.5 - 10.5 mm; head and pronotum shiny copper and purple

above and shiny copper to bronze on the sides; elytra shiny bright copper to green

with wavy wrinkles on surface and two large white dots on rear half of margin; upper parts of legs light and translucent with a tinge of green.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Scrub forest floor (250 - 300 m).

Behaviour: Active during fall monsoon.

Seasonality: September to November.

Map 160 Pale-legged Tethys Tiger Beetle, Jansenia choriodista.

Purple-margined Tethys Tiger Beetle, Jansenia prothymoides (Horn) (Plate 21) [Map 161] Identification: Length 8-9 mm; head and pronotum shiny copper-black above and

blue-green to violet on the sides; elytra black to copper-black with a wide shiny purple band along the sides, surface rough and uneven with no light markings.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern Western Ghats in southern India. Open floor of moist forests (750 m).

Behaviour: Sometimes occurs with the more common Bristled Tethys Tiger Beetle (J. semisetigerosa).

Map 161 Purple-margined Tethys Tiger Beetle, Jansenia prothymoides.

Seasonality: May to July.

151

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Speckled Tethys Tiger Beetle, Jansenia crassipalpis (Horn) (Plate 21) [Map 162] Identification: Length 10 - 13.5 mm; head and pronotum shiny copper with green and red reflections; elytra black with copper and blue-green reflections and a broad copper green band along sides with three whitish spots along the margin - a small oval near

the front, a large oval in the middle and a crescent following the margin at the rear end.

Subspecies and morphological variants: No subspecies have been discovered.

Distribution and habitats: Endemic to southern India. Along paths through shrubby forested slopes (300 - 700 m) in

Map 162 Speckled Tethys Tiger Beeth Jansenia crassipalpis.

Behaviour: Unknown.

Seasonality: June to October.

Red-headed Tethys Tiger Beetle, Jansenia tetrastacta (Wiedemann) (Plate 21) [Map 163] Identification: Length 9 - 10 mm; head and pronotum crimson above and shiny blue to green on the sides; elytra shiny dark black or copper black with a wide band of violet, blue, green, and red iridescence along the sides, two white spots along rear margin.

The middle spot varies in size from absent to distinct.

Subspecies and morphological variants: Two subspecies have been described, but

the differences are so slight that some experts consider them a single form.

J. t. tetrastacta - the middle spot on each elytron is distinct. J. t. delhiensis - the middle spot on each elytron is absent to faint.

Map 163 Red-headed Tethys Tiger Beetli Jansenia tetrastacta; A, J. t. tetrastacta, 1 J. t. delhiensis.

Distribution and habitats: Open areas on floor of secondary growth forests am occasionally into adjacent grassy fields (200 - 550 m).

J. t. tetrastacta - eastern and northeastern India and Nepal.

Species Accounts

J. t. delhiensis - presently known only from the Union Territory near New Delhi and

Uttarakhand.

Behaviour: Attracted to lights at night.

Seasonality: June and July.

Green-backed Tethys Tiger Beetle, Jansenia chlorida (Chaudoir) (Plate 21) [Map 164] Identification: Length 10 - 10.5 mm; head and pronotum green occasionally copper-

green; elytra uniformly dull green with a copper margin and two large whitish markings on the rear margin, one an oval at mid-margin and the other a narrow band

along the curved apex.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and

habitats:

Endemic

to south India. Floor of scrub forest and

adjacent open areas (430 m).

*

Behaviour: Active during the fall monsoons.

Map 164 Green-backed Tethys Tiger Beede,

Seasonality: October to November.

Jansenia chlorida.

Green-legged Tethys Tiger Beetle, Jansenia psarodea (Acciavatti and Pearson) (Plate 21) [Map 165] Identification: Length 10 - 11.5 mm; head and pronotum shiny copper above with blue-green between eyes and blue-green on the sides; elytra copper-black with shiny black-bronze and purple band along sides, two large yellow patches on rear margin and notably rounded sides of elytra. Upper parts of legs metallic green.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to in southern India (Karnataka, Tamil Nadu). Floor of scrub forest and adjacent openings or sandy roads (916 m).

Map 165 Green-legged Tethys Tiger Beetle, Jansenia psarodea.

Behaviour: Unknown.

Seasonality: May to October. 153

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Large-lipped Tethys Tiger Beetle, Jansenia rostrulla (Acciavatti and Pearson) (Plate 22) [Map 166] Identification: Length 10.1 mm; head and pronotum green or copper; elytra copperbrown with smooth surface and longitudinal row of distinct pits reflecting green, metallic copper band along sides with two yellow spots on rear margin. Long, ivory-coloured labrum.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Floor of scrub forests (300 m). Behaviour: Active during fall monsoons.

Seasonality: September to November.

Map 166 Large-lipped Tethys Tiger Beetle, Jansenia rostrulla.

White-banded Tethys Tiger Beetle, Jansenia grossula (Horn) (Plate 23) [Map 167] Identification: Length 8 mm; head and pronotum dark bronze with violet reflections above and dark blue-back sides; elytra

distinctly oval-shaped, dull dark bronze with a large whitish band extending almost

entirely across the mid elytra and a large spot at the rear margin.

Subspecies and morphological variants: No subspecies have been described for this very rare species.

Distribution and

habitats: Apparently

a rare species as its existence is based on single specimen collected in Southern Tamil Nadu.

Behaviour: Unknown.

Seasonality: Unknown.

154

Map

167 White-banded Tethys Tiger Beetle, Jansenia grossula.

Species Accounts

Blue-shouldered Tethys Tiger Beetle, Jansenia venus (Horn) (Blate 22) [Map 168] Identification: Length 11-12 mm; head and pronotum reddish-copper with bright blue-green patches; elytra reddish-copper

with iridescent blue-violet band along sides especially wide at shoulder, two large yellow markings - middle one crosses mid elytra and round one at rear margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and habitats: Endemic to southern India. Shaded floor of coffee plantations, moist mossy rocks along mountain streams and mature forest above 700 m.

Behaviour: Flies to land on undergrowth

Map 168 Blue-shouldered Tethys Tiger Beetle, Jansenia venus.

leaves when disturbed.

Seasonality: May to June.

Blue-margined Tethys Tiger Beetle, Jansenia stuprata (Horn) (Blate 22) [Map 169]__________________________________ Identification: Length 9-10 mm; head and pronotum shiny crimson above and bright green on the sides; elytra dull black-bronze

with iridescent copper crescent at shoulder, iridescent blue along sides, and violet black along the inner edges, pale yellow line across elytra at middle and a small spot at the rear margin. Similar to the Spindle-necked Tethys

Tiger beetle (J. fusissima) but which has a

yellow-orange spot at the shoulder.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Habitat unknown, but occurs at higher elevations of the Western Ghats (60-1250 m).

Map 169 Blue-margined Tethys Tiger Beetle, Jansenia stuprata.

Behaviour: Unknown. Seasonality: May to June. 155

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Spindle-necked Tethys Tiger Beetle, Jansenia fusissima (Acciavatti and Pearson) (Plate 22) [Map 170] Identification: Length 8.5 - 9.5 mm; head and pronotum shiny crimson above with

green and purple sides; elytra shiny copper with contrasting blue-green pits and a black

and purple band along edge, yellow-orange spot on shoulder, narrow pale yellow band

across middle of elytron and small yellow spot at rear margin. Similar to the Bluemargined Tethys Tiger beetle (J. stuprata) which has a copper crescent at the shoulder.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Habitat unknown, but occurs

Map 170 Spindle-necked Tethys Tiger Beetle, Jansenia fusissima.

above 900 m in the Western Ghats.

Behaviour: Unknown. Seasonality: May.

Twin-spotted Tethys Tiger Beetle, Jansenia chloropleura (Chaudoir) (Plate 22) [Map 171] Identification: Length 10-12 mm; head and pronotum brilliant copper-red above

and metallic blue, violet and green on the sides; elytra copper brown or olive green

with a narrow blue-green band along the side and inner edges, two yellowish spots along rear margin.

Subspecies and morphological variants: A subspecies has been described for the population in Bhutan.

Distribution and habitats: Northern and northeastern India,

Nepal,

Bhutan, and

Bangladesh. Boulders and rock areas on mountain sides and above streams (150 3600 m).

Map 171 Twin-spotted Tethys Tiger Beetle, Jansenia chloropleura.

Behaviour: Runs along rocky trails and often flies to boulders, muddy banks, and near vertical surfaces to escape danger.

Seasonality: April to August. 156

Species Accounts

Green-sided Tethys Tiger Beetle, Jansenia viridicincta (Horn) (Plate 22) [Map 172] Identification: Length 9-12 mm; head and pronotum bright reddish-copper above and shiny violet green on the sides; elytra reddish-copper with

narrow green and

violet stripe on the outer edge and green stripe down the inner edge, yellowish white markings, if present, one or two small spots on the rear third.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to Western Ghats of southern India. Road cuts and open spaces within forest near steep slopes (700- 1000 m).

Map 172 Green-sided Tethys Tiger Beetle, Jansenia viridicincta.

Behaviour: Unknown.

Seasonality: May to June.

Purple-sided Tethys Tiger Beetle, Jansenia azureocincta (Bates) (Plate 22) [Map 173] Identification: Length 8-9 mm; head bright copper-red with iridescent blue area between eyes; pronotum bright copper red

above with purple band on sides and blue­ green in the middle; elytra copper-red with outer sides iridescent purple and blue and inner edge blue, two whitish spots on rear half.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and habitats: Endemic Western Ghats of southern India (Maharashtra, Goa, Karnataka). Road cuts and open areas near steep banks and cliffs (600 - 700 m).

to

Map 173 Purple-sided Tethys Tiger Beetle, Jansenia azureocincta.

Behaviour: Unknown. Seasonality: June to July.

157

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Crimson Tethys Tiger Beetle, Jansenia rugosiceps (Chaudoir) (Plate 22) [Map 174] Identification: Length 11-12 mm; head copper-red with bright blue or green sides;

pronotum purple and green; elytra dull

crimson with broad green central stripe branching to side and blue inner and outer edges, two yellowish spots on rear third. The similar Sandur Tethys Tiger Beetle (Jansenia sandurica) has a brighter reddish-maroon and blue upper surface with no green

centre stripe. It also has larger elytra spots and thicker hair-like setae on the sides of the pronotum than even the northwestern forms of the Crimson Tethys Tiger Beetle.

Subspecies and morphological variants:

MaP 174 Crimson Tethys Tiger Beetle,

Individuals in the northwestern part of its

Jansenta ruZ0SicePs-

range have more setae on the sides of the pronotum, and the yellowish spots on the elytra are larger and more distinct.

Distribution and habitats: Endemic to southern India (Tamil Nadu, Kerala, Karnataka). Rock faces of large boulders (500 - 2000 m).

Behaviour: Often forages on termites. Seasonality: May to December.

Sandur Tethys Tiger Beetle, Jansenia sandurica (Acciavatti and Pearson) (Plate 22) [Map 175] Identification: Length 8-10 mm; Head and pronotum bright copper-red above with metallic blue sides; elytra dull copperred with interrupted blue outer sides and complete blue band along inner edges, two large yellowish spots toward centre of rear. The similar Crimson Tethys Tiger Beetle (J. rugosiceps) has a duller reddish-maroon and blue upper surface with a green centre stripe. It also has smaller elytra spots and

fewer setae on the sides of the pronotum.

Subspecies and morphological variants: No subspecies are known for this highly localized species.

158

Map 175 Sandur Tethys Tiger Beetle, Jansenia sandurica.

Species Accounts

Distribution and habitats: Found on large boulders in scrub forest only near Sandur, Bellary District in southern India (Karnataka).

Behaviour: When disturbed runs to other side of boulder. Rarely flies.

Seasonality: June to July.

Copper Tethys Tiger Beetle, Jansenia motschulskyana (Horn) (Plate 23) [Map 176] Identification: Length 9-10.5 mm; head and pronotum shiny copper above and

green on the sides; elytra dull copper with

two dull black patches and shiny purple, green and copper along outer edges with one small whitish dot on rear.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to Western Ghats of southern India. Road cuts near open steep slopes and scrub forest floor (700- 1100 m) in

Behaviour: Unknown.

Seasonality: May to July.

Map 176 Copper Tethys Tiger Beetle, Jansenia motschulskyana.

Metal-lipped Tethys Tiger Beetle, Jansenia indica (Fleutiaux) (Plate 23) [Map 177] Identification: Length 7-8 mm; head and pronotum shiny black and wrinkled above

and bight dark purple and blue on the sides; elytra bright copper along the sides and the rest of the surface dull blackish bronze with raised velvety black areas and no light markings.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to western India in Maharashtra. Paths and along road cuts through forest (600 - 800 m). Behaviour: Unknown.

Seasonality: June to August.

Map 177 Metal-lipped Tethys Tiger Beetifi Jansenia indica.

Carpet-backed Tethys Tiger Beetle, Jansenia reticulella (Acciavatti and Pearson) (Plate 23) [Map 178] Identification: Length 7-8 mm; head and pronotum shiny black and heavily wrinkled above and shiny black or purple on the sides; elytra shiny bronze on the sides and the rest bronze-black with a labyrinth of raised areas across the surface, but no light markings. Labrum long and metallic copper.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Forest floor of mature forest on coastal plains.

Behaviour: Unknown.

Map 178 Carpet-backed Tethys Tiger

Seasonality: June to July.

Beetle, Jansenia reticulella.

Bronze-spotted Tethys Tiger Beetle, Jansenia tetragrammica (Chaudoir) (Plate 23) [Map 179] Identification: Length 10-13 mm; head and pronotum dull black with shiny copper

areas; elytra dull, velvety black with faint copper or metallic green at the shoulder and along the outer edge, surface with

many green and bronze pits and flecks and distinctive large and circular greenish patches, two large circular yellow spots

on rear of each elytron. Large prominent eyes. Similar to the Flat-headed Tethys Tiger

Beetle (J. appianata), which is smaller and its eyes are much less prominent.

Subspecies and morphological variants: NO subspecies have been described.

^ap 179 Bronze-spotted Tethys Tiger Beetle, Jansenia tetragrammica.

Distribution and habitats: Endemic to southern India (Karnataka, Kerala, Tamil Nadu). Sandy roads through shrubby

secondary forest (170-1900 m).

Behaviour: Unknown.

Seasonality: May to October.

160

Species Accounts

Flat-headed Tethys Tiger Beetle, Jansenia appianata (Acciavatti and Pearson) (Plate 23) [Map 180] Identification: Length 9.5-11 mm; head and pronotum dull black with copper flecks; elytra dull black with copper or metallic

green shoulder and outer edges, many green and copper flecks and large circular, metallic green patches, two large yellow oval spots on rear half of each elytron. Eyes small and barely protruding. Similar to the Bronze-spotted Tethys Tiger Beetle (J. tetragrammica), which is larger and its eyes

are much more prominent.

Subspecies and morphological variants: No subspecies have been described.

Map 180 Flat-headed Tethys Tiger Beetle,

Distribution and habitats: Endemic to Jansenia appianata. southern India (Tamil Nadu).

Habitat unknown, but occurs in the Western Ghats (2000 - 2500 m).

Behaviour: Unknown.

Seasonality: May to October.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Carpet-backed Tethys Tiger Beetle, Jansenia reticulella (Acciavatti and Pearson) (Plate 23) [Map 178] Identification: Length 7-8 mm; head and pronotum shiny black and heavily wrinkled above and shiny black or purple on the sides; elytra shiny bronze on the sides and the rest

bronze-black with a labyrinth of raised areas across the surface, but no light markings. Labrum long and metallic copper.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Forest floor of mature forest on coastal plains.

Behaviour: Unknown. Seasonality: June to July.

Map

178 Carpet-backed Tethys Tiger Beetle, Jansenia reticulella.

Bronze-spotted Tethys Tiger Beetle, Jansenia tetragrammica (Chaudoir) (Plate 23) [Map 179] Identification: Length 10-13 mm; head and pronotum dull black with shiny copper

areas; elytra dull, velvety black with faint

copper or metallic green at the shoulder and along the outer edge, surface with many green and bronze pits and flecks and distinctive large and circular greenish patches, two large circular yellow spots on rear of each elytron. Large prominent eyes. Similar to the Flat-headed Tethys Tiger

Beetle (J. appianata), which is smaller and its eyes are much less prominent.

Subspecies and morphological variants: No subspecies have been described.

Map 179 Bronze-spotted Tethys Tiger Beetle, Jansenia tetragrammica.

Distribution and habitats: Endemic to southern India (Karnataka, Kerala, Tamil Nadu). Sandy roads through shrubby

secondary forest (170-1900 m).

Behaviour: Unknown.

Seasonality: May to October.

160

Species Accounts

Flat-headed Tethys Tiger Beetle, Jansenia applanata (Acciavatti and Pearson) (Plate 23) [Map 180] Identification: Length 9.5-11 mm; head and pronotum dull black with copper flecks; elytra dull black with copper or metallic

green shoulder and outer edges, many green and copper flecks and large circular, metallic green patches, two large yellow oval spots on rear half of each elytron. Eyes small and barely protruding. Similar to the Bronze-spotted Tethys Tiger Beetle (J. tetragrammica), which is larger and its eyes

are much more prominent.

Subspecies and morphological variants: No subspecies have been described.

Map 180 Flat-headed Tethys Tiger Beetle,

Distribution and habitats: Endemic to Jansenia applanata. southern India (Tamil Nadu).

Habitat unknown, but occurs in the Western Ghats (2000 - 2500 m).

Behaviour: Unknown.

Seasonality: May to October.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Pygmy Tiger Beetles, genus Glomera The two endemic species that make up this genus are the smallest tiger beetles found in India« Adults have smooth elytra with glossy purple-black margins, but no light markings. They also lack noticeable hairs (setae) on their underside. They have a uniquely coiled structure within the bulbous male genitalia that indicates they have been evolutionarily isolated for a long time.

Key XIX: Pygmy Tiger Beetles, genus Glomera Identification Key to Glomera Species la.

Body entirely without hairs on thorax,

Immaculate Pygmy Tiger

only finely hairy on abdomen, elytra with

Beetle, Glomera belloides

wide glossy purple-black lateral margins

extend completely from shoulder to tip

1b.

Body with sparse hairs only on underside

Pale-legged Pygmy Tiger

of thorax and fine hairs on abdomen,

Beetle,

elytra with wide glossy purple-black lateral

Glomera ochrocnemis

margins extend only from front to outer

angles

Immaculate Pygmy Tiger Beetle, Glomera belloides (Horn) (Plate 23) [Map 181] Identification: Length 5.5 - 6 mm; head and pronotum black, bronze, or copper above and blue-green and purple on the

sides; elytra with glossy purple-black band

along outer edge and broad shiny coppergreen down the centre, no light markings; underside of the abdomen with few to no hair-like setae. Similar to the Pale-legged Pygmy Tiger Beetle (G. ochrocnemis) but distinguished by the lack of setae on the

abdomen.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Along forest paths.

Behaviour: Unknown.

Seasonality: May to June. 162

Map 181 Immaculate Pygmy Tiger Beetle, Glomera belloides.

Species Accounts

Pale-legged Pygmy Tiger Beetle, Glomeraochrocnemis (Acciavatti and Pearson) (Plate 23) [Map 182] Identification: Length 6 - 6.5 mm; head and pronotum shiny bronze above and bluish-

black to purple on the sides; elytra dull black with iridescent purple, blue or copper along the sides and lobes projecting toward the middle, no light markings; sparse hair-like setae on the underside of the rear abdomen. Similar to the Immaculate Pygmy Tiger

Beetle (G. belloides) but distinguished by the hair-like setae on the end of the abdomen.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to southern India. Along road cuts (200 - 600 m).

Behaviour: Unknown.

Seasonality: May to June.

Map 182 Pale-legged Pygmy Tiger Beetle, Glomera ochrocnemis.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Lustrous Tiger Beetles, genus Setinteridenta The single species in this genus is found only in northern India, Nepal, and Myanmar. It is distinguished by unique structures within the male genitalia, labrum with setae inserted between teeth along the front margin, and its small shiny dark bronze body.

Lustrous Tiger Beetle, Setinteridenta rhytidopteroides (Horn) (Plate 23) [Map 183] Identification: Length 7-8 mm; head dark bronze above and iridescent blue-green

on the sides; pronotum greenish-bronze to copper above and purple on the sides; elytra

shiny dark bronze to copper brown with broad band of blue-green and purple along the sides, two ivory oval marks on the rear half.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern and northeastern India, Nepal, and Myanmar. Grassy meadows, croplands, stream banks

along the base of the Himalayas (400 - 1100 m).

Behaviour: Unknown.

Seasonality: June to August.

Map

183 Lustrous Tiger Setinteridenta rhytidopteroides.

Beetle,

Species Accounts

Rounded-pronotum Tiger Beetles, genus Cylindera The taxonomy of this genus is in great flux. It includes more than 230 species and presently is divided into 12 subgenera, some of which are confined to the Western Hemisphere, Africa or Eurasia. As more molecular studies are done on this immense taxon, there are certain to be many cryptic species recognized leading to changes in status and names. Although many species currently assigned to this genus externally appear quite different, they are apparently related by virtue of several shared features; relatively consistent internal male genitalic structures, small size (7.5 to 9 mm), and moderate amounts of flattened hairs (setae) on various body segments.The 43 species found in India (eight of which are endemic) are presently placed into four subgenera (Cylindera, Ifasina, Eriodera, and Eugrapha).

The key to species and their accounts that follow, along with their images, should confidently allow identification of the large number of Cylindera tiger beetle species that may be encountered throughout India. However, the small size of many of these species often requires that specimens be collected or imaged for closer examination and accurate identification.

Key XX: Rounded-pronotum Tiger Beetles, genus Cylindera Identification Key to Cylindera Species la.

Elytra markings not forming extensive patterns

2

rather as separate bands or spots often reduced

to small dots or entirely missing, elytra surface

sculpturing may include raised polished areas, dull smooth areas, brilliant metallic iridescent

areas, or glossy black areas, labrum varied in colour and texture, either entirely or partially dark or entirely white or tawny, metallic or not, body underside with moderate amounts of

flattened setae either in limited areas on certain

segments or entirely absent

1b:

Elytra markings forming an extensive standard

30

pattern of lunules either separated into spots

or forming bands with middle band often

distinctively bent and curved inward or all lunules joining a continuous lateral band,

labrum nonmetallic pale tan or white, body

underside with dense flattened setae on most

segments

165

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

2a. (la.)

Labrum with either large white centre or pale

3

brown with darker front edge, elytra surface irregularly sculptured each with only two oval

spots on their back halves near outer margins;

occurs only in southern India

2b.

Labrum uniformly coloured and textured, dark

4

at centre, metallic or not, or entirely whitish or pale brown, with or without dark margins, elytra surface varied in sculpture and pattern

of markings, but never with combination of irregular surface and two oval spots only on

back halves near margins; found in many parts of India

3a. (2a.)

Body surfaces almost entirely bare, elytra surface furrowed down the middle, labrum dark

3b.

Dot-lipped Tiger

Beetle,

with large white centre

Cylindera paradoxa

Body surfaces covered by abundant setae lying

Fluted Tiger Beetle,

flat, elytra surface rough from raised polished

Cylindera lacunosa

areas, labrum dull and pale brown with a darker front edge 4a. (2b.)

Elytra distinctly widened toward tips with

5

edges smooth and without minute teeth along posterior margins, elytra centre with large well

defined irregularly shaped smooth patches, elytra markings only two irregular lateral spots

or elytra entirely without 4b.

Elytra parallel sided toward tips with small or

6

minute teeth along posterior edges, centre

without smooth patches, elytra markings varied in number and placement with one or

more located in centre or entirely unmarked 5a. (4a.)

Elytra

sutural

margin

distinct,

metallic

contrasting with remainder of surface and two

elongate markings along lateral margin 5b.

Elytra sutural margin diffuse, metallic blending

dromicoides Terrestrial Tiger

with remainder of surface, elytra markings

Beetle,

reduced to two small lateral dots on most

Cylindera delavayi

specimens, absent on some specimens

166

Fleet Tiger Beetle, Cylindera

Species Accounts

6a. (4b.)

Elytra markings a broad middle band sloping

Slope-striped Tiger

downward across the middle, a complete apical

Beetle, Cylindera obliquefasciata

lunule with wide anterior extension, small lunule at shoulder and small spot at centre

6b.

Elytra markings narrow or broken into two or

7

more irregular spots, bands or small dots, or elytra entirely without markings

7a. (6b.)

Labrum entirely dark in middle and over entire

8

surface on most specimens, but a whitish or pale tan area may be present on each side of a

darkened middle on a few specimens, darkened

labrum nonmetallic shiny brown to black on some specimens, metallic copper-green on others, purplish black on others

7b.

Labrum not dark in middle, either entirely

24

whitish or pale brown, with or without dark margins, never with metallic reflections

8a. (7a.)

Elytra marking present at outer shoulder on

9

most specimens 8b.

Elytra marking absent at outer shoulder on

14

most individuals or if minute marking present

on other specimens then remainder of elytra without spots 9a. (8b.)

Most of apical elytra margin with a separate and

10

distinct band on some specimens or attached to

an anterior apical spot on others

9b.

Most of apical elytra margin lacking any marking

11

or only anterior apical spot present with a short

marginal extension reaching no more than half way to suture 10a. (9a.)

Labrum of most specimens pale tan on each

Ten-spotted Tiger

side of a dark, brownish black middle and all

Beetle, Cylindera decempunctata

margins darkened, labrum on a few specimens entirely darkened; elytra spots large; humeral and apical markings as single bands on some

specimens; apical marginal extension broad

and reaching suture; pronotum laterally with several irregular rows of flattened setae; female elytra with slight lateral expansion at middle

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

10b.

11a. (9b.)

Shade-loving Tiger

Labrum pale tan on each side of a dark, brownish black middle, all margins darkened; lateral elytra margin with a wide copper-green band inside of a broad and purple band extending from outer humeral angle to suture, encompassing much of apical spot

Violet-sided Tiger

Beetle,

Cylindera anelia

Beetle, Cylindera limitisca

11b.

Labrum entirely darkened, nonmetallic shiny brown or black; lateral elytra margin with only a diffuse copper-green humeral area; purple margin diffuse, narrowly reaching suture, not encompassing apical spot

12

12a. (11b.)

Pronotum entirely devoid of setae; legs pale, femurs nearly translucent with a pale brown stripe only along anterior margins; only on Great Nicobar Island, India

Striped-legged Tiger

12b.

Pronotum with many flattened setae along entire lateral margins; legs dark, femurs entirely opaque with metallic colours; found on continental India

13

13a. (12b.)

Entire outer edge of pronotum with many flattened setae coming from two to several nearly regular rows of large, closely spaced punctures; middle elytra band complete, only slightly oblique and narrowed at middle; small, moderately deep punctures on elytra centre, especially around terminus of middle band

13b.

168

Labrum entirely brownish black on most specimens, two pale tan areas on a few specimens; elytra spots small surrounded by a darkened area; humeral marking divided into two spots; apical marking as a single band on a few specimens; marginal extension of apical marking narrow and on most specimens not reaching suture; pronotum laterally bare or with one irregular row of flattened setae; female elytra with wide lateral expansion at middle

Entire outer edge of pronotum with sparse flattened setae coming from one to two irregular rows of small, widely spaced punctures; middle elytra band on most specimens represented by two irregularly shaped spots, medial one situated posteriorly, on some specimens joined by a thin oblique line; elytra centre with minute, shallow punctures, area around terminus of middle band nearly smooth

Beetle,

Cylindera dartista

Stream Tiger Beetle, Cylindera subtilesignata

Black-lipped Tiger Beetle,

Cylindera spinolae

Species Accounts

14a. (8b.)

Lateral elytra margins glossy black to purple­ black with colour bulging toward centre

14b.

Purple-footed Tiger

Beetle, Cylindera umbropolita

Lateral elytra margins shiny iridescent purple,

15

blue or copper, either abruptly or gradually

contrasting with colour at centre

15a. (14b.)

Elytra either with small, narrow moderately

16

impressed iridescent punctures, or surface nearly smooth, especially on rear third 15b.

Elytra with wide, shallowly impressed iridescent

Shy Tiger Beetle,

punctures many of which coalesce, especially

Cylindera viduata

on rear third 16a. (15a.)

Front margin of pronotum with curved ridges

Hump-headed Tiger

in the centre and merging with those on the

Beetle, Cylindera

disc such that middle line extends onto anterior

cydobregma

reflexed margin, elytra laterally shiny black lacking a contrasting metallic margin or humeral area, elytra surface dull, with numerous, smooth

areas, slightly shiny and mostly on apical third 16b.

Anterior reflexed margin of pronotum medially with transverse

ridges

not

17

merging with

those on the disc, middle line of pronotum

not extending onto anterior reflexed margin,

elytra laterally metallic, margin and humeral

area diffusely iridescent on some specimens,

abruptly contrasting in colour with remainder of surface on other specimens, elytra surface with only punctate areas on apical third 17a. (16b.)

Elytra shoulders with a distinctive iridescent blue-green

crescent

ending

abruptly

18

and

contrasting with dull, nonmetallic, dark colour

laterally and at centre, lateral edge of elytra

narrowly or not at all metallic 17b.

Elytra shoulders of some specimens with an

20

inconspicuous diffuse iridescent area lacking a distinct end and gradually merging with shiny,

metallic lateral margin and nonmetallic centre, elytra of other specimens without any diffuse

iridescence at shoulders, lateral edge of elytra

broadly and diffusely metallic

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

18a. (17a.)

Pronotum

lateral

margins with

numerous,

flat setae from small, submarginal punctures

in

multiple rows, front of head coarsely,

Deceptive Tiger

Beetle,

Cylindera fallaciosa

transversely ridged at middle; labrum with a large, broad central ridge down its middle 18b.

Pronotum lateral margins with sparse, flat setae

19

from small submarginal punctures or bare, front

of head finely, transversely ridged at middle; labrum with a thin, narrowly raised central ridge on some specimens, other specimens without

any central ridge 19a. (18b.)

Pronotum sides with sparse flattened hairs

Channel-headed

originating from a few to many small, irregularly

Tiger Beetle,

arranged punctures; elytra dark, copper-brown,

Cylindera collicia

with

numerous contrasting small, shallow,

blue-green punctures on back third; labrum

metallic copper, middle ridge thin, narrowly raised and darkened by extension of marginal

pigmentation along midline 19b.

Pronotum sides bare, a few inconspicuous hairs

Rough Tiger Beetle,

on a few specimens; elytra black, nearly smooth

Cylindera severini

on back third, either with noncontrasting

minute punctures or small iridescent flecks; labrum metallic copper, lacking a middle ridge and darkened extension of marginal pigment

along midline

20a. (17b.)

Elytra with four spots, most large and elongate

21

with middle two on some specimens narrowly

joined, surface uniformly punctured, pronotum laterally with numerous large setae bearing

punctures often occurring in multiple rows 20b.

Elytra of most specimens with three or fewer

Tiger Beetle,

few specimens), surface in its middle with a dull

Cylindera belli

longitudinal area surrounding a small central

spot and with distinctly fewer punctures than remainder of surface, pronotum laterally varied, bare or with sparse appressed setae originating

from a few to many irregularly arranged

punctures

170

Shiny-shouldered

small oval spots (minute fourth dot on only a

Species Accounts

Body dull, dark purple-black dorsally, elytra with four nearly circular spots alternating between medial and lateral positions from anterior to posterior, elytra surface with small, noncontrasting punctures shallow basally becoming nearly smooth apically

Four-dotted Tiger

21b.

Body shiny, copper, copper-green or green dorsally, elytra with whitish yellow spots, only anterior spot consistently circular, others irregular in shape, elytra surface with large, contrasting blue-green punctures

22

22a. (21b.)

Elytra surface covered with large punctures and small sunken areas

Karl's Tiger Beetle,

22b.

Elytra surface with large, contrasting blue-green punctures deepest basally becoming shallower toward tips

23

23a. (22b.)

Wide ridges on top of head, 8 to 11 nearly complete ridges between eye and midline, female with side of thorax below shoulder of elytra with a middle groove or small pit along back margin, occurs in northern half of India

Green-lipped Tiger

21a. (20a.)

23b.

Narrow ridges on top of head, 12 to 16 nearly complete ridges between eye and midline, female with side of thorax below shoulder of elytra with a large, deep cavity at middle along back margin, occurs only in southern India

Beetle, Cylindera nietneri

Cylindera karli

Beetle, Cylindera viridilabris

Copper-lipped Tiger Beetle,

Cylindera labioaenea

24a. (7b.)

Cheeks on most specimens with flat hairs, on some inividuals sparse hairs only near bottom, on other individuals hairs missing, if cheeks lack hairs then elytra with long, narrow macula at sides and second small segment of middle legs lack a subapical seta

25

24b.

Cheeks entirely bare, elytra lack a long, narrow macula at margin, second small segment of middle legs always with a subapical seta

28

25a. (24a.)

Labrum short, transverse, not projecting forward, small elytra markings, elytra surface smooth and evenly contoured

26

25b.

Labrum long, projecting forward, elytra surface unmarked, rough and unevenly contoured

Pitted Tiger Beetle,

Cylindera foveolata

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

26a. (25a.)

Elytra marking at shoulder divided into two dots

White-margined

Tiger Beetle, Cylindera kaleea 26b.

Elytra marking at shoulder either missing or

27

only a front dot

27a. (26b.)

Elytra without any marking at shoulder, two

Small-lipped Tiger

small spots at middle of elytron, one marginal

Beetle,

and one in the middle, found in northeastern

Cylindera sikhimensis

India 27b.

Elytra marked with a small basal dot at shoulder,

one oval spot at middle of elytron, only on Great

Beetle, Cylindera discreta

Nicobar Island, India 28a. (24b.)

Dark-spotted Tiger

Elytra with spot at shoulder, middle band

Brown-lipped Tiger

oblique or transverse, enlarged at margin and

Beetle,

extending as a narrow or slightly enlarged

Cylindera paudpilina

marking onto centre, apical marking a long spot

touching margin

28b.

Elytra without spot at shoulder, three spots

29

aligned in a row on centre of each elytron

29a. (28b.)

Labrum pale yellow on middle, metallic blue­

Pit-lined Tiger

green on each lateral quarter, six to eight sensory

Beetle, Cylindera

bristles on labrum, eyes small, not prominent,

seriepunctata

numerous moderately deep punctures on elytra centre contrasting with surface colour

29b.

Labrum entirely pale and tawny, eight to ten sensory bristles on labrum, eyes large,

prominent,

elytra

centre

nearly

without

Honey-faced Tiger

Beetle,

Cylindera melitops

punctures, shallow non-contrasting punctures

only at elytra base

30a. (1b.)

Head, pronotum, all sides of the thorax and

Mountain

abdomen, entirely covered by long flattened

Stream Tiger

hairs, also hooked hairs on the longest leg

Beetle, Cylindera

segments, elytra markings form a divided

albopunctata

pattern of four spots along outer edges and two

oval spots at centres, occurs in northern India

30b.

Body never entirely covered with long flattened hairs, elytra markings never forming isolated

spots both along edges and at centre, not restricted to northern India 172

31

Species Accounts

31a. (30b.)

Head bare below eyes on cheeks and above

32

labrum 31b.

32a. (31a.)

Head with sparse flattened hairs below eyes on

Ragged Tiger Beetle,

cheeks and above labrum

Cylindera sublacerata

Elytra at shoulder unmarked

Bar-shouldered

Cylindera singalensis 32b.

33a. (31b.)

Elytra at shoulder marked with band or spots

33

Elytra middle band on descending portion parallel to suture with very irregular and

Fragmented-

band Tiger Beetle, Cylindera

fragmented edges along inner margins

mesoepisternalis 33b.

Elytra middle band with descending portion

34

parallel to suture with smooth and wavy edges

along inner margins

34a. (33b.)

Elytra band at shoulder divided into two dots

35

34b.

Elytra band at shoulder continuous, not divided

37

35a. (34a.)

Elytra

band

along

outer

rear

margin

36

incompletely formed, marginal portion narrow,

either long or short, and separated from

rounded inner spot 35b.

Elytra band along outer rear margin completely

Dwarf Tiger Beetle,

formed, marginal portion widened at each end

Cylindera brevis

and broadly connected to large rounded inner

spot or both each end narrowly separated 36a. (35a.)

Pronotum at sides with sparse flattened hairs

Slender Tiger Beetle,

on bottom three-quarters, few to no setae on

Cylindera procera

upper quarter; marginal portion of rear elytra

band very short, reduced to a short elongate

spot restricted to outer rear angle; fourth male antennal segment with only a few thin,

hairs 36b.

Pronotum

at

sides

uniformly covered

by

Clubbed Tiger

moderate to abundant flattened hairs; marginal

Beetle,

portion of rear elytra band long, extending from

Cylindera bigemina

outer rear angle to suture; fourth male antennal

segment with clubbed comb of wide setae

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

37a. (34b.)

Mandibles with a long terminal tooth equal or

38

exceeding half its total length 37b.

Mandibles with a short terminal apical tooth

39

less than half its entire length

38a. (37b.)

Elytra band at tip a single continuous marking

Long-toothed Tiger Beetle,

Cylindera cognata 38b.

Elytra band at tip broken into two dots

Erudite Tiger Beetle,

Cylindera erudita 39a. (37b.)

Elytra shoulder band meets middle band by a

40

marginal line

39b.

Elytra shoulder band separated from middle

42

band

40a. (39a.)

Elytra middle band at its inner end extending

Wavy-lined Tiger

posteriorly along suture nearly to apical band;

Beetle,

anterior portion of apical band long, slightly

Cylindera venosa

sinuate, extending more than halfway toward transverse portion of middle band

40b.

Elytra middle band at its inner end notextending

41

along suture posteriorly; anterior portion of apical band short, recurved, extending medially

for only a short distance

41a. (40b.)

Elytra with anterior band abruptly hooked at its

tip on middle of surface

Fish Hook Tiger

Beetle, Cylindera ancistridia

41b.

Elytra with anterior band gradually recurved at

its tip on middle of surface

42a. (39b.)

Elytra shoulder band projecting on to middle of

surface and ending in a bulge

Dune Tiger Beetle, Cylindera agnata

Mudflat Tiger Beetle, Cylindera

grammophora

42b.

174

Elytra shoulder band projecting only slightly on

Little Tiger Beetle,

to middle of surface and not ending in a bulge

Cylindera minuta

Species Accounts

Dot-lipped Tiger Beetle, Cylindera paradoxa (Horn) (Plate 24) [Map 184] Identification: Length 7.5 mm; head and pronotum bright green and violet reflections on the sides; elytra shiny dark bronze to black

with longitudinal row of green pits on the surface and two oval white spots on rear half of each elytron. Male labrum with large white centre; body entirely without hair-like setae.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Southern India and Sri Lanka. Floor of open forest, brushy areas, fallow fields, and road cuts (0 - 650 m).

Behaviour: Rarely flies, but it is attracted to lights at night.

184 Dot-lipped Cylindera paradoxa.

Map

Tiger

Beetle,

Seasonality: February to October.

Fluted Tiger Beetle, Cylindera lacunosa (Putzeys)(Plate 24) [Map 185] Identification: Length 7.5 - 9 mm; bronze; elytra with polished raised areas and two

oval white spots on rear half.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Southern India and Sri Lanka. Coastal forest openings. Behaviour: Feeble flier and occasionally attracted to lights at night. Seasonality: May to November.

Map 185 Fluted Tiger Beetle, Cylindera lacunosa.

Slope-striped Tiger Beetle, Cylindera obliquefasciata (Adams) (Plate 24) [Map 186] Identification: Body small to medium (9-11 mm); head and pronotum metallic copper-green with greenish reflections; elytra dull black-green, each with humeral band divided into two small spots, one at front angle, one at middle, a separate middle band wide and distinctly descending inward from near lateral margin onto

175

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

across the middle, and a complete apical band with forward pointing end enlarged; body copper-green ventrally with sparse, flattened hairs on sides, but surface bare in middle.

Subspecies and morphological variants: Two subspecies have been described for this widespread species, but only C. o. descendens occurs in India.

Distribution and habitats: The subspecies C. o. descendens occurs in extreme northern India in Kashmir west to Uzbekistan and Russia and east to China. Open grassland and on moist soils (1500 m).

Map

186 Slope-striped Cylindera obliquefasciata.

Tiger

Beetle,

Behaviour: Larvae and adults occur on floor of valley forests near large rivers or in meadow terraces in moist loam soils.

Seasonality: May to June.

Fleet Tiger Beetle, Cylindera dromicoides (Chaudoir) (Plate 24) [Map 187] Identification: Length 11-12 mm; head and pronotum bright copper to dull brown or green; elytra dark velvety brown patches surrounded by bright bronze on shoulders

outer edge and band down length of the inner edges, two elongate white spots at mid-margin and rear, elytral sides peculiarly

widened at rear. Somewhat similar to the Terrestrial Tiger Beetle (C. delavayi), but larger

body and different-shaped elytra distinguish it.

Subspecies and morphological variants: No subspecies have been described, but larger individuals tend to occur at higher elevations, and at some sites the head and pronotum are green, blue or purple.

Map 187 Fleet Tiger Beetle, Cylindera dromicoides.

Distribution and habitats: Northern and northeastern India, Nepal, Bhutan, and China. Floor of oak woodlands to fir forests (1000 - 3700 m).

Behaviour: Although this species has wings, it rarely flies. Runs quickly along clay trails and open areas in the leaf litter. Often hides under dead leaves on the ground.

Seasonality: May to September.

176

Species Accounts

Terrestrial Tiger Beetle, Cylindera delavayi (Fairmaire) (Plate 24) [Map 188] Identification: Length 7 - 10 mm; head and pronotum copper or green; elytra dull bronze to blue or green with two velvety dark areas

and two small yellow spots on rear margin. Sides of elytra are parallel. Somewhat similar to the Fleet Tiger Beetle (C. dromicoides), but

smaller body and different-shaped elytra

distinguish it.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Uttarakhand to northeastern

India, Bhutan, Nepal, China,

Vietnam, Laos, Thailand, and Myanmar. Forest paths, dirt roads in moist forests (500 - 2900 m).

Map 188 Terrestrial Tiger Beetle, Cylindera delavayi.

Behaviour: Although it has wings, it rarely flies. Also occurs in isolated patches within what appears to be continuous habitat.

Seasonality: April to October.

Pitted Tiger Beetle, Cylindera foveolata (Schaum) (Plate 24) [Map 189] Identification: Length 8 mm; head shiny black above and blue-green on the sides; pronotum shiny black above and copper to blue-green on the sides; elytra shiny blackish bronze with row of blue-green to copper pits running along inner edge,

surface uneven forming an irregular pattern, no light markings.

Subspecies and morphological variants: No subspecies have been described for this widespread species.

Distribution and habitats: Widespread across India except desert northwest, Map 189 Pitted Tiger Beetle, Cylindera Bangladesh, Nepal, Myanmar, Thailand, foveolata. Laos, Cambodia, Vietnam, China, Indonesia, and Philippines. Open areas of mature forest, road cuts and during heavy monsoons moves out into scrub forest floor (150-1400 m).

Behaviour: Often hides in leaf litter to escape danger. Seasonality: April to October. 177

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Hump-headed Tiger Beetle, Cylindera cyclobregma (Acciavatti and Pearson) (Plate 24) [Map 190] Identification: Length 7.3 mm; head and pronotum black and head with distinctive bump between the eyes; elytra dull back

with shiny back margin and with one small

white dot close to the inner edge, elytra noticeably expanded toward the rear.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India, Myanmar, Laos, and Vietnam. Habitat

undescribed, but occurs at altitudes from 900 to 1000 m.

Behaviour: Unknown.

Map 190 Hump-headed Tiger Beetle,

Seasonality: April to November.

Cylindera cyclobregma.

Shy Tiger Beetle, Cylindera viduata (Fabricius) (Plate 24) [Map 191] Identification: Length 7.5 - 8.5 mm; head and pronotum copper-green above and

blue-green on the sides; elytra dull greenish

black to copper-black with a narrow velvety black stripe running along the inner edge, three small white spots on the rear half of each elytron.

Subspecies and morphological variants: No subspecies have been described because this widespread species has a surprisingly uniform appearance throughout its vast geographical range.

Distribution and habitats: Northern and

Map 191 Shy Tiger Beede, Cylindera viduata.

northeastern India east to Philippines, Papua New Guinea, and China. Common in a wide range of disturbed habitats including forest paths, fields, soccer pitches, and along lake and river edges (120-1400 m).

Behaviour: Flies relatively long distances when disturbed and is attracted to lights at night.

Seasonality: April to September. 178

Species Accounts

Karl's Tiger Beetle, Cylindera karli Cassola (Plate 24) [Map 192] Identification: Length 6.5 - 8 mm; head bronze with green and reddish reflections;

pronotum bronze with coppery reflections; elytra reddish bronze with a green band along

outer edge and surface covered with green punctures throughout, four small whitish spots and ovals on rear half of each elytron. Similar to the Ten-spotted Tiger Beetle (C decempunctata), the Green-lipped Tiger Beetle (C viridilabris), and the Black-lipped Tiger Beetle (C. spinolae), but distinguished from all of them by the extensive green punctures across the surface of the elytra.

Subspecies and morphological variants:

Map 192 Karl’s Tiger Beetle, Cylindera karli.

No subspecies have been described.

Distribution and habitats: Highly localized and endemic to southeastern India in Andhra Pradesh.

Behaviour: Unknown.

Seasonality: July.

Green-lipped Tiger Beetle, Cylindera viridilabris (Chaudoir) (Plate 25) [Map 193] Identification: Length 6.5 - 8 mm; head and pronotum dark bronze above and shiny blue-green on the sides; elytra dark brown or black with metallic blue-green punctures, shiny areas at the shoulder and along the sides, four thin to small yellowish spots and lines on each elytron; labrum copper green. Similar to the Copper-lipped Tiger Beetle (C. labioaenea), which is distinguished by its copper-bronze coloured labrum and more southern geographical range.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern India, Pakistan,

Nepal,

Thailand,

Map

193 Green-lipped Cylindera viridilabris.

Tiger

Beetle,

Bangladesh,

Cambodia, and China. Moist shrubby parks and forested areas (200 - 2000 m).

Behaviour: Attracted to lights at night. Seasonality: May to July.

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Copper-lipped Tiger Beetle, Cylindera labioaenea (Horn) (Plate 25) [Map 194] Identification: Length 7-8 mm; head and pronotum shiny copper and blue-green

above and bright blue and green on the sides; elytra dull copper-green to green with shiny blue green at shoulder and four small yellowish spots on each elytron; labrum

metallic bronze to copper. Similar to the Green-lipped Tiger Beetle (C. viridilabris), which

is recognized by its green-coloured labrum and more northern geographical range.

Subspecies and morphological variants: Two subspecies have been described, but only C. /. fuscocuprascens, with its copper

segments on the underside of the thorax, has been found in India.

Map 194 Copper-lipped Tiger Beetle, Cylindera labioaenea.

Distribution and habitats: Southern India. Floor of moist scrub to mature forests (350 - 600 m).

Behaviour: Attracted to lights at night.

Seasonality: April to September.

Deceptive Tiger Beetle, Cylindera fallaciosa (Horn) (Plate 25) [Map 195] Identification: Length 7-8 mm; head and pronotum copper above and shiny blue-green and violet on the sides; elytra

dull dark brown to black with shiny blue­

green crescent on the shoulders and four small whitish dots on rear two-thirds of each elytron. Similar to the Copper-lipped Tiger Beetle (C. labioaenea) from which it is distinguished by the shiny crescents on the shoulders of the elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India, Bangladesh, Myanmar, Thailand, Laos, Vietnam, and China. Paths and openings in forests.

Behaviour: Attracted to lights at night. Seasonality: April to August. 180

Map 195 Deceptive Tiger Beetle, Cylindera fallaciosa.

Species Accounts

Rough Tiger Beetle, Cylindera severini (Horn) (Plate 25) [Map 196] Identification: Length 7-8 mm; head and pronotum shiny copper above and purple to blue-green on the sides; elytra dull brownish black except for a shiny blue green crescent on the shoulders. Similar to the Deceptive Tiger Beetle (C fallaciosa), but separated by

geographical range.

Subspecies and morphological variants: Some males are all black (melanistic).

Distribution and habitats: Endemic to peninsular India. Forest paths and roads (20 -1050 m).

Behaviour: Attracted to lights at night.

Map 196 Rough Tiger Beetle, Cylindera

Seasonality: May to July.

severini.

Channel-headed Tiger Beetle, Cylindera collicia (Acciavatti and Pearson) (Plate 25) [Map 197] Identification: Length 7.5 - 9.5 mm; head and pronotum copper brown above with metallic green on the sides; elytra dull dark copper-brown with iridescent blue­ green crescent at shoulders and four small whitish spots on each elytron. Similar to the Rough Tiger Beetle (C. severini) and the Green-lipped Tiger Beetle (C. viridilabris) but

separated by geographical range.

Subspecies and morphological variants: At one time considered to be only a local form of the Green-lipped Tiger Beetle (C.

virdilabris), but subtle differences in female coupling sulcus, sculpture of the head, and

patterns of punctures on the elytra have been used to separate it as a full species.

Map 197 Channel-headed Tiger Beetle, Cylindera collicia.

Distribution and habitats: Endemic to southern portion of the Western Ghats in southern India. Moist forest floor and road cuts (150 - 900 m). Behaviour: Attracted to lights at night. Seasonality: April to July. 181

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Four-dotted Tiger Beetle, Cylindera nietneri (Horn) (Plate 25) [Map 198] Identification: Length 7 - 7.5 mm; head and pronotum dull purple black above and shiny purple-green on the sides; elytra dull purple

black with four small whitish spots on each elytron. Very similar to the Copper-lipped Tiger Beetle (C. labioaenea) and may prove

to be a melanic form of its subspecies C. /.

fuscocuprascens.

Subspecies and morphological variants: Only males of this rare form have been found, and its status as a full species needs more information on specimens over a wider geographical area.

Distribution and habitats: In India only

Map U ___ 198 Four-dotted Cylindera nietneri.

Tiger

Beetle,

known from a single specimen in Andhra Pradesh; also occurs in Sri Lanka. Moist habitats.

Behaviour: Unknown. Seasonality: August.

Shiny-shouldered Tiger Beetle, Cylindera belli (Horn) (Plate 25) [Map 199] Identification: Length 7-8 mm; head and pronotum shiny bronze black to coppergreen above and blue green on the sides; elytra shiny bronze-black with a dull black area in the middle and metallic copper to

blue shoulder and sides, three whitish spots on each elytron.

Subspecies and morphological variants: Blackish elytra in the northern part of its

range and copper to copper-green elytra in

the south.

Distribution and habitats: Endemic to peninsular India. Moist shade in bushy areas and more open adjacent sites in later monsoon (200-1500 m).

Behaviour: Unknown. Seasonality: May to July. 182

Map 199 Shiny-shouldered Tiger Beetle, Cylindera belli.

Species Accounts

Purple-footed Tiger Beetle, Cylinders umbropolits (Horn) (Plate 25) [Map 200] Identification: Length 7.5 - 8 mm; head and pronotum copper to blue green above and blue green on the sides; elytra shiny copper

with dull black areas, shiny black edges and one or two small whitish spots on each elytron; lower legs shiny purple.

Subspecies and morphological variants: Two subspecies have been described:

C. u. umbropolita - narrow shiny black edges to the elytra.

C. u. lucidinigrosa - wide shiny black edges that extend into the centre of each elytron.

Distribution and habitats: shaded forest floor: - MaP 200 Purple-footed Tiger Beetle, -

,

.

.

Cylindera umbropolita; A, C. u. umbropolita,

r

C. u. umbropolita - southern portion of Western Ghats in South India (800 - 1700 m).

B> c. u. lucidinigrosa.

C. u. lucidinigrosa - central portion of Western Ghats (100 - 650 m) in South India. Intergrades predominate in central Karnataka.

Behaviour: Unknown.

Seasonality: May to July.

Pit-lined Tiger Beetle, Cylinders seriepunctsts (Horn) (Plate 26) [Map 201] Identification: Length 7.5 - 8.5 mm; head and pronotum green bronze to bronze above and metallic green on the sides; elytra dull green bronze to bronze with shiny blue-green edges, deep blue

punctures and three whitish spots on each elytron; labrum pale brown in the centre and blue-green on the sides.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India, Nepal, and Bhutan. Sand and gravel

along mountain streams and occasionally adjacent open forests and agricultural

Map 201 Pit-lined Tiger Beetle, Cylindera seriepunctata.

fields (180-2000 m).

Behaviour: Unknown. Seasonality: April to July.

183

A. Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Honey-faced Tiger Beetle, Cylindera melitops (Acciavatti and Pearson) (Plate 26) [Map 202] Identification: Length 8 mm; head and pronotum shiny green bronze to bronze

above and metallic green on the sides; elytra dull green bronze to bronze with no punctures, shiny and broad blue green

along edges, and three whitish spots on each elytron; labrum all pale brown. Similar to the Pit-lined Tiger Beetle (C seriepuntata), which has a pale brown labrum with metallic blue-green on its sides.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to northeastern India in Arunachal Pradesh.

202 Honey-faced Cylindera melitops.

Map

Tiger

Beetle,

Tiger

Beetle,

Stream edges (200 - 350 m).

Behaviour: Unknown. Seasonality: May to June.

Black-lipped Tiger Beetle, Cylindera spinolae (Gestro) (Plate 26) [Map 203] Identification: Length 7-8 mm; head and pronotum bright copper above and blue­ green on the sides; elytra dull brownish with greenish inner edges and five or more

small circular and irregular whitish markings on each elytron; labrum completely shiny brownish black. Similar to Brown-lipped Tiger Beetle (C. paucipilina), which is smaller, has

a

lighter-coloured

labrum,

and

a

complete middle line on the elytron.

Subspecies and morphological variants: Some individuals uniformly dull brownish

(nearly melanic). Another subspecies has been described from Thailand.

Map

203 Black-lipped Cylindera spinolae.

Distribution and habitats: Northern India, Nepal, Bangladesh, Myanmar, Thailand, Laos, Cambodia, Vietnam, and China. Floor

of moist forests (120-1600 m).

Behaviour: Often flies up to land on leaves of undergrowth plants when disturbed. Seasonality: April to October. 184

Species Accounts

Striped-legged Tiger Beetle, Cylindera dartista Acciavatti and Pearson (Plate 26) [Map 204] Identification: Male body small (7.1-7.4 mm), a very distinctive species with pale tawny legs with longest segments nearly translucent except for a brown stripe on anterior margins, elytra marked with three separate markings, front and back ones completely formed, middle one divided into irregularly shaped spots. This species was discovered recently on Great Nicobar Island in the eastern side of the Bay of Bengal. Its closest relatives, Black-lipped Tiger Beetle (C. spinolae) and Brown-lipped Tiger Beetle (C. paucipilina), from the adjoining mainland of Southeast Asia, both have elytra with incomplete front and rear markings, but a complete but narrow middle band.

Map 204 Striped-legged Tiger Beetle, Cylindera dartista.

Subspecies and morphological variants: None known. Distribution and habitats: Known only from Campbell Bay, Great Nicobar Island. Adults apparently inhabit moist jungle forests.

Behaviour: Runs on the forest floor and flies to leaf surfaces of low shrubs. Seasonality: May.

Brown-lipped Tiger Beetle, Cylindera paucipilina (Acciavatti and Pearson) (Plate 26) [Map 205] Identification: Length 6-8 mm; head blue­ green and purple with a narrow copper area between the eyes; pronotum shiny golden copper on top and blue-green on sides; elytra dull brownish with copper green and purple bands running along outer edge, three whitish spots and a thin line running obliquely across each elytron; labrum pale brown. Similar to the Black-lipped Tiger Beetle (C. spinolae), which is

larger, has darker labrum, and an incomplete middle line on the elytron.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Forest floor (900

Map 205

Brown-lipped Tiger Cylindera paucipilina.

Beetle,

-1100 m) in Northeastern India, Bangladesh,

Myanmar, Thailand, Laos, and China. Forest floor (900 - 1100 m).

Behaviour: Unknown.

Seasonality: May to November.

185

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Violet-sided Tiger Beetle, Cylindera limitisca (Acciavatti and Pearson) (Plate 26) [Map 206] Identification: Length 8 -10 mm; head blue­ green and purple with copper between eyes; pronotum shiny golden copper above and

blue-green on the sides; elytra dull brownish with copper band and wide purple strip along outer edge, copper-green along inner edge, three whitish spots and line connecting two spots in middle; labrum dark brownish black with pale brown areas on the sides. Similar to the Black-lipped Tiger Beetle (C. spinolae),

which is larger, has darker labrum, narrow purple band along outer edge of elytra, and an incomplete middle line on the elytron.

Subspecies and morphological variants: No subspecies have been described.

Map

206 Violet-sided Cylindera limitisca.

Tiger

Beetle,

Distribution and habitats: Endemic to northeastern India. Forest floor (200 - 330 m) in

Behaviour: Unknown. Seasonality: May to June.

Stream Tiger Beetle, Cylindera subtilesignata (Mandi)

(Plate 26) [Map 207]

Identification: Length 7-8 mm; head and pronotum copper above and blue-green on

the sides; elytra dull, dark bronze with copper shoulder, shiny blue to green on outer and

inner edges and whitish double spot at shoulder, complete but narrow middle band

and small spot at rear; labrum dark brownish black. Similar to the Black-lipped Tiger Beetle (C. spinolae), the Ten-spotted Tiger Beetle (C. decempunctata), the Violet-sided Tiger Beetle (C. limitisca), and the Brown-lipped Tiger Beetle (C. paucipilina), but is separated

from these others by female coupling sulcus shape, labrum colour, width of purple band

along elytra outer edge, and body size.

Map 207 Stream Tiger Beetle, Cylindera subtilesignata.

Subspecies and morphological variants: No subspecies have been described. Distribution and habitats: Endemic to northern India. Exposed rocks and boulders in mountain streams and adjacent forest floor during high water (450 - 3600 m).

Behaviour: Attracted to lights at night. 186 Seasonality: April to July.

Species Accounts

Ten-spotted Tiger Beetle, Cylindera decempunctata (Dejean) (Plate 26) [Map 208] Identification: Length 9-10 mm; head and pronotum copper to copper green above and blue-green on the sides; elytra dull,

dark copper green or brown with shallow blue punctures and five small yellowishwhite spots on each elytron; labrum usually nonmetallic dark brown in the centre and

pale brown on the sides.

Subspecies and morphological variants: The markings on the elytra are broader and sometimes connected in northern India.

Distribution and habitats: Northern and northeastern India, Pakistan, Nepal, Bangladesh,

Myanmar,

Thailand,

Laos,

Map

208 Ten-spotted Cylindera decempunctata.

Tiger

Beetle,

Vietnam, and Cambodia. Openings in secondary forest and shrubby areas (100-1600 m).

Behaviour: Often attracted to lights at night.

Seasonality: May to October.

Shade-loving Tiger Beetle, Cylindera anelia (Acciavatti and Pearson) (Plate 26) [Map 209] Identification: Length 8 - 10 mm; head and pronotum copper above and purple to blue­ green on the sides; elytra dull, dark copperbrown with five whitish spots on each elytron (shoulder spot often hard to see),

and female elytra distinctly widened in the

middle; labrum usually all dark. Similar to the Ten-spotted Tiger Beetle (C. decempunctata), which has a bi-coloured labrum, the female has more parallel sided elytra, and habitat/ distribution does not overlap.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Endemic to ,

...................................................

.

7209 Cylindera anelia.

Shade-loving

Tiger

Beetle,

southwestern India. Heavily shaded and moist forest floor, often near streams (100 - 650 m).

Behaviour: Unknown.

Seasonality: May to July. 187

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Small-lipped Tiger Beetle, Cylindera sikhimensis (Mandi) (Plate 27) [Map 210] Identification: Length 7-8 mm; head and pronotum dark bronze-green above and shiny green on the sides; elytra dull bronze

with three whitish spots on the rear half of

each elytron; labrum light brown.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India and Myanmar. In open areas and along

roads (150-950 m).

Behaviour: Unknown. Seasonality: April to July.

Map

210 Small-lipped Cylindera sikhimensis.

Dark-spotted Tiger Beetle, Cylindera discreta (Schaum

Tiger

Beetle,

(Plate 27) [Map 211]

Identification: Length 6 mm; body dorsally greenish with head black and green on the

sides; pronotum almost cylindrical with sparse white hairs along edges; elytra blackish brown in the middle with dark diffused area almost covering entire centre and only the side margins bright green; elytra markings a small shoulder dot, a large, circular spot at centre and an apical band with its anterior branch only slightly widened.

Subspecies and morphological variants: Numerous subspecies have been described

based primarily on differences in their markings. Although these have been assigned to this species in the past, most have been elevated to

Map

211 Dark-spotted Cylindera discreta.

Tiger

Beetle,

species because subsequent study revealed adults exhibited distinctly different features across widely isolated locations, including Southeast Asia, the Philippines, Indonesia, and

Australia. Currently three subspecies are provisionally recognized. However, to date, only C. d. nicobarica, has been documented from India on Great Nicobar Island.

Distribution and habitats: The most widespread subspecies, C. d. discreta, occurs on Indonesia on Sumatra, less than 200 km from Great Nicobar Island. There is a

chance it could eventually be found on Great Nicobar Island, but presently C. d. nicobarica is the only subspecies known from India and is apparently endemic to this island. Only one specimen is known.

Behaviour: Unknown.

188 Seasonality: March.

Species Accounts

White-margined Tiger Beetle, Cylindera kaleea (Bates) (Plate 27) [Map 212] Identification: Length 9 - 10 mm; head and pronotum shiny copper above and metallic blue-green on the sides; elytra dull brown

to blackish with four whitish spots on rear

half of each elytron; labrum non-metallic and ranging in colour from entirely dark to entirely pale.

Subspecies and morphological variants: The size, shape and number of light markings on the elytra and the colour of the labrum vary throughout its range; three additional subspecies have been described from China and Japan.

Distribution and habitats: Forest paths,

Map 212 White-margined Tiger Beetle, Cylindera kaleea.

along lakes and rivers and in open areas of villages (350 - 1400 m) in northern India (Assam, Arunachal Pradesh, Meghalaya, Sikkim, Uttarakhand), Myanmar, Thailand, Laos, Vietnam, China, and Taiwan.

Behaviour: Unknown.

Seasonality: May to July.

Mountain Stream Tiger Beetle, Cylindera albopunctata (Chaudoir) (Plate 27) [Map 213] Identification: Length 10-11 mm; olive green to dull brown green above; elytra shiny

with wide blue punctures and six pale yellow markings on each elytron; underside of body heavily covered with white hair-like setae.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern and northeastern India, Pakistan, Nepal, Bhutan, and China. Boulders and exposed bare areas

along mountain streams (350 - 4000 m) in

Behaviour: Often fly from boulder to boulder along mountain streams. Attracted to lights at night.

Map 213 Mountain Stream Tiger Beetle, Cylindera albopunctata.

Seasonality: May to October. 189

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Ragged Tiger Beetle, Cylindera sublacerata (Solsky)

(Plate 27) [Map 214]

Identification: Length 7.5 - 8 mm; head and pronotum shiny green to copper-green; elytra

dull green to copper-green with purple to

blue punctures and yellowish-white shoulder

crescent, curved middle band and rear crescent.

Subspecies and morphological variants: Four

additional

subspecies

have

been

described from across southwest northward into Russia and China.

Asia

Distribution and habitats: Extreme northern India in Kashmir east to Russia and Kazakhstan. Open sandy beaches along lakes and rivers.

Behaviour: Larvae occur in sandy or loam sandy soils near rivers, lakes, and ponds.

Map 214 Ragged Tiger Beetle, Cylindera sublacerata.

Seasonality: May to August.

Fragmented-band Tiger Beetle, Cylindera mesoepisternalis (Horn) (Plate 27) [Map 215] Identification: Length 8 mm; body dorsally copper green, head and pronotum shiny and elytra duller with purple or blue punctures; body underneath along the sides at the

middle with much dense, long setae; elytra with three white bands all connected by a wide continuous lateral margin extending from shoulder to tip at suture, bands project inward with ends of front and rear bands

blunt and broad, ends of middle band sinuate and descending parallel to suture

and fragmented along inner margins. Similar to the Ragged Tiger Beetle, (C. sublacerata), whose middle band is also fragmented, but

Map 215 Fragmented-band Tiger Beetle,

underside of its body has much sparser and Cylindera mesoepistemalis. shorter setae. Similar to the Fish HookTiger Beetle, (C. ancistridia), except its shoulder band forms a distinctive forward bend at its end that looks like a fish hook and the middle band is not fragmented along inner margins near the suture.

Subspecies and morphological variants: None known.

Distribution and habitats: Northern India and Pakistan. The only specimen known from India is a single male labeled Skardo (Kashmir). Habitat unknown.

Behaviour: Unknown. 190 Seasonality: Unknown.

Species Accounts

Clubbed Tiger Beetle, Cylindera bigemina (Klug) (Plate 27) [Map 216] Identification: Length 9-10 mm; head and pronotum copper above and partially

green on the sides; elytra copper-green with thin yellowish line at shoulder and at

rear, a round spot on the front and rear and an elbow-shaped mark in the middle; male antennae with comb of bristles.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Widespread in India, Pakistan, and Nepal. Grassy edges of sandy river banks, rice paddies and adjacent fallow fields and open forests (150-2150 m):

Behaviour: Attracted to lights at night.

Map 216 Clubbed Tiger Beetle, Cylindera bigemina.

Seasonality: May to October.

Slender Tiger Beetle, Cylindera procera (Horn) (Plate 27) Identification: Length 8 - 8.5 mm; head and pronotum shiny copper bronze; elytra dull copper-brown with small green punctures

and small white spots and bands, rear marking reduced to a spot; no comb of bristles *on male antennae. Considered by some experts to be the same species as the Clubbed Tiger Beetle (C. bigemina).

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Known from only two male specimens labeled "India." Habitat unknown. Behaviour: Unknown.

Seasonality: Unknown. 191

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Dwarf Tiger Beetle, Cylindera brevis (Horn) (Plate 28) [Map 218] Identification: Length 7 - 8.5 mm; head and pronotum shiny copper-green; elytra

dull copper-brown to green with purple punctures and whitish markings made up of spots and thin bands; male with no comb of bristles on antennae. Similar to Clubbed Tiger Beetle (C. trigemina), which is larger

and females have different shaped coupling sulcus on thorax.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern India, Afghanistan, and Pakistan. Grassy edges of muddy ponds, reservoirs, and lakes (200 1000 m).

Map 218 DwarfTiger Beetle, Cylindera brevis.

Behaviour: Unknown.

Seasonality: June to October.

Little Tiger Beetle, Cylindera minuta (Olivier) (Plate 28) [Map 219] Identification: Length 7 - 8.5 mm; head and pronotum shiny copper or copper-green above and metallic blue-green on the sides; elytra dull green and copper with three thin ivory-coloured lines at the shoulder, mid­

margin and rear.

Subspecies and morphological variants: Although widespread, no subspecies have been described.

Distribution and habitats: Widespread in India and Nepal east to Philippines. Wide range of muddy and sandy habitats from water's edge to grassy areas (20 - 1000 m).

Behaviour: Often together with the Dune Tiger Beetle (Cylindera agnata) and Sand Tiger Beetle (Calomera angulata).

Seasonality: February to December. 192

Map 219 Little Tiger Beetle, Cylindera minuta.

Species Accounts

Long-toothed Tiger Beetle, Cylindera cognata (Wiedemann) (Plate 28) [Map 220] Identification: Length 9-10 mm; head and pronotum copper above with green reflections on the sides; elytra dark black-green with broad whitish shoulder line "C" shaped

not connected along the edge to the rightangled and sinuate middle band, rear marking

a large dot connected to a band across the rear margin; last tooth on mandible extends almost half the length of the mandible.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Widespread in India, Pakistan, Nepal, and Bangladesh:

Sandy beaches of rivers and adjacent moist

Map 220 Long-toothed Tiger

Beetle,

Cylindera cognata.

grassy areas (500 - 900 m).

Behaviour: Attracted to lights at night. Occurs together with the Erudite Tiger Beetle (C. erudita).

Seasonality: June to October.

Erudite Tiger Beetle, Cylindera erudita (Wiedemann) (Plate 28) [Map 221] Identification: Length 8 - 10 mm; head and pronotum copper above with bright green to purple on the sides; elytra dull deep blue­

black with shoulders and front part shiny

green, white markings broad with shoulder line "C" shaped and connected along the edge to*the right-angled middle band, rear marking divided into two.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern India, Afghanistan, and Nepal. Moist open areas often in grass above sandy riverbanks (350 - 2500 m).

Map 221 Erudite Tiger Beetle, Cylindera erudita.

Behaviour: Attracted to lights at night. Seasonality: May to October. 193

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Dune Tiger Beetle, Cylindera agnato (Fleutiaux) (Plate 28) [Map 222] Identification: Length 9-10 mm; head and pronotum shiny copper-brown above with green reflections on the sides; elytra dull

brown to greenish with yellowish-white lines

along the outer edges and three sinuate lines at the shoulder, mid-margin and rear connected to the edge and extending into the interior of the elytra. Similar to the Fish

Hook Tiger Beetle (C ancistridia}, which has a distinctive fish hook shape to its shoulder

marking, and larger size to distinguish it.

Subspecies and morphological variants: Some variation in size and subtle morphological characteristics of female coupling sulcus.

Map 222 Dune Tiger Beetle, Cylindera agnata.

Distribution and habitats: Widespread in India except central and northwestern parts, Nepal, and Myanmar. Broad sandy

beaches and adjacent dunes along rivers (10 - 1000 m).

Behaviour: Unknown.

Seasonality: June to November.

Fish Hook Tiger Beetle, Cylindera ancistridia (Acciavatti and Pearson) (Plate 28) [Map 223] Identification: Length 11 mm; head and pronotum shiny copper; elytra copper green

with continuous whitish line along edge and

connected to three broad whitish lines into the interior of the elytra, the shoulder mark

has a distinctive hook at the end resembling a fish hook. Similar to the Dune Tiger Beetle (C. agnata), but the abbreviated "C" shape of its shoulder marking and smaller size distinguish it.

Subspecies and morphological variants: No subspecies have been described for this extremely localized species.

Distribution and habitats: Known only from wet sand beaches along the Ghaggar River in northern India (Punjab).

Behaviour: Unknown.

194 Seasonality: July.

Map 223 Fish Hook Tiger Beetle, Cylindera ancistridia.

Species Accounts

Wavy-lined Tiger Beetle, Cylindera venosa (Kollar)

(Plate 28) [Map 224]

Identification: Length 8-9 mm; head, pronotum and elytra shiny green with copper reflections; entire margin of elytra with a whitish-yellow, narrow stripe, shoulder

marking "C'shaped and connected to margin, middle band connected to the margin, long, sinuate, angled forward and then bending toward the rear extending almost to the end, rear marking connected to margin and extending forward as a short wavy line.

Subspecies and morphological variants: Some individuals are almost blackish in

background colour.

Distribution and habitats: Northern and . Map 224 Wavy-lined Tiger Beetle, Cylindera eastern

India,

Nepal,

Bangladesh,

and

venosa.

Myanmar. Margins of rivers, lakes and ponds with sandy beaches and little or no vegetation (35 - 3105 m).

Behaviour: Often abundant on the banks of rivers in the early monsoon, where it is extremely swift both running and flying. Attracted to lights at night. Seasonality: March to December.

Mudflat Tiger Beetle, Cylindera grammophora (Chaudoir) (Plate 28) [Map 225] Identification: Length 8 - 8.5 mm; head and pronotum green with copper reflections; elytra dull dark green with blue punctures and three unconnected whitish markings

on each elytron - shoulder marking in the

shape of a "C", middle marking right-angled and elbów-shaped, and rear marking a stalked spot connected to a short line along rear end of the elytra margin.

Subspecies and morphological variants: No subspecies have been described.

Distribution

and

habitats:

Northern and northeastern India, Pakistan, Nepal, and Bangladesh. Open sandy and muddy beaches along rivers (100-1340 m).

Map 225 Mudflat Tiger Beetle, Cylindera grammophora.

Behaviour: Attracted to lights at night. Seasonality: May to September. 195

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Bar-shouldered Tiger Beetle, Cylindera singalensis (Horn) (Plate 28) [Map 226] Identification: Length 8 - 8.5 mm; head, pronotum and elytra shiny bronze-green

with varied markings with irregular margins; no markings at the base of the shoulder, but thick white bar to the rear of the shoulder extending straight across the elyton from

the margin halfway to the midline, whitish middle marking curved and separated into blotches, rear marking complete and on some individuals connected by a wide border to the other markings.

Subspecies and morphological variants: Considerable individual variation in the extent of the marginal band width and

connection of markings on elytra.

Map 226 Bar-shouldered Tiger Beetle, Cylindera singalensis.

Distribution and habitats: Southern India and Sri Lanka. Dry, open sandy habitats along the sea coast and major rivers farther inland (0 - 50 m).

Behaviour: Unknown. Seasonality: July to October.

Species Accounts

Hook-legged Tiger Beetles, genus Myriochila The characteristics of this genus include peculiar coiled structures within the male genitalia; light-coloured markings fused together along the sides of the elytra, hooked hair-like setae on the longest upper leg segments (femora), and only four hairs arising near the front edge of a pale tan or white labrum (upper lip). The 47 species of this genus are divided into two subgenera, both of which are well represented throughout Africa, across southern Eurasia to Australia. Seven species are found in India, but none of them is endemic.

Key XXI: Hook-legged Tiger Beetles, genus Myriochila Identification Key to Myriochila Species la.

Legs at least partly pale and tawny; elytra

2

markings expanded 1b.

Legs completely dark metallic; elytra markings

3

divided, shortened or missing 2a. (la.)

Elytra bands along lateral margin all separate

Yellow-legged Tiger Beetle,

and distinct, divided into spots, two longest

Myriochila fastidiosa

segments of legs (femur and tibia) on most specimens partly pale with some metallic

reflections, on other specimens majority of these segments translucent tan

2b.

Elytra bands along lateral margin touching

Strife Tiger Beetle,

or joined into a continuous band at least

Myriochila litigiosa

in front half, two longest segments of legs

(femur and tibia) entirely pale tan 3a. (lb.)

Elytra shoulder band with basal portion

4

reduced to a small dot or entirely missing

3b.

Elytra shoulder band with basal portion

5

elongate although it may be separated from or joined to apical end

4a. (3a.)

Labrum one toothed in male, three toothed

Rippled Tiger Beetle,

in female; female elytra with a slightly shiny

Myriochila undulata

area at middle in centre on basal third 4b.

Labrum on three toothed in both sexes;

Imperfect Tiger Beetle,

female elytra without a slightly shiny area on

Myriochila atelesta

basal third 5a. (3b.)

Band at shoulder of elytra not divided, short and not reaching middle of elytra

Wavering Tiger Beetle,

Myriochila dubia

197

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

5b.

Band at shoulder of elytra divided in two,

6

apical spot located at middle of elytra on

some specimens, humeral band complete

and broadly joined to lateral expansion of middle band on other specimens

6a. (5b.)

6b.

First short segment (trochanter) of rear legs

Melancholic Tiger Beetle,

translucent, pale tan

Myriochila melancholica

First short segment (trochanter) of rear legs

Distinctive Tiger Beetle,

opaque, metallic violet to green-black or reddish

Myriochila distinguendo

Yellow-legged Tiger Beetle, Myriochila fastidiosa (Dejean) (Plate 29) [Map 227] Identification: Length 8-10 mm; head and pronotum shiny blue-green or copper; elytra

dull greenish bronze to copper with yellowish

markings - shoulder marking in a "C" shape, middle marking a short and broad disconnected band along outer edge with a small tail curving

into the interior of the elytron, and the rear

marking a broad band along the rear end of the elytron with a small tail extending forward, and an isolated spot on the rear of the elytron and near its inner margin. Legs

vary from light coloured to greenish.

Subspecies and morphological variants: Considerable individual variation in the degree of connections between markings on the elytra and leg colour. In some areas

Map 227 Yellow-legged Tiger Beetle, Myriochila fastidiosa-, A, M. f fastidiosa, B, M. f leucoloma.

brownish, greenish and bluish individuals occur together, perhaps a result of aging and exposure to the elements. Presently two subspecies are recognized from India,

but not all experts accept this division: M. f fastidiosa - elytral markings separate and distinct, many of which are divided into spots; upper legs with metallic reflections. M. f. leucoloma - all elytral markings usually joined by 3 continuous band; upper legs

light coloured with no metallic reflections.

Distribution and habitats: M. f. fastidosa - occurs on old fields, grasslands, forest paths, and floor of scrub forest (50 - 1340 m) in central, southern, northeastern India, Pakistan, Bangladesh, Myanmar, and Sri Lanka. M. f. leucoloma - found along edge of muddy ponds, stony rivers, and lakes in

northern India.

Behaviour: Attracted to lights at night. 198 Seasonality: July to November.

Species Accounts

Strife Tiger Beetle, Myriochila litigiosa (Dejean) (Plate 29) [Map 228] Identification: Length 8-10 mm; head and pronotum shiny blue-green or copper; elytra dull greenish bronze to copper with yellowish markings - shoulder marking re­

shaped, middle marking often connected by a broad band along outer edge to the middle and rear markings. Legs generally all light coloured.

Subspecies and morphological variants: Some individual variation in elytra markings

and leg colour.

Distribution and habitats: Northern India, Pakistan, and Nepal. Edges of muddy ponds

and lakes (5 - 1000 m).

. Map 228 Strife Tiger Beetle, Myriochila litigiosa.

Behaviour: Unknown. Seasonality: May to July.

Rippled Tiger Beetle, Myriochila ondulata (Dejean) (Plate 29) [Map 229] Identification: Length 10-12 mm; head and pronotum shiny green; elytra dull green, but female often with shiny area at front, thin whitish markings - shoulder patch reduced to a small spot or absent, middle marking with short wide bar along outer edge and a short curved line extending into the interior, and rear marking complete.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Widespread India except desert northwest and northeastern states, Nepal, Pakistan, Sri in

Lanka, and Bangladesh. Occupies edges of

small, temporary muddy ponds, lakes, rivers,

Map 229 Rippled Tiger Beetle, Myriochila undulata.

old fields, scrub forest and grassy meadows (50 - 3000 m).

Behaviour: Attracted to lights at night. Seasonality: May to December.

199

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Wavering Tiger Beetle, Myriochila dubia (Horn) (Plate 29) [Map 230] Identification:

Length

10

mm;

head,

pronotum and elytra shiny greenish bronze with

purple

punctures;

elytra

markings

include a complete "C" -shaped shoulder marking, middle marking a right-angled elbow, sometimes disrupted into separate spots and lines on some individuals, and the

rear marking complete. Similar to the Rippled

Tiger Beetle (M. undulata) but distinguished by the complete shoulder marking.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northeastern India,

Pakistan,

Nepal,

Myanmar,

and

Map 230 Wavering Tiger Beetle, Myriochila dubia.

Thailand. Sandy banks of rivers and occasionally muddy areas (600 - 1550 m).

Behaviour: Sometimes occurs together with the Rippled Tiger Beetle (M. undulata) in muddy habitats. Attracted to lights at night.

Seasonality: April to June.

Imperfect Tiger Beetle, Myriochila atelesta (Chaudoir) (Plate 29) [Map 231] Identification: Length 10-11 mm; head, pronotum and elytra dark brown with copper

reflections; white elytra markings include a white spot above the shoulder, middle marking reduced to a short band along the

outer edge and a spot in the interior, and the rear marking complete, females lack any shiny area at the front of the elytra.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northern and northwestern India, Nepal, Bangladesh, and China. Edges of muddy ponds and water tanks (250-1400 m).

Behaviour: Often occurs together with the Melancholic Tiger Beetle (M. melancholica).

200 Seasonality: July to August.

Map 231 Imperfect Tiger Beetle, Myriochila atelesta.

Melancholic Tiger Beetle, Myriochila melancholica (Fabricius) (Plate 29) [Map 232] Identification: Length 9.5-10 mm; head, pronotum and elytra dull greenish with copper reflections; elytral markings pale and narrow - shoulder "C" shaped often with a disrupted dot at the lower end, middle marking a disrupted right angle with a

broad base along the middle margin, and

rear marking complete; female with shiny spot on front third.

Subspecies and morphological variants: Several subspecies have been described for this cosmopolitan species, but only M. m. melancholica has been found in India.

Distribution and habitats: Throughout Europe, Africa, and Asia and enters south

Map 232 Melancholic Myriochila melancholica.

Tiger

Beetle,

Asia in northern India, Pakistan, Afghanistan, Nepal, and China. Occurs along bare edges of muddy ponds and lakes (150-1000 m).

Behaviour: Attracted to lights at night.

Seasonality: March to September.

Distinctive Tiger Beetle, Myriochila distinguendo (Dejean) (Plate 29) [Map 233] Identification: Length 10 - 12.5 mm; head and pronotum copper-green to green; elytra dull brown to black with thin whitish markings variable in size and shape but often consist of six irregular spots on each elytron, female with large an distinctive

shiny area on front quarter of elytra.

Subspecies and morphological variants: Some individuals with dark green head and

pronotum.

Distribution

and

habitats:

Southern

India and Sri Lanka. River edges inland and brackish mud puddles near the ocean (5 -

300 m).

Map 233 Distinctive Myriochila distinguenda.

Tiger

Beetle,

Behaviour: Attracted to lights at night.

Seasonality: January to October. 201

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Hairy-sided Tiger Beetles, genus Salpingophora This genus is characterized by unique male genitalia, eyes bulging out sideways, and extremely dense hair-like white setae on the underside. The five species in this genus are found on tidal flats, muddy estuaries and river deltas along the coast from the Red Sea to the Arabian Sea. One species is found in India.

Hairy-legged Tiger Beetle, Salpingophora maindroni (Horn) (Plate 29) [Map 234] Identification: Length 14-16 mm; Head and pronotum shiny dark bronze to bronze

green; elytra bronze green with broad and irregular tan band along the outer edge; dense white hair-like setae on the underside of the body and extending onto the upper

parts of the front and middle legs.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: Northwestern India in Gujarat and Pakistan. Clay and compact soils near water.

Behaviour: Unknown. Seasonality: July to October.

234 Hairy-legged Salpingophora maindroni.

Map

Tiger

Beetle,

Species Accounts

Flat-eyed Tiger Beetles, genus Hypaetha The fourteen species of this genus inhabit sandy ocean beaches from South Africa and Arabia to Indonesia and Australia. Distinguishing characteristics include a broadly concave area on the head between the extremely flat outward bulging eyes, a distinctive fold evident on the pronotum within its side margins, and white hair-like setae so thick on the underside that they hide the body segments. Three of these species occur along the coastal beaches of India.

Key XXII: Flat-eyed Tiger Beetles, genus Hypaetha Identification Key to Hypaetha Species 1a.

Two longitudinal bands along each margin of each wing cover (elytron)

2

1b.

One longitudinal band only along lateral margin of each wing cover (elytron) ’

Asian Coastal Tiger Beetle, Hypaetha biramosa

2a. (la.)

Both longitudinal bands on each wing cover about equal in length, separate or narrowly joined near middle; body large (14-19 mm)

Four-lined Tiger Beetle, Hypaetha quadrilineata

2b.

Inner longitudinal band on each wing cover (elytron) shorter than outer one -broadly joined to outer band at middle by a short transverse band; body small (8 mm)

Sandspit Tiger Beetle, Hypaetha copulata

Four-lined Tiger Beetle, Hypaetha quadrilineata (Fabricius) (Plate 30) [Map 235] Identification: Length 14-19 mm; head and pronotum copper with some green reflections; elytra dark greenish bronze with two wide

yellow-white bands running the length of each

elytron, the one along the outer edge usually narrower; underside of body densely covered with white hair-like setae that extend onto the upper parts of the legs.

Subspecies and morphological variants:The two longitudinal bands on the elytra merge together at various points in some individuals.

Distribution and habitats: East and west coasts of India, Iran, Pakistan, Sri Lanka, Myanmar, Thailand, and Malaysia. Sandy beaches along the coast and inland on large river deltas.

Map 235 Four-lined Tiger Beetle, Hypaetha quadrilineata.

Behaviour: Individuals disperse away from the ocean beaches along lowland river beaches at some times of the year. They are strong fliers.

Seasonality: January to October.

203

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindeiidae

Asian Coastal Tiger Beetle, Hypaetha biramosa (Fabrictus) (Plate 30) [Map 236] Identification: Length 10-13 mm; Head, pronotum and elytra dark bronze or bronze­

green with copper reflections; markings on elytra ivory white border along outer edges

bulging inward at the middle and at the rear.

Subspecies and morphological variants: Some individuals are dark blue-green almost

black and show considerable individual variation in the width of the ivory-white markings.

Distribution and habitats: East and west coasts of India, Andaman and Nicobar Islands, Bangladesh, Sri Lanka, Myanmar, Thailand, Malaysia, and Indonesia. Undisturbed outer sandy ocean beaches.

Map 236 Asian Coastal Tiger Beetle, Hypaetha biramosa.

Behaviour: Often forage in intertidal areas running in shallow water. Seasonality: February to December.

Sandspit Tiger Beetle, Hypaetha copulate (Schmidt-Goebel) (Plate 30) [Map 237] Identification:

Length 8 mm; head, pronotum and elytra shiny bronze with green reflections toward the sides; broad whitish band along the entire outer margin of the elytra continuing forward around the front and then back along the inner edge one fourth the length, middle marking extending at right angles from the outer edge toward the interior and then turning at sharp right angle toward the rear; underside of body without dense white hair-like setae.

Subspecies and morphological variants: One

additional

subspecies

has

described from the Arabian Peninsula.

Distribution

and

habitats:

been Map 237 Sandspit Tiger Beetle, Hypaetha copulata.

Extreme northwestern India in Gujarat, Iran, Pakistan, Saudi Arabia, United Arab Emirates, and Oman. Sandy seashore beaches and stream edges leading into the ocean.

Behaviour: Runs quickly to avoid disturbances and flies short distances only occasionally. 204 Seasonality: April to October.

Species Accounts

Thick-thighed Tiger Beeties, genus Callytron The eleven species of tiger beetles in this genus occur from Arabia to Japan and Indonesia. They share long, slender legs that have the bases characteristically swollen. There are only a few hair-like setae on the underside of the body. The markings on the elytra are reduced to a narrow continuous band along die edges or completely absent They occupy muddy substrates at river mouths and brackish areas along the coast Three species are found around the southern margins of India and one reaches its Indian Ocean archipelagoes.

Key XXIII: Thick-thighed Tiger Beetles, genus Callytron Identification Key to Callytron Species la.

Elytra marked with a narrow, continuous

2

lateral band from shoulder to apical suture

1b.

Spotless Tiger Beetle,

Elytra without markings

Callytron malabaricum

2a. (la.)

Pronotum underside bare anterior to front

Side-banded Tiger Beetle,

legs, four setae near from edge of labrum

2b.

Callytron gyllenhalii

Pronotum underside with setae anterior to

Dull Tiger Beetle,

front legs, eight setae near middle of labrum

Callytron limosum

Side-banded Tiger Beetle, Callytron gyllenhalii (Dejean)

(Plate 30) [Map 238]

Identification: Length 9 - 9.5 mm; head, pronotum, and elytra shiny greenishbronze; elytra with a thin white border

along the entire outer edge; female elytra oval-shaped. Similar to the Dull Tiger beetle (C. limosum), which has a dull elytral surface.

Subspecies and morphological variants: Two subspecies have been described based on the presence or absence of the white outer border on the elytra. Only the form (C. g. gyllenhalii) with the white border is

known to definitely, albeit rarely, occur in India. However, an unlabeled specimen with no white border on the elytra may be from India. This form is considered a subspecies,

C. g. immarginatum, by some experts.

238 Side-banded Tiger Beetle, Callytron gyllenhalii; A, C. g. gyllenhalii, B, C. g. immarginatum.

Map

Distribution and habitats: Western India in Maharashtra and Pakistan. Coastal muddy areas and clay patches along streams running into the ocean. 205

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Behaviour: Occasionally found together with the rare Sandspit Tiger Beetle (H. copulata) and the more common Four-lined Tiger Beetle (H. quadrilineata).

Seasonality: July to August.

Dull Tiger Beetle, Callytron limosum (Saunders) (Plate 30) [Map 239] Identification: Length 9 mm; head and pronotum dull green with copper reflections; elytra dull green with narrow white band along entire outer edge. Similar to the Side-

banded Tiger beetle (C. gyllenhalii), which

has a shiny elytra surface.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: East coast of India and occasionally more inland and Andaman and Nicobar Islands, Sri Lanka, Myanmar, Thailand, and China. Brackish

muddy areas along coastal deltas and inland on coastal plain rivers and salt lakes.

Map 239 Dull Tiger Beetle, Callytron limosum.

Behaviour: Quick to fly from disturbances and difficult to see against the dark substrates. Often hides in the burrows of ghost crabs when disturbed.

Seasonality: March to September.

Spotless Tiger Beetle, Callytron malabaricum (Fleutiaux and Maindron) (Plate 30) [Map 240] Identification: Length 7.5 - 9.5 mm; Head and pronotum bronze above with blue­

green sides; elytra dull bronze almost black with no light markings.

Subspecies and morphological variants: No subspecies have been described.

Distribution and habitats: West coast of India and Pakistan. Muddy areas at or near the sea shore and along coastal rivers and deltas.

Behaviour: Unknown. Seasonality: June to July. Map 240 Spotless Tiger Beetle, Callytron 206

malabaricum.

Species Accounts

Enigmatic Tiger Beetles, genus Apteroessa This is the most mysterious tiger beetle in India as it has not been encountered in South India since its original discovery there over 200 hundred years ago in the Eighteenth Century. Its stout legs, lack of flight wings, large body, unusually large head, and its elytral surfaces covered with hair-like setae provide few clues as to its phylogenetic position within the family. Numerous researchers have tried to find additional specimens near its purported type locality, but without success. There is only one species in the genus.

Enigmatic Tiger Beetle, Apteroessa grossa (Fabricius) (Plate 30) [Map 241] Identification: Length 21-22 mm; black above on all surfaces; head and pronotum are large and about the same width; eyes small; elytra oval, three large yellow spots on each elytron with white hair-like setae along their rear margins and scattered over much

of the upper surface; head and pronotum honey-combed with pits that are also deep

in front half of elytra; lacks wings.

Subspecies and morphological variants: No subspecies of this highly localized

species have been described.

Distribution

and

habitats:

Since

its

discovery in South India, apparently at Ammayanayakanur (Tamil Nadu), in the

Map 241 Enigmatic Tiger Beetle, Apteroessa grossa.

late 1700's no specimens have been collected nor individuals observed despite the efforts of numerous tiger beetle enthusiasts to discover its whereabouts.

Behaviour: Unknown.

Seasonality: Unknown.

207

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Plate 1

Metallic Tiger Beetles and Arboreal Tiger Beetles Metallic Tiger Beetles, Grammognatha 1 Eurasian Metallic Tiger Beetle, Grammognatha euphratica Widespread in southern Europe and southwestern Asia, rare in northwest desert of India. Large, robust, metallic green with maroon reflections. Large yellow spot on back half of each elytron, legs yellow. (p. 30)

Arboreal Tiger Beetles, Tricondyla 2 Pleated Arboreal Tiger Beetle, Tricondyla femorata Southern India and Sri Lanka. Black and shiny. Long cylinder-like thorax with slight constriction at neck. Elytra with fine wrinkles running across width. (p. 32) 3 Satin Arboreal Tiger Beetle, Tricondyla gounellei Endemic to southern India. Black and velvet-like. Long cylinder-like thorax with slight constriction at neck. Elytra with shallow, uniform pits. (p. 32) 4 Thick-necked Arboreal Tiger Beetle, Tricondyla macrodera Northeastern India to Vietnam. Brown and shiny. Relatively short and thick thorax with deep constriction at neck. Elytra wrinkled on front half and smooth towards rear. (p. 33) 5 Lumpy Arboreal Tiger Beetle, Tricondyla tuberculata Northeastern India to Myanmar. Dull black to dark brown. Relatively short thorax with moderate constriction at neck. Elytra deeply wrinkled in front and distinctly pitted towards rear. (p. 33)

208

Plate 1

209

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Plate 2

Bark Tiger Beetles and Brush Tiger Beetles I Bark Tiger Beetles, Derocrania 1 Camel-backed Bark Tiger Beetle, Derocrania longesulcata Endemic to Western Ghats of southern India. Shiny black. Thorax distinctly constricted at front third. Many small but well-defined grooves running length of elytra. Rear half of elytra expanded abruptly into a bulbous shape (p. 35) 2 Flat-headed Bark Tiger Beetle, Derocrania honorei Endemic to Western Ghats of southern India. Shiny black. Thorax distinctly constricted at front third, irregular grooves running length of elytra. Rear two-thirds of elytra expanded gradually. (p. 35) 3 Long-necked Bark Tiger Beetle, Derocrania dembickyi Endemic to Western Ghats of southern India. Shiny black. Thorax gradually constricted in front fourth. Shallow but distinct grooves running length of elytra. Rear two-thirds of elytra expanded gradually. (p. 36) 4 Short-necked Bark Tiger Beetle, Derocrania brevicollis Endemic to Western Ghats of southern India. Shiny black. Thorax relatively short and gradually constricted at front fourth. Irregular, shallow grooves running length of elytra. Rear two-thirds of elytra expanded gradually. (p. 36)

Primitive Brush Tiger Beetles, Protocollyris 5 Nilgiri Brush Tiger Beetle, Protocollyris nilgiriensis Endemic to Nilgiri Hills of southern India. Dark blue. Thorax long and narrow. Elytra slightly expanded towards rear and with distinctive pitting spread evenly over entire surface. (p. 38) 6 Stout Brush Tiger Beetle, Protocollyris pacholatkoi Endemic to Nilgiri Hills of southern India. Dark blue. Thorax relatively short and thick. Elytra nearly parallel-sided and with shallow pitting over their entire surface. (p. 38) 7 Short-lipped Brush Tiger Beetle, Protocollyris brevilabris Eastern India to Malaysia. Dark blue to green. Thorax short and thick. Elytra nearly parallel-sided with shallow, irregular pitting. (p. 39) 8 Brittle Brush Tiger Beetle, Protocollyris fragilis Endemic to Kerala in southern India. Blue to greenish. Extremely slender head and thorax contrast with relatively broad elytra. Tiny labrum. (p. 39)

Brush Tiger Beetles, Neocollyris 9 Maroon-spotted Brush Tiger Beetle, Neocollyris purpureomaculata Northeastern India to China. Small. Head and thorax black with green reflections. Elytra bright green often with two large maroon to purple bands across each elytron. (p. 49)

210

Plate 2

211

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Plate 3

Brush Tiger Beetles II Brush Tiger Beetles, Neocollyris 1 Ingrid’s Brush Tiger Beetle, Neocollyris ingridae Endemic to northeastern India. Small. Head shiny dark purple-green to black. Long and slender thorax shiny dark green. Elytra green with small oblong reddish patch on side of each elytron. (p. 49) 2 Thin Brush Tiger Beetle, Neocollyris macilenta Endemic to northeastern India. Small. Head and relatively short thorax shiny green. Elytra parallel-sided and green with small oblong reddish patch on margin of each elytron. (p. 50) 3 Ringed Brush Tiger Beetle, Neocollyris annulicornis Endemic to Northeastern India. Small. Head and relatively short neck shiny blue and green. Elytra green and conspicuously widened towards rear. (p. 50) 4 Yellow-legged Brush Tiger Beetle, Neocollyris roeschkei Endemic to northeastern India. Medium size. Head dark violet. Thorax very short and bright blue. Elytra blue-green with violet borders and triangular shaped reddish spot on mid-margin of each elytron. Bright yellow narrow band across front edge of elytra. Antennae and legs completely yellow. (p. 51) 5 Meghalaya Brush Tiger Beetle, Neocollyris pearsoni Endemic to northeastern India. Small. Blue. Thorax short and broad. Elytra with small yellowish oblong patches on mid-margin. (p. 51) 6 Blue-feeler Brush Tiger Beetle, Neocollyris cyaneipalpis Endemic to Western Ghats of southern India. Small. Head and thorax shiny blue. Labrum short and completely black with blue reflections. Elytra blue-green with small light areas along mid-margin. Long antennae. (p. 52) 7 Four-lined Brush Tiger Beetle, Neocollyris quadrisulcata Endemic to southern India. Small. Shiny blue. Relatively short antennae. Labrum black with yellow centre. (p. 52) 8 Graceful Brush Tiger Beetle, Neocollyris redtenbacheri Northeastern India to Myanmar. Medium size. Bright green to blue green. Labrum metallic blue. Small yellow spots on mid-margin of elytra. Last segment of abdomen yellow. (p. 53) 9 Husky Brush Tiger Beetle, Neocollyris schaumi Endemic to Andaman Islands. Medium size. Blue head. Dark green thorax and elytra. Labrum metallic green. (p. 53)

212

Plate 3

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Plate 4

Brush Tiger Beetles III 1 Red-legged Brush Tiger Beetle, Neocollyris bonellii Widespread throughout India except desert northwest and extend east to Indonesia. Medium size. Head and thorax dark blue to black. Elytra bright blue. Upper parts of legs reddish-orange. (p. 54) 2 Long-lipped Brush Tiger Beetle, Neocollyris distincta Endemic to peninsular India. Medium size. Bright blue to green or violaceous. Legs red. Long labrum black. (p. 54) 3 Nepal Brush Tiger Beetle, Neocollyris nepalensis Northeastern India to Myanmar. Medium size. Head black with violet reflections. Body long, robust and blue. (p. 55) 4 . Orange-spotted Brush Tiger Beetle, Neocollyris hiekei Endemic to northeastern India. Small. Head black with violet reflections. Thorax bright violet. Elytra bright blue-violet with large bright orange spots on mid-margin and at shoulder. (p. 55) 5 Blood-spotted Brush Tiger Beetle, Neocollyris cruentata Northeastern India to Thailand. Medium size. Large head. Short, thick thorax. Body blue, green violaceous or brownish. Elytra often with dark red stripe on shoulders and irregular reddish patch at mid-margin. Outermost segments of hind legs light-coloured. (p. 56) 6 Round-headed Brush Tiger Beetle, Neocollyris batesi Northeastern India to Laos. Medium size. Head black, round and large with relatively small eyes. Thorax blue-violet. Elytra roughly pitted, blue with no reddish marks. (p. 56) 7 Golden Brush Tiger Beetle, Neocollyris orichalcina Southern and northeastern India to Laos. Medium size. Large head and short, black thorax with blue and violet reflections. Elytra shiny violet or blue with large and irregular pitting. (p. 57) 8 Dark-legged Brush Tiger Beetle, Neocollyris fuscitarsis Northeastern India to Vietnam. Medium to large. Brilliant blue, rarely violet or green. Thorax short. Elytra with uniform shallow rounded pits and indistinct red, oblong spot mid-margin. (p. 57) 9 Broad-headed Brush Tiger Beetle, Neocollyris insignis Northeastern India to Nepal. Large. Head large and black with blue reflections. Thorax and elytra blue rarely brown with violaceus reflections. Elytra long and broaden gradually towards rear; their central surfaces covered with deep labyrinth of connected depressions. (p. 58)

214

Plate 4

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Plate 5

Brush Tiger Beetles IV Brush Tiger Beetles, Neocollyris 1 Blue Brush Tiger Beetle, Neocollyris smaragdina Northeastern India to Myanmar. Large. Head and thorax dark blue. Elytra bright blue and parallel-sided. (p. 58) 2 Sapphire Brush Tiger Beetle, Neocollyris saphyrina Northeastern India to Laos. Medium to large. Head large and thorax short, black with blue reflections. Elytra bright blue. (p. 59) 3 Eminent Brush Tiger Beetle, Neocollyris egregia Endemic to northeastern India. Large. Entirely violet. Head rounded and wide. Elytra with short, thin light line at mid-margin. Deep and convoluted pits in surface at middle of elytra. (p. 59) 4 Blue-lipped Brush Tiger Beetle, Neocollyris similis Rare species in India with record from Tamil Nadu and uncertain records from northeastern India to Laos. Medium size. Head and thorax blue-black. Labrum long and dark blue. Elytra blue with dense pits forming a confluence of polygons towards centre. (p. 60) 5 Red-tipped Brush liger Beetle, Neocollyris rufipalpis Northeastern India to Vietnam. Medium size. Head and thorax blue, short, and wide. Elytra blue-green, short and expand towards rear. Labrum long and metallic blue. (p. 60) 6 Large-necked Brush liger Beetle, Neocollyris crassicornis India to Taiwan. Medium size. Large head black with blue reflections. Short, thick, blue thorax. Parallel-sided elytra blue to violaceous with their surfaces covered by a network of interlocking punctures. Short, thick antennae. (p. 61) 6a N. c. crassicornis -- mainland form smaller with smoother elytra surface. 6b N. c. andamana -- Andaman Islands form larger, more elongate, and with coarser elytra sculpturing. 7 Greyish Brush Liger Beetle, Neocollyris subclavata Southern India to Malaysia. Medium to large. Blue to grey. Head large and round. Thorax long with distinct lines running from side to side on its upper surface. Indistinct reddish patch at mid margin of elytra. (p. 61) 7a N. s. subclavata -- blue and finer elytra microsculpture, northern part of Western Ghats. 7b N. s. continentalis -- steel grey with blue reflections and coarser elytra microsculpture, southern part of Western Ghats and Andaman islands.

216

Plate 5

217

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Plate 6

Brush Tiger Beetles V Brush Tiger Beetles, Neocollyris 1 Brown Brush Tiger Beetle, Neocollyris acuteapicalis Endemic to northeastern India. Medium size. Dark brown with blue reflections along sides. Thorax long. Antennae short. Rufous oval patch on mid margin of elytra (p. 62) 2 Hairy Brush Tiger beetle, Neocollyris multipilosa Rare species endemic to southern India. Medium size. Dull black. Long, thin thorax covered with white hairs. Elytra with uniform pits. (p. 62) 3 Tapered Brush Tiger Beetle, Neocollyris attenuata Northern and northeastern India west to Pakistan and east to China. Medium size. Long and oval head and thorax black with blue or violet reflections. Elytra bright green and expanded in width towards rear. (p. 63) 4 Slim Brush Tiger Beetle, Neocollyris brancuccii Endemic to Northeastern India. Medium size. Head and thorax dark green. Elytra bright green with copper reflections. Body extremely elongated and thin. Labrum rounded and black with pale yellow centre. (p. 63) 5 Long-necked Brush Tiger Beetle, Neocollyris brendelli Endemic to Northeastern India. Medium size. Head black with violet reflections and elongated with relatively small eyes. Thorax black with violet reflections and extremely narrow column at front and bulbous at rear. Elytra narrow and long. Labrum rectangular and brown. (p. 64) 6 Lesser Brush Tiger Beetle, Neocollyris parvula Endemic to southern India. Small. Bright blueviolet. Head narrow. Thorax round and narrow in front and expanded towards rear. Elytra widened distinctly towards rear. (p. 64) 7 Long-legged Brush Tiger Beetle, Neocollyris maindroni Endemic to southern mdia. Small. Dark green to blackish with green to copper reflections. Head narrow and elongated. Thorax extremely long and thin with distinct cross lines. Elytra short. (p. 65) 8 Short-necked Brush Tiger Beetle, Neocollyris kollari Endemic to India. Medium size. Bright blue. Head rounded. Thorax short. Elytra surface with uniform small pits. (p. 65) 9 Diamond-headed Brush Tiger Beetle, Neocollyris variicornis Northeastern India and Andaman Islands to Vietnam. Medium size. Triangular-shaped head black with violet reflections. Long vase-shaped thorax black with green or blue reflections. Elytra bright green, dark blue, or purple often with a reddish mark at mid-margin and small one at shoulder. (p. 66) 218

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Brush Tiger Beetles VI Brush Tiger Beetles, Neocollyris 1 Pacholatko’s Brush Tiger Beetle, Neocollyris pacholatkoi Endemic to northeastern India. Medium size. Head bluish-black with short yellow-brown antennae, Thorax bluish-black and shaped like a flask, covered with distinct white hairs. Elytra greenish black with small yellow band across middle, punctures form rows across centre. (p. 66) 2 Spindle-necked Brush Tiger Beetle, Neocollyris variitarsis Northeastern India to Vietnam. Medium size. Elongate head black with violet or blue reflections. Thorax black with violet or blue reflections and expanding from narrow front to distinctly widened rear. Elytra blue or rarely green, long, and parallel-sided with narrow reddish line across middle. (p. 67) 3 Bottle-necked Brush Tiger Beetle, Neocollyris ampullicollis Endemic to southern india. Medium to large. Black with violet reflections. Head round. Thorax long with distinct constriction in front and extremely expanded rear. Elytra long. (p. 67) 4 Furrow-necked Brush Tiger Beetle, Neocollyris plicicollis Endemic to southern India. Medium size. Head and thorax bright dark blue. Elytra bronze to olive with large yellow patch on mid-margin. (p. 68) 5 Metallic Brush Tiger Beetle, Neocollyris metallica Endemic to southern India. Small to medium. Shiny and metallic dark green or blue with violaceous patches on front of elytra. Elytra parallel-sided with small pale spot at mid-margin. ( p. 68) 6 Smooth-necked Brush Tiger Beetle, Neocollyris subtileflavescens Endemic to southern India. Medium size. Bright dark brown-violet. Thorax short and stout. Elytra parallel-sided with small light yellow spot at mid-margin. (p. 69) 7 Wrinkle-necked Brush Tiger Beetle, Neocollyris rugata Endemic to southern India. Medium size. Head black with violet reflections. Thorax and elytra dark green. Thorax long and slender. Elytra expanding slightly to rear and yellowish spot at mid-margin. (p. 69) 8 Yellow-spotted Brush Tiger Beetle, Neocollyris conspicua Endemic to southern India. Small. Head and thorax black with blue reflections. Elytra dark brown with large yellow patch mid-margin and small reddish spot at shoulder. (p. 70) 9 Greenish Brush Tiger Beetle, Neocollyris vannideki Endemic to southern India. Small. Head and thorax coppery green. Extremely long and slender thorax with constriction. Elytra parallel-sided and bronze with distinctive patch of yellow-red on mid-margin. (p. 70) 220

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Brush Tiger Beetles VII Brush Tiger Beetles, Neocollyris 1 Least Brush Tiger Beetle, Neocollyris flava Endemic to southern India. Small. Head and short thorax bright green. Elytra brown with copper reflections and large yellow patch mid-margin, sides expanding towards rear. (p. 71) 2 Blackish Brush Tiger Beetle, Neocollyris juengeri Endemic to southern India. Large. Bright blackish grey. Thorax long and elytra long expanding slightly towards rear. (p. 71) 3 Charcoal Brush Tiger Beetle, Neocollyris anthracina Endemic to southern India. Large. Head and thorax shiny black with green reflections. Thorax extremely long and constricted. Elytra long, thin, parallel-sided and matte-black with shades of dark green. (p. 72) 4 Brown-kneed Brush Tiger Beetle, Neocollyris nilgirica Endemic to southern India. Medium size. Greenish-bronze. Thorax long and slender with strong cross striations. Elytra long and narrow with yellow mark at mid-margin. (p. 72) 5 Small-eyed Brush Tiger Beetle, Neocollyris andrewesi Endemic to southern India. Large. Black with blue reflections. Thorax long and stout with distinct constriction. Elytra expanded slightly towards rear. (p. 73) 5a N. a. andrewesi -- somewhat smaller. Western Ghats of Karnataka. 5b N. a. regia -- somewhat larger with minor differences in anatomy. Western Ghats of Kerala and Tamil Nadu. 6 Chestnut Brush Tiger Beetle, Neocollyris rubens Northeastern India to Vietnam. Large. Chestnut brown with darker head. Thorax strongly constricted forming two humps. Elytra surfaces smooth except for 5 or 6 deep and sharp wrinkles in their middle. Elytra edges slightly constricted in middle. (p. 73) 7 Narrow-necked Brush Tiger Beetle, Neocollyris compressicollis Northeastern India to Vietnam. Large. Chestnut brown with darker head. Thorax strongly constricted forming two humps. Elytra surface with polygons and wrinkles almost entire length. Elytra edges slightly constricted in middle. (p. 74) 8 Long-backed Brush Tiger Beetle, Neocollyris apteroides Endemic to Northeastern India. Medium to large. Shiny dark brown. Thorax short with distinct constriction towards front. Elytra long and only slightly expanded towards rear. Elytra surfaces with deep polygons in middle and faint reddish spots at mid-margin and at shoulder. (p. 74) 222

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Brush Tiger Beetles VIII Brush Tiger Beetles, Neocollyris 1 Pitted Brush Tiger Beetle, Neocollyris foveifrons Northeastern India to Vietnam. Large. Shiny red to matte brown or occasionally bluish. Thorax short and stout. Elytra long and expanded towards their rear, and surfaces with distinct folds and wrinkles in middle. (p. 75) 2 Assam Brush Tiger Beetle, Neocollyris assamensis Endemic to northeastern India. Medium to large. Dark reddish brown. Thorax long with distinct constriction near front. Elytra exceptionally long with moderate widening at their rear, and surface with deep folds restricted to middle. (p. 76) 3 Rufous Brush Tiger Beetle, Neocollyris coapteroides Endemic to northeastern India. Large. Head and thorax brownish. Elytra bright reddish brown, long, slender, and slightly widened at rear. (p. 76) 4 Thin-spotted Brush Tiger beetle, Neocollyris smithii Northeastern India to Myanmar. Large size. Dark blue to black. Thorax long and stout. Elytra long with large light brown patches at shoulder and rear. (p. 76)

Large Brush Tiger Beetles, Collyris 5 Notch-lipped Brush Eiger Beetle, Collyris longicollis Northeastern India and Nepal. Large. Blue to dark blue or blackish. Thorax relatively short and stout with cross striations. Elytra parallel-sided with metallic blue and rounded pits in front and rear and polygons in middle. (p. 78) 6 Violet Brush Tiger Beetle, Collyris dohrnii Southern India and Sri Lanka. Large. Violaceus brown to purple. Short, thick thorax smooth on top. Elytra long and gradually widening toward rear. (p. 78) 7 Short-winged Brush Liger Beetle, Collyris brevipennis Endemic to southern ndia. Large. Bright blue. Thorax long with obvious cross striations. Elytra surface with deep polygons in middle but more rounded ones towards front and rear. (p. 79) 8 Smooth Brush Tiger Beetle, Collyris subtilesculpta Endemic to southern India. Large. Head black or violet. Thorax dark blue, long and larger at base. Elytra dark to bright blue, long, parallelsided with relatively small surface pits along their entire length. (p. 79) 9 Punctured Brush Tiger Beetle, Collyris dormeri Northeastern India to Laos. Large. Head and stout thorax black with violet reflections. Elytra violaceous brown with extensive punctures and folding on central third of elytra surface. (p. 80) 224

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Knobby Tiger Beetles I Knobby Tiger Beetles, Therates 1 Brown-legged Knobby Tiger Beetle, Therates annandalei Endemic to northeastern India. Head and thorax shining greenish-black. Elytra shining black, each with long brownish yellow front mark, small yellow central dot, and light ye low patch covering rear humps. (p. 82) 2 Vellow-spotted Knobby Tiger Beetle, Therates nepalensis Northeastern India and Nepal. Head and thorax shiny greenish-black. Elytra shiny greenish or violet black with markings orange in front and yellow in middle and rear. (p. 82) 3 Brown-spotted Knobby Tiger Beetle, Therates jendeki Endemic to northeastern India. Head and thorax shiny greenish-black. Elytra shiny black, each with broad brownish yellow front marking, right-angled brownish yellow central dot, and brown rear patch. (p. 83) 4 Dark-lipped Knobby Tiger Beetle, Therates arunachalcolus Endemic to Northeastern India. Head and thorax shiny greenish-black. Elytra shiny brownish-black, each with long yellow front mark, broad middle band, and yellow rear mark that reaches humps. (p. 83) 5 Ingrid’s Knobby Tiger Beetle, Therates ingridae Endemic to northeastern India. Head and thorax shiny black. Elytra with yellow, variable-shaped front mark, yellow central dot, and indistinct pale crescent at rear. (p. 84) 6 Brown-bellied Knobby Tiger Beetle, Therates sausai Endemic to northeastern India. Head shiny greenish-black with reddish forehead. Thorax greenish-black. Elytra shiny brownish-black, each with large “J”-shaped brownish-yellow front mark, yellow horizontal central dot, and pale yellow rear patch. (p. 84) 7 Brown-vented Knobby Tiger Beetle, Therates dohertyi Endemic to northeastern India. Head and thorax shiny greenish-black. Elytra shiny brownish-black, each with brownish-yellow “C”-shaped front mark, short yellow central band, and light yellow rear patch. Labrum yellowish. (p. 85) 8 Red-fronted Knobby Tiger Beetle, Therates waagenorum Endemic to northeastern India. Head shiny greenish-black with extensive reddish forehead. Thorax shiny greenish-black. Elytra shiny brownish-black, each with brownish-yellow front mark that may be separated into three large dots or coalesced to connect to each or may join with yellow horizontal central dot, and pale yellow rear patch. (p. 85) 9 Wrinkled Knobby Tiger Beetle, Therates westbengalensis Endemic to northeastern India. Head and thorax greenish-black. Elytra yellow-brown, each with brighter yellow patches at rear end and a long narrow stripe running along side from shoulder two thirds length of elytra. (p. 86) 226

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Knobby Tiger Beetles II, Wrinkled Tiger Beetles, Apothecary Tiger Beetles, Jungle Tiger Beetles, and Glossy Tiger Beetles I Knobby Tiger Beetles, Therates 1 Hennig’s Knobby Tiger Beetle, Therates hennigi Endemic to northeastern India. Head and thorax shiny black with blue-green or violet reflections. Front halves of elytra orange, rear halves with two large patches of shiny black in middle and rear surface separated by a broad bright yellow middle band extending entire width of elytra and a large yellow area at rear end of elytra. (p. 86) 1a T. h. hennigi - orange front half of elytra connected to yellow central band by a narrow bright strip. Meghalaya and Assam. 1b T. h. dormeri - orange front half of elytra separated from central band. Arunachal Pradesh to Manipur.

Wrinkled Tiger Beetles, Rhytidophaena 2 Ornamental Wrinkled Tiger Beetle, Rhytidophaena limbata Northern India to Pakistan and Bangladesh. Elytra shiny crimson occasionally with violet tinge on shoulders. Thorax green or bright blue on its sides. Central line between elytra and sides green with two distinct white spots along outer margins. (p. 87) 3 Plain Wrinkled Tiger Beetle, Rhytidophaena inornata Endemic to northern India. Head dark coppery-bronze with purple sides. Thorax and elytra bronze-green. Elytra with two white spots along outer margins in rear half. (p. 88)

Apothecary Tiger Beetles, Prothyma 4 Azure Tiger Beetle, Prothyma proxima India and Sri Lanka. Head and thorax dark green to black. Elytra dark blue with two small white spots along outer margin. (p. 89) 5 Assam Tiger Beetle, Prothyma assamensis Northeastern India and Nepal. Head, thorax and elytra shiny olive with blue-green reflections. Elytra with three pale whitish spots on outer edges. (p. 89)

Jungle Tiger Beetles, Heptodonta 6 Beautiful Jungle Tiger Beetle, Heptodonta pulchella Northern India to Vietnam. Dull olive green. Elytra surfaces unmarked with fine sculpturing on surface. (p. 90)

Glossy Tiger Beetles, Pronyssa 7 Hennig’s Glossy Tiger Beetle, Pronyssa hennigi Endemic to northeastern India. Head and thorax dark, coppery green. Elytra shiny coppery-green with three white spots along margin. (p. 91) 8 Green Glossy Tiger Beetle, Pronyssa nodicollis Northeastern India to Vietnam. Bright green with black-green reflections. Elytra with three white spots along margins. (p. 92) 228

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Glossy Tiger Beetles II and Beautiful Tiger Beetles I Glossy Tiger Beetles, Pronyssa 1 Kraatz’s Glossy Tiger Beetle, Pronyssa kraatzi Endemic to northeastern India. Bluish green. Elytra with blue sides and no light spots (p. 92) 2 Assam Glossy Tiger Beetle, Pronyssa assamensis Endemic to northeastern India. Copper brown with reflections. Elytra sides blue with two distinct white spots at middle and rear. (p. 93) 3 Montane Glossy Tiger Beetle, Pronyssa montanea Endemic to northeastern India. Head and thorax blue-green. Elytra green with blue-violet sides and with three white spots along margins. (p. 93) (Fig. 30.1)

Beautiful Tiger Beetles, Calochroa 4a Yellow-spotted Tiger Beetle, Calochroa flavomaculata India west to Pakistan and east to Philippines. Velvety dark black to greenish or bluish-black. Thorax with no hair-like setae along the side of the upper edge (4b). Elytra each with three pale yellow spots in a row down their centres. (p. 97) 5a Six-dotted Tiger Beetle, Calochroa sexpunctata East coast of India and Andaman Islands to Philippines. Head and thorax shiny bronze and metallic green. Thorax with hair-like setae on along the side of the upper edge (5b). Elytra dull black with green sides and three oval spots in a row down their centres. (p. 97) 6 Green Tiger Beetle, Calochroa whithillii Endemic to southern India. Head and thorax green, blue-green to blue. Elytra dull green, blue-green or blackish-green with shiny green sides and a single small white dot in middle near margin. (p. 98) 7 Greater Spotted Tiger Beetle, Calochroa octonotata Northern India to Myanmar. Very large. Head and thorax deep purple-blue and metallic green with bright copper-red reflections. Elytra blue and green with copper reflections and three large yellow markings on each elytron. (p. 98) 230

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Beautiful Tiger Beetles II Beautiful Tiger Beetles, Calochroa 1 Blackish Tiger Beetle, Calochroa tritoma Northeastern India to Thailand. Head and thorax black to blue-green. Elytra dull black with bold yellow-orange band running down each elytron from shoulder towards middle, aligned with two large yellow dots in middle and rear of each elytron. (p. 99) 2 Lesser Spotted Tiger Beetle, Calochroa octogramma Northern India to China. Head and thorax shiny black with bright copper and green reflections. Elytra dull black each with four pale yellow spots. (p. 99) 3 Safranek’s Tiger Beetle, Calochroa safraneki Endemic to Maharashtra in western India. Head and thorax coppery. Elytra dark greenish-brown with one shoulder spot and three larger oval or round spots in a row on each elytron. (p. 100) 4 Black-headed Tiger Beetle, Calochroa fabriciana Endemic to southern India. Head and thorax shiny black to bronze. Elytra dull velvety black with narrow green sides and three large orange oval spots on each elytron. (p. 100) (Fig. 32.2) 5 Four-spotted Tiger Beetle, Calochroa assamensis Northeastern India to Myanmar. Head and thorax greenish-black. Elytra dull, dark greenish-black and purple, each large yellow shoulder spot and larger yellow spot at mid-margin. (p. 101) 6 Bicoloured Tiger Beetle, Calochroa bicolor India west to Pakistan and east to Thailand. Head and thorax metallic green to copper. Elytra dull black, dark indigo, or blue-green with or without three small spots arranged in a row along length of elytron. End of abdomen under elytra reddish-orange. (p. 101) 6a C. b. bicolor -- dull dark blue to green elytra with no light markings. Northeastern India. 6b C. b. atavus -- three large yellow spots on each elytron. Northwestern India to Pakistan. 6c C. b. haemorrhoidalis -- copper head and thorax; elytra black, each with small yellow shoulder spot and two large yellow spots along elytron. East central and southern India to Pakistan. 6d C. b. xanthospilota -- copper head and thorax. Elytra black, each with small yellow spots. Southwestern India and Sri Lanka. 232

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Beautiful Tiger Beetles III and Temperate Tiger Beetles I Beautiful Tiger Beetles, Calochroa 1 Orange-striped Tiger Beetle, Calochroa hamiltoniana Endemic to southern India. Body slender. Head and thorax green or bronze. Elytra green-bronze with narrow orange and black stripes lengthwise on each elytron. (p. 102)

Temperate Tiger Beetles, Cicindela Large number of species that share male genitalic characters as well as DNA molecules. Occur at higher latitudes in Asia, Europe and North America. All species have medium to large bodies, bulky heads with eyes less prominent. Elytra are generally dark and boldly marked with broad yellow or white bands or spots. 2 Granulated Tiger Beetle, Cicindela granulata Northern India to Kazakhstan. Head and thorax shiny black. Elytra dull black to purple-black with highly varied yellow markings. (p. 105) 3a Cobalt Tiger Beetle, Cicindela cyanea Northeastern India to Nepal and Bangladesh. Head and thorax dull black to shiny blue or green. Elytra dull blue or blackish green with no to small light markings. Some individuals have a wide lengthwise orange stripe down side border of each elytron (3b). (p. 105) 4 Yellow-shouldered Tiger Beetle, Cicindela angulicollis Endemic to southern India. Head and thorax black. Elytra dull black, each with three large yellow to white bands. Labrum black with large yellow spot in middle. (p. 106) 5 Shivah Tiger Beetle, Cicindela shivah Highly localized and endemic to central Western Ghats of southern India. Dull black above and below. Cheeks on head with hair-like setae. Thorax covered with dense dark hair-like setae. Elytra with shiny purple sides and three bold oval yellow spots on each elytron. Labrum all black (p. 106) 6 Dappled Tiger Beetle, Cicindela guttata Endemic to northern India. Head and thorax shiny dark green. Elytra dull dark green or black-green with bold crescent-shaped and rounded, yellowish markings. Cheeks on head with white hairs restricted to lower part. (p. 109) 7 Ghats Tiger Beetle, Cicindela andrewesi Endemic to Western Ghats of southern India. Slender body. Head and thorax shiny black. Elytra dull black with bold yellow-orange spots or longitudinal curved lines running length of each elytron. Cheeks on head with no white hairs. (p. 109) 7a C. a. andrewesi -- three oblong yellow spots along length of each elytron. Southern parts of Western Ghats. 7b C. a. zingaroana -- orange elytra markings coalesced into two lines running length of each elytron. Northern parts of Western Ghats.

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Temperate Tiger Beetles II and Margined Tiger Beetles I Temperate Tiger Beetles, Cicindela 1 Broad-striped Tiger Beetle, Cicindela princeps Endemic to northeastern and southern India. Head and thorax shiny black. Elytra dull black with bold, yellow band in front, broad yellow band across entire middle, and narrow yellow margin on rear. (p. 107) 1a C. p. princeps -- elytra surfaces dull with non-contrasting punctuation or no punctation on unbanded areas, yellow bands on elytra broad. Southern India. 1b G p. ducalis -- elytra surfaces with distinct punctuation on unbanded areas, yellow bands narrow. Central and northeastern India. 1c G p. pochoni -- yellow shoulder band and central band connected. Nilgiri Hills of Tamil Nadu. 2 Golden-banded Tiger Beetle, Cicindela aurofasciata Endemic to highlands of southern India. Head and thorax shiny black or dark bronze. Elytra black with front yellow bands pointing obliquely back to almost meet middle yellow bands that cross both elytra. (p. 108) 3 Lowland Tiger Beetle, Cicindela goryi Endemic to lowlands of southern India. Head and thorax shiny black or dark bronze. Elytra black with broad yellow bands across front and middle fused to form an “X”. (p. 108) 4 Transverse-banded Tiger Beetle, Cicindela calligramma Southern India and Sri Lanka. Head and thorax shiny copper, greenish copper, or green. Elytra dull green or brown, each with three separate broad, separated, or fused light yellow markings. Entire cheek with long white hairs. (p. 110)(Fig. 7.1)

Margined Tiger Beetles, Calomera 5 Brackish Tiger Beetle, Calomera littoralis Northwestern India to Ukraine. Head and thorax dark brown, black or greenish brown. Elytra dark brown with varied ivory-coloured markings that are variously connected or separated into as many as six large spots. (p. 112) 6 Beach Tiger Beetle, Calomera aulica Northwestern India to southern Europe and East Africa. Head and thorax brown, green, dark copper-green or nearly black. Elytra dark brown with copper reflections and bold ivory-coloured markings. (p. 113) 7 Sand Tiger Beetle, Calomera ungulata Throughout India west to Afghanistan and east to Japan. Male elytra slender, but female elytra significantly expanded at mid-body. Bronze to dark copper, occasionally green or nearly black. Elytra with standard pattern of ivory-coloured markings (p. 113)

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Margined Tiger Beetles II and Ornamented Tiger Beetles I Margined Tiger Beetles, Calomera 1 Riverine Tiger Beetle, Calomera fowleri Endemic to peninsular India. Male slender and female considerably stouter. Head and thorax dark brown to bronze with copper-green reflections. Elytra brown to copper with a disrupted pattern of thin, light markings often separated into spots and short lines. (p. 114) 2 Combed Tiger Beetle, Calomera plumigera India to Myanmar. Head and thorax bronze with copper-green reflections. Elytra dark brown to olive green or copper with standard elytral pattern of light yellowish markings often connected along elytra margin. Male with comb of bristles on antennae, (p. 115) (Fig. 32.3) 2a C. p. plumigera -- smaller, brighter copper, elytra with thinner markings on elytra. Southern India. 2b C. p. macrograptina -- larger, dark brown to olive-green, elytra with bolder markings. Northern India to Myanmar. 3 Riverbank Tiger Beetle, Calomera cardoni Throughout India and Sri Lanka. Brown to blackish brown. Standard elytral pattern but light markings interrupted. (p. 115) 4 Waterfront Tiger Beetle, Calomera chloris Northern India west to Afghanistan and east to Laos. Bright green, copper green, black green or blue. Elytra with three bold, white spots along each margin in rear halves, and surfaces distinctly granulate with punctures. (p. 116) 5 Forest Stream Tiger Beetle, Calomera funerea India west to Pakistan and east to Vietnam. Dull greenish black, green bronze or black. Elytra with three or four thin white spots along each outer margin and surfaces granulate with patches not punctate. (p. 117) 5a C. f. funerea -- copper-brown with four spots along each elytra margin. Eastern India to Vietnam. 5b C. f. assimilis -- greenish-black to green with three spots along each elytra margin. India west to Pakistan and east to China. 6 Anamalai Tiger Beetle, Calomera quadripunctulata Highly localized and endemic to Anamalai Hills of southern India. Head and thorax copper to violet-green. Elytra bronze or bronze green with two white spots along margins in rear half of each elytron. (p. 117)

Ornamented Tiger Beetles, Cosmodela 7 Iridescent Tiger Beetle, Cosmodela duponti Peninsular and northeastern india to Vietnam. Head and thorax greenish-purple. Elytra dull dark blue, black with green and purple iridescence on sides and along middle line separating elytra, each with four white spots. (p. 120) (Fig. 31.1)

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Ornamented Tiger Beetles II and Small-eyed Tiger Beetles Ornamented Tiger Beetles, Cosmodela 1 Intermediate Tiger Beetle, Cosmodela intermedia Northern India to Afghanistan. Head and thorax iridescent copper on top and blue-green on sides. Elytra olive-green to purple-green with five separate white spots on each elytron. (p. 119) 1a C. i. intermedia -- olive-green elytra with small dots. Northwestern India. 1b G i. chitwanae -- dark purple-green elytra with thicker white markings and middle ones joined to form broad band. Eastern Nepal and likely adjacent northeastern India. 2 White-spotted Tiger Beetle, Cosmodela fleutiauxi Northeastern India to China. Head and thorax greenish-purple. Elytra dull dark blue, black with green and purple iridescence on sides and along middle line separating elytra. Four white spots on each elytron, middle two often connected. (p. 120) 3 Twin-spotted Tiger Beetle, Cosmodela didyma HYPOTHETICAL for Nicobar Islands. Head dark greenish to black olive. Thorax blue-green to brassy reddish black. Elytra dull black with shoulders and all margins usually bright blue-green, each elytron marked with live yellow spots. (p. 121) 4 Goblet-marked Tiger Beetle, Cosmodela diehli Great Nicobar Island to Indonesia. Head greenish-red. Thorax glossy, bare, greenish coppery with distinct blue violet. Elytra dull black with narrow green-copper margins, inner edge and rear fringed with bright green, and each elytron marked with five spots. (p. 122) 5 Coppery-green Tiger Beetle, Cosmodela juxtata Northern India to Vietnam. Head and thorax metallic copper edged with green. Elytra greenish black with iridescent inner edges; each elytron with small yellow shoulder dot and three large, yellow spots in a row along its length. (p. 122) 6 Arched Tiger Beetle, Cosmodela virgula Northern India to Vietnam. Head and thorax metallic copper edged with green. Elytra greenish black with iridescent inner edges; each elytron with one small, yellow, shoulder dot, and three larger yellow spots, the middle one crescent-shaped, aligned along its length. (p. 123)

Small-eyed Tiger Beetles, Plutacia 7 Opulent Tiger Beetle, Plutacia dives Endemic to India. Head and thorax shiny green and copper red. Elytra velvety green with metallic copper edges and marked with three obliquely transverse bands. Male antennae without comb of stiff bristles. (p. 124) 8 Ridged Tiger Beetle, Plutacia notopleuralis Single specimen known only from northeastern India. Head and thorax black with sides shiny green. Elytra dull black with bold yellow markings at shoulder and middle. Male antennae each with comb of stiff bristles. (p. 125)

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Reticulated Tiger Beetles I Reticulated Tiger Beetles, Lophyra 1a Braided Tiger Beetle, Lophyra catena India west to Pakistan and east to Myanmar. Head and thorax copper on top and green or blue on sides. Elytra dull copper with intricate pattern of wide, yellowish-white lines that obscure background colour. Cheeks covered with thick mat of white hairs (1b). (p. 128) 2 Waxen Tiger Beetle Lophyra cerina Highly localized and endemic to coast of extreme southern India. Head and thorax bright copper-red. Elytra yellowish-orange with waxy lustre, each with five small blue spots along their length near midline. (p. 128) 3 Bald Tiger Beetle, Lophyra striatifrons India and Nepal. Head and thorax copper. Elytra blackish-green with three standard whitish markings supplemented by oval spots and stripes. Cheeks with white hairs. (p. 129) 4 Harlequin Tiger Beetle, Lophyra histrio Northwestern India to Saudi Arabia. Head and thorax copper. Elytra dull, copper-green with standard ivory-coloured markings plus “L”-shaped markings at front. Cheeks bare. (p. 129) 5 Lattice Tiger Beetle, Lophyra cancellata India to Vietnam. Head and thorax copper to copper-green. Elytra dull, dark copper-green, green or blue with standard ivory-coloured markings, light spots along inner edges ot elytra broken up into two or four short lines. Cheeks bare (5c). (p. 130) 5a L. c. cancellata -- copper head and thorax, and dull dark copper-green elytra with broad markings. Southern India to Sri Lanka. 5b L. c. intemperata -- copper green to blue head and thorax, and elytra with narrow markings. Northern India to Vietnam. 6 Belted Tiger Beetle, Lophyra striolata india to Philippines. Head and thorax copper above and bright copper-green, blue-green and purple on sides. Elytra velvety black or greenish black with white or yellow markings forming variable pattern of separated longitudinal stripes and bands that do not reach outer edge. Hairs on cheeks. (p. 130)

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Reticulated Tiger Beetles II, Woolly Tiger Beetles, and Tethys Tiger Beetles I Reticulated Tiger Beetles, Lophyra 1 Lined Tiger Beetle, Lophyra lineifrons Northern India to Vietnam. Head and thorax bright copper. Elytra velvety black, each with white, narrow markings at shoulder and small spots at middle and rear. Hairs on cheeks. (p. 131) 2 Tangled Tiger Beetle, Lophyra lefroyi Endemic to northern India. Head and thorax bright copper-red above and shiny blue-green on sides. Elytra velvety black, each with fourteen white lines and spots. Cheeks without hairs. (p. 132) 3 Tiny-spotted Tiger Beetle, Lophyra parvimaculata Northern India and Nepal. Head and thorax copper-brown above and shiny blue green on sides. Elytra velvety black or greenish-black with white or yellow markings reduced to circles and elongated spots. Cheeks with no hairs, and male with bristle comb on each antenna. (p. 132) 4 Plaited Tiger Beetle, Lophyra vittigera Northern India west to Pakistan and east to Bangladesh. Head and thorax dark copper to greenish copper. Elytra dull velvety green with ivory, many ivory lines and two spots towards rear. Cheeks with few hairs. (p. 133) 5 Many-spotted Eiger Beetle, Lophyra multiguttata Northern India east to Myanmar. Head and thorax dark metallic copper above and green and blue-copper on sides. Elytra greenish black with large yellow marking at shoulder and five or six smaller spots that often run together on each elytron. Cheeks distinctly but sparsely covered with hairs. (p. 133)

Woolly Tiger Beetles, Chaetodera 6 White Tiger Beetle, Chaetodera albina Northern India west to Pakistan. Head and thorax greenish-copper and covered by dense white, flat hairs (setae). Elytra almost completely covered by four broad whitish stripes with narrow intermittent dark copper areas. (p. 134) 7 Twenty-spotted Tiger Beetle, Chaetodera vigintiguttata Northeastern India west to Pakistan. Head and thorax copper above and violet on sides. Elytra black-green with ten oval and round yellowish spots on each elytron. (p. 135)

Tethys Tiger Beetles, Jansenia 8 Flightless Tethys Tiger Beetle, Jansenia westermanni Endemic to southern India. Head and thorax copper. Elytra oval-shaped, dull black edged with shiny copper, each with a medium yellow spot at mid margin and large patch at rear. Runs quickly but does not fly. (p. 44) (Fig. 7.7) 9 Green-spotted Tethys Tiger Beetle, Jansenia pseudodromica Highly localized and endemic to southern India. Head and thorax copper-brown. Elytra oval-shaped, dull black with copper margins, each with yellow central spot and two short longitudinal bands near margin at front and rear. (p. 145) 244

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Tethys Tiger Beetles II Tethys Tiger Beetles, Jansenia 1 Nathan’s Tethys Tiger Beetle, Jansenia nathanorum Highly localized and endemic to southern India. Head copper with green golden-green reflections. Thorax copper-bronze. Elytra dark bronze to shiny black with small and larger white dots on each elytron. (p. 145 ) 2 Shaggy Tethys Tiger Beetle, Jansenia dasiodes Endemic to southern India. Head and thorax copper and blue-green. Elytra shiny black streaked with copper or copper-green; each elytron with two longitudinal furrows, usually with two ivory spots on rear half. (p. 146) 3 Undulated Tethys Tiger Beetle, Jansenia biundata Endemic to southern India. Dark brown. Elytra surface rough with two longitudinal furrows down the centre of each elytron, and two light oblong marks along rear margin. (p. 146) 4 Bristled Tethys Tiger Beetle, Jansenia semisetigerosa Endemic to southern India. Head and thorax shiny copper and blue-green. Elytra shiny black with copper streaks, single longitudinal furrow on each elytron, and two small light spots along rear margin. (p. 147) 5 Waved Tethys Tiger Beetle, Jansenia vestiplicatica Endemic to southern India. Head and thorax bright copper and copper-green. Elytra shiny copper-black with a wide copper band and a narrow bluish green band running along outer margins; each elytron with indistinct longitudinal furrow on front half and two oblong light spots. (p. 147) 6 Band-lipped Tethys Tiger Beetle, Jansenia legnotia Endemic to southern India. Head and thorax shiny copper and copper-green. Elytra shiny black along raised centre contrasting with copper or copper-green along margin, and two light spots along rear margin of each elytron. (p. 148) 7 Streaked Tethys Tiger Beetle, Jansenia plagatima Endemic to southern India. Head and thorax shiny copper and copper-green. Elytra shiny black down centre with copper or copper-green along sides and two small white spots on rear half of margins. (p. 148) 8 Bengaluru Tethys Tiger Beetle, Jansenia bangalorensis Highly localized and endemic to southern India. Head and thorax shiny copper and blue-green. Elytra shiny copper-bronze; small whitish yellow spots on mid margin and rear of each elytron. (p. 149) 9 Purple-banded Tethys Tiger Beetle, Jansenia ostrina Endemic to central and eastern India. Head and thorax shiny copper and blue green. Elytra shiny black down centre and broad copper-green and purple band down sides, broad ridge on surface running length of both elytra with two whitish spots on each. (p. 149)

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Tethys Tiger Beetles III Tethys Tiger Beetles, Jansenia 1 Burnished Tethys Tiger Beetle, Jansenia corrugatosa Endemic to southern India. Head and thorax shiny copper, blue-green, and bronze. Elytra shiny black with indistinct bronze longitudinal band with narrow, purple edge and two white spots along rear of each; surface with no ridges or irregularities. (p. 150) 2 Pitted Tethys Tiger Beetle, Jansenia cratera Endemic to southern India. Head and thorax shiny copper with patches of bronze-green. Elytra copper to bronze in centre grading into diffuse copper green with a narrow purple edge, and two large, white marks on rear margin of each elytron. (p. 150) 3 Pale-legged Tethys Tiger Beetle, Jansenia choriodista Endemic to southern India. Head and thorax shiny copper and purple above and shiny copper to bronze on sides. Elytra shiny bright copper to green with wavy wrinkles on their surfaces and two large, white dots on rear half of margins. Upper parts of legs light with a tinge of green. (p. 151) 4 Purple-margined Tethys Tiger Beetle, Jansenia prothymoides Endemic to southern India. Head and thorax shiny copper-black, blue-green to violet. Elytra black to copper-black with a wide shiny purple band along sides; their surfaces rough and uneven with no light markings. (p. 151) 5 Speckled Tethys Tiger Beetle, Jansenia crassipalpis Endemic to southern India. Head and thorax shiny copper with green and red reflections. Elytra black with copper and blue-green reflections and a broad copper green band along their sides, each with three whitish spots. (p. 152) 6 Red-headed Tethys Tiger Beetle, Jansenia tetrastacta Northern and eastern India to Nepal. Head and thorax crimson above and shiny blue to green on sides. Elytra shiny dark black or copper black with a wide band of violet, blue, green, and red iridescence along sides with one or two white spots at rear of each elytron. (p. 152) 6a J. t. tetrastacta -- Eastern and northeastern India to Nepal. 6b J. t delhiensis --Union Territory near New Delhi. 7 Green-backed Tethys Tiger Beetle, Jansenia chlorida Endemic to southern India. Head and thorax green occasionally copper-green. Elytra uniformly dull green, each with a copper margin and two large whitish markings on rear margin. (p. 153) 8 Green-legged Tethys Tiger Beetle, Jansenia psarodea Endemic to southern India. Head and thorax shiny copper above with blue-green between eyes and on sides. Eytra copper-black with shiny black-bronze and purple band along notably rounded sides; two large yellow patches on rear margin of each. (p. 153)

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Tethys Tiger Beetles IV Tethys Tiger Beetles, Jansenia 1 Large-lipped Tethys Tiger Beetle, Jansenia rostrulla Endemic to southern India. Head and thorax green or copper. Elytra copper-brown with smooth surfaces and longitudinal row of distinct pits reflecting green. Metallic copper band along sides with two yellow spots on rear margin of each. Long, ivory-coloured labrum. (p. 154) 2 Blue-shouldered Tethys Tiger Beetle, Jansenia venus Endemic to southern India. Head and thorax reddish-copper with bright blue-green patches. Elytra reddish-copper with iridescent blue-violet band along sides widest at shoulders, two large yellow markings on each. (p. 155) 3 Blue-margined Tethys Tiger Beetle, Jansenia stuprata Endemic to southern India. Head and thorax shiny crimson above and bright green on sides. Elytra dull black-bronze with iridescent copper crescent at shoulders, iridescent blue along sides, and violet black along inner edges, pale yellow line across elytra at middle and a small spot at rear margins. (p. 155) 4 Spindle-necked Tethys Tiger Beetle, Jansenia fusissima Endemic to southern India. Head and thorax shiny crimson above with green and purple sides. Elytra shiny copper with black and purple band along edges, yellow-orange spot on shoulders, narrow pale yellow band across middle of elytron and small yellow spot at rear margin of each. (p. 156) 5 Twin-spotted Tethys Tiger Beetle, Jansenia chloropleura Northern India to Nepal and Bangladesh. Head and thorax brilliant copper-red above and metallic blue, violet and green on sides. Elytra copper brown or olive green with a narrow blue-green band along outer and inner edges, two yellowish spots along rear margin of each. (p. 156) 6 Green-sided Tethys Tiger Beetle, Jansenia viridicincta Endemic to southern India. Head and thorax bright reddish-copper above and shiny violet green on sides. Elytra reddish-copper with narrow green and violet stripe on outer edges and green stripe down inner edges, yellowish white markings. (p. 157) 7 Purple-sided Tethys Tiger Beetle, Jansenia azureocincta Endemic to southern India. Head bright copper-red with iridescent blue area between eyes. Thorax bright copper red above with purple band on sides and blue-green in middle. Elytra copper-red with outer edges iridescent purple and blue and inner edges blue, two whitish spots on rear half of each. (p. 157) (Fig. 7.3) 8 Crimson Tethys Tiger Beetle, Jansenia rugosiceps Endemic to southern India. Head copper-red with bright blue or green sides. Thorax purple and green. Elytra dull crimson with broad green central stripe branching to each side and blue inner and outer edges, two yellowish spots on rear third (p. 158) 9 Sandur Tethys Tiger Beetle, Jansenia sandurica Highly localized and endemic to southern India. Head and thorax bright copper-red above with metallic blue sides. Elytra dull copper-red with interrupted blue outer sides and complete blue band along inner edges; two large yellowish spots towards centre on rear of each elytron. (p. 158)

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Plate 23 Tethys Tiger Beetles V, Pygmy Tiger Beetles, and Lustrous Tiger Beetles Tethys Tiger Beetles, Jansenia 1 White-banded Tethys Tiger Beetle, Jansenia grossula Highly localized and endemic to southern India. Head and thorax dark bronze with violet reflections and dark blue-back sides. Elytra distinctly oval-shaped, dull dark bronze with a large whitish band extending almost entirely across mid elytra and a large spot at rear margin of each elytron. (p. 154) 2 Copper Tethys Tiger Beetle, Jansenia motschulskyana Endemic to southern India. Head and thorax shiny copper above and green on sides. Elytra dull copper with two dull black patches and shiny purple, green and copper along outer edges with one small whitish dot on rear. (p. 159) 3 Metal-lipped Tethys Tiger Beetle, Jansenia indica Endemic to western India. Head and thorax shiny black; surfaces wrinkled above and bright dark purple and blue on sides. Elytra bright copper along sides and rest of surface dull blackish bronze with raised velvety black areas and no light markings. (p. 159) 4 Carpet-backed Tethys Tiger Beetle, Jansenia reticulella Endemic to southern India. Head and thorax shiny black, heavily wrinkled above and shiny black or purple on sides. Elytra shiny bronze on sides and rest bronze-black with labyrinth of raised areas across surface but no light markings. (p. 160) 5 Bronze-spotted Tethys Tiger Beetle, Jansenia tetragrammica Endemic to southern India. Head and thorax dull black with shiny copper areas. Elytra dull, velvety black with faint copper or metallic green and distinctive large and circular greenish patches. Two large yellow spots on rear of each elytron. Large prominent eyes. (p. 160) 6 Flat-headed Tethys Tiger Beetle, Jansenia applanata Endemic to southern India. Head and thorax dull black with copper flecks. Elytra dull black with copper or metallic green shoulders and outer edges, metallic green patches; each elytron with two large yellow oval spots on rear half. Eyes small and barely protruding. (p. 161)

Pygmy Tiger Beetles, Glomera 7 Immaculate Pygmy Tiger Beetle, Glomera belloides Endemic to southern India. Head and thorax black, bronze, or copper above and blue-green and purple on sides. Elytra with glossy purple-black band along outer edge and broad shiny copper-green down centre, no light markings. No hairs on underside. (p. 162) 8 Pale-legged Pygmy Tiger Beetle, Glomera ochrocnemis Endemic to southern India. Head and thorax shiny bronze above and bluish-black to purple on sides. Elytra dull black with iridescent purple, blue or copper along sides and as lobes projecting toward middle, no light markings. Sparse hairs on underside at rear of abdomen. (p. 163)

Lustrous Tiger Beetles, Setinteridenta 9 Lustrous Tiger Beetle Setinteridenta rhytidopteroides Northern India to Myanmar. Head dark bronze and iridescent blue-green on sides. Thorax greenish-bronze to copper above and purple on sides. Elytra shiny dark bronze to copper brown with broad band of blue-green and purple along sides, and two ivory oval marks on rear half of each elytron. (p. 164) 252

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Rounded-thorax Tiger Beetles I Rounded-thorax Tiger Beetles, Cylindera 1 Dot-lipped Tiger Beetle, Cylindera paradoxa Southern India and Sri Lanka. Head and thorax bright green with violet reflections. Elytra shiny dark bronze to black with longitudinal row of green surface pits and two oval white spots on rear half of each elytron. Male labrum with large white centre. (p. 175) 2 Fluted Tiger Beetle, Cylindera lacunosa Southern India and Sri Lanka. Bronze. Elytra with polished raised areas and two oval white spots on rear half of each. (p. 175) 3 Slope-striped Tiger Beetle, Cylindera obliquefasciata Northern India west to Kazakhstan and east to China. Head and thorax metallic copper-green. Elytra dull black-green, each with two small spots and two distinct light bands. (p. 175) 4 Fleet Tiger Beetle, Cylindera dromicoides Northern India to China. Head and thorax bright copper to dull brown or green. Elytra with irregular, dark, velvety brown patches surrounded by bright bronze patches; each with two elongate white spots at mid-margin and rear. Sides of elytra widened noticeably at rear. Flightless. (p. 176) (Fig. 7.7) 5 Terrestrial Tiger Beetle, Cylindera delavayi Northern India to Laos. Head and thorax copper or green. Elytra dull bronze to blue or green, each with two velvety dark areas and two small yellow spots on rear margin. Sides of elytra parallel. Flightless. (p. 177) 6 Pitted Tiger Beetle, Cylindera foveolata India to Laos. Head shiny black and blue-green. Thorax shiny black and copper. Elytra shiny blackish bronze with no light markings. (p. 177) 7 Hump-headed Tiger Beetle, Cylindera cyclobregma Northeastern India to Vietnam. Head and thorax black, and head with distinctive but small bump between eyes. Elytra dull black with shiny black margin, noticeably expanded toward rear and with one small white dot on each. (p. 178) 8 Shy Tiger Beetle, Cylindera viduata Northern India to Papua New Guinea. Head and thorax copper. Elytra dull greenish black to copper-black; each elytron with narrow velvety black stripe running along inner edge and three small white spots. (p. 178) 9 Karl’s Tiger Beetle, Cylindera karli Highly localized and endemic to Andhra Pradesh in southeastern India. Head and thorax bronze. Elytra reddish bronze with green band along outer edges and upper surfaces surface covered with green punctures, four small whitish spots and ovals on each elytron (p. 179)

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Rounded-thorax Tiger Beetles II Rounded-thorax Tiger Beetles, Cylindera 1 Green-lipped Tiger Beetle, Cylindera viridilabris Northern India west to Pakistan and east to Cambodia. Head and thorax dark bronze. Elytra dark brown or black with shiny areas at shoulder and along sides, four thin to small yellowish spots and lines on each elytron. Labrum copper green. (p. 179) 2 Copper-lipped Tiger Beetle, Cylindera labioaenea Southern India to Sri Lanka. Head and thorax shiny copper and blue-green. Elytra dull copper-green to green with shiny blue-green shoulders and four small yellowish spots on each elytron. Labrum metallic bronze to copper. (p. 180) 3 Deceptive Tiger Beetle, Cylindera fallaciosa Northeastern India to Vietnam. Head and thorax copper. Elytra dull dark brown to black with shiny blue-green crescent on shoulders and four small whitish dots on rear two-thirds of each elytron. (p. 180) 4 Rough Tiger Beetle, Cylindera severini Endemic to peninsular India. Head and thorax shiny copper. Elytra dull brownish black except for a shiny blue green crescent on shoulders; four, small whitish dots in an alternating pattern on each elytron. (p. 181) 5 Channel-headed Tiger Beetle, Cylindera collicia Endemic to southern India. Head and thorax copper. Elytra dull dark copper-brown with iridescent blue-green crescent at shoulders and four small whitish spots on each elytron. (p. 181) 6 Four-dotted Tiger Beetle, Cylindera nietneri Highly localized in southern India to Sri Lanka. Head and thorax dull purple black. Elytra dull purple black with four small whitish spots on each elytron. (p. 182) 7 Shiny-shouldered Tiger Beetle, Cylindera belli Endemic to peninsular India. Head and thorax dull purple black. Elytra dull purple black with four small whitish spots on each elytron. (p. 182) 8 Purple-footed Tiger Beetle, Cylindera umbropolita Endemic to southern India. Head and thorax copper to blue green. Elytra shiny copper with dull black areas, shiny black edges and one or two small whitish spots on each elytron. Lower legs shiny purple. (p. 183) 8a C. u. umbropolita -- narrow shiny black edges to elytra. Southern Western Ghats. 8b C. u. lucidinigrosa -- wide shiny black edges that extend into centre of each elytron. Central Western Ghats.

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Rounded-thorax Tiger Beetles III Rounded-thorax Tiger Beetles, Cylindera 1 Pit-lined Tiger Beetle, Cylindera seriepunctata Northeastern India and Nepal. Head and thorax green bronze to bronze. Elytra dull green bronze to bronze with shiny bue-green edges and three whitish spots on each elytron. Blue-green labrum pale brown in centre. (p. 183) 2 Honey-faced Tiger Beetle, Cylindera melitops Highly localized and endemic to southern India. Head and thorax shiny green bronze to bronze. Elytra dull green bronze to bronze, shiny and broad blue green along edges, three whitish spots on each elytron. Labrum pale brown. (p. 184) 3 Black-lipped Tiger Beetle, Cylindera spinolae Northeastern India to Vietnam. Head and thorax bright copper. Elytra dull brownish with greenish inner edges and five or more small circular and irregular whitish markings on each elytron. Labrum shiny brownish black. (p. 184) 4 Striped-legged Tiger Beetle, Cylindera dartista Highly localized and endemic to Great Nicobar Island. Head and thorax dark copper and green. Elytra brownish with three separate markings. Legs pale. (p. 185) 5 Brown-lipped Tiger Beetle, Cylindera paucipilina Northeastern India to Laos. Head blue-green, purple, and copper. Thorax shiny golden. Elytra dull brownish with copper green and purple bands running along outer edge, and with three whitish spots and a thin line. Labrum pale brown. (p. 185) 6 Violet-sided Tiger Beetle, Cylindera limitisca Endemic to Northeastern India. Head blue-green, purple, and copper. Thorax shiny golden copper. Elytra dull brownish with copper band and wide purple strip along outer edge, copper-green along inner edge, and three whitish spots and a line. Labrum dark brownish black with pale brown sides. (p. 186) 7 Stream Tiger Beetle, Cylindera subtilesignata Northern India west to Pakistan and east to Malaysia. Head and thorax copper. Elytra dull, dark bronze with copper shoulder, shiny blue to green on outer and inner edges and whitish spots and narrow middle band. Labrum dark brownish black. (p. 186) 8 Ten-spotted Tiger Beetle, Cylindera decempunctata Northern India west to Pakistan and east to Vietnam. Head and thorax copper to copper green. Elytra dull, dark copper green or brown and five small yellowish-white spots on each elytron. Labrum pale brown with dark brown centre. (p. 187) 9 Shade-loving Tiger Beetle, Cylindera anelia Endemic to southwestern India. Head and thorax copper. Elytra dull, dark copper-brown with five whitish spots on each elytron and female elytra distinctly widened in middle. Labrum usually all dark. (p. 187)

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Rounded-thorax Tiger Beetles IV Rounded-thorax Tiger Beetles, Cylindera 1 Small-lipped Tiger Beetle, Cylindera sikhimensis Northeastern India and Myanmar. Head and thorax dark bronze-green. Elytra dull bronze; each elytron with three whitish spots. Labrum light brown. (p. 188) 2 Dark-spotted Tiger Beetle, Cylindera discreta Nicobar Island to Australia. Head and thorax black and green. Elytra blackish brown in middle with bright green margins, each with two spots and thin, rear band. (p. 188) 2a C. d. discreta -- widespread subspecies found on nearby Indonesian Sumatra; elytra blackish brown with five, widely spaced spots on each elytron. 2b C. d. nicobarica -- single specimen known only from Great Nicobar Island; dark green with small, white maculations reduced to shoulder dot, central oval spot and a thin, rear band on each elytron. 3 White-margined Tiger Beetle, Cylindera kaleea Northeastern India to Japan. Head and thorax shiny copper. Elytra dull brown to blackish with four whitish spots on each. Labrum highly variable in colour - dark to pale. (p. 189) 4 Mountain Stream Tiger Beetle, Cylindera albopunctata Northern India to China. Olive green to dull brown green above. Elytra shiny with six pale yellow markings on each elytron. Underside of body completely covered with white, hair-like setae. (p. 189) 5 Ragged Tiger Beetle, Cylindera sublacerata Northern India to Russia. Head and thorax shiny green or copper-green. Elytra dull green or copper-green, each with three white crescents. (p. 190) 6 Fragmented-band Tiger Beetle, Cylindera mesoepisternalis Northern India and Pakistan. Head and thorax shiny green. Elytra duller with three white bands all connected by a wide continuous stripe along lateral margin of each. (p. 190) 7 Clubbed Tiger Beetle, Cylindera bigemina India to Pakistan. Head and thorax copper. Elytra copper-green with thin yellowish lines and round spots. Male antennae with comb of bristles. (p. 191) 8 Slender Tiger Beetle, Cylindera procera India with no exact location documented. Head and thorax shiny copper bronze. Elytra dull copper-brown, each with small white spots and bands. No comb of bristles on male antennae. (p. 191)

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Rounded-thorax Tiger Beetles V Rounded-thorax Tiger Beetles, Cylindera 1 Dwarf Tiger Beetle, Cylindera brevis Northern India to Afghanistan. Head and thorax shiny copper-green. Elytra dull copper-brown to green with whitish spots and thin bands. Male with no comb of bristles on antennae. (p. 192) 2 Little Tiger Beetle, Cylindera minuta India to Philippines. Head and thorax shiny copper or copper-green above. Elytra dull green and copper, each with three thin ivory-coloured lines. (p. 192) 3 Long-toothed Tiger Beetle, Cylindera cognata India west to Pakistan and east to Bangladesh. Head and thorax copper with green reflections. Elytra dark black-green with broad whitish lines and dots. (p. 193) 4 Erudite Tiger Beetle, Cylindera erudita Northern India to Afghanistan. Head and thorax copper. Elytra dull deep blue-black with shoulders and front part shiny green with bold white markings. (p. 193) 5 Dune Tiger Beetle, Cylindera agnata India to Myanmar. Head and thorax shiny copper-brown above with green reflections. Elytra dull brown to greenish with yellowish-white lines. (p. 194) 6 Fish Hook Tiger Beetle, Cylindera ancistridia Endemic to northern India. Head and thorax shiny copper. Elytra copper green, each with continuous whitish line along edge and connected to three broad whitish lines. Hook-shaped shoulder mark. (p. 194) 7 Wavy-lined Tiger Beetle, Cylindera venosa Northern India to Myanmar. Body shiny green with copper reflections. Elytra with complex pattern of thin, connected, white lines. (p. 195) 8 Mudflat Tiger Beetle, Cylindera grammophora Northern India to Bangladesh. Head and thorax green with copper reflections. Elytra dull dark green with three unconnected whitish markings on each. (p. 195) 9 Bar-shouldered Tiger Beetle, Cylindera singalensis Southern India and Sri Lanka. Body shiny bronze-green. Elytra with varied and irregular pattern of fragmented, whitish markings. (p. 196)

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Hook-legged Tiger Beetles and Hairy-sided Tiger Beetles Hook-legged Tiger Beetles, Myriochila 1 Yellow-legged Tiger Beetle, Myriochila fastidiosa India west to Pakistan and east to Myanmar. Head and thorax shiny hlue-green or copper. Elytra dull greenish bronze to copper with thin yellowish markings. Legs pale to greenish. (p. 198) 1a M. f. fastidiosa -- elytra markings separate and distinct. Upper legs with metallic reflections. Throughout India except north. 1b M. f. leucoloma -- all elytral markings joined by a continuous band. Upper legs light coloured with no metallic reflections. Northern India. 2 Strife Tiger Beetle, Myriochila litigiosa Northern India to Pakistan. Head and thorax shiny blue-green or copper. Elytra dull greenish bronze to copper with yellowish markings. Legs generally all light coloured. (p. 199) 3 Rippled Tiger Beetle, Myriochila undulata India to Pakistan. Head and thorax shiny green. Elytra dull green with thin whitish markings. (p. 199) 4 Wavering Tiger Beetle, Myriochila dubia Northeastern India to Thailand. Shiny greenish bronze with purple punctures. Elytra markings include a complete “C” -shaped shoulder band. (p. 200) 5 Imperfect Tiger Beetle, Myriochila atelesta Northern and northeastern India to China. Dark brown with copper reflections. White elytra markings include a white spot above shoulder. (p. 200) 6 Melancholic Tiger Beetle, Myriochila melancholica India west to Europe and east to China. Body dull, greenish with copper reflections. Elytra markings pale and narrow. (p. 201) 7 Distinctive Tiger Beetle, Myriochila distinguenda Southern India and Sri Lanka. Copper-green to green. Elytra dull brown to black with thin whitish markings. (p. 201)

Hairy-sided Tiger Beetles, Salpingophora 8 Hairy-legged Tiger Beetle, Salpingophora maindroni Northwestern India and Pakistan. Head and thorax shiny dark bronze to bronze green. Elytra bronze green with broad and irregular tan band along their outer edges. (p. 202)

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Plate 30

Flat-eyed Tiger Beetles, Thick-thighed Tiger Beetles, and Enigmatic Tiger Beetle Flat-eyed Tiger Beetles, Hypaetha 1 Four-lined Tiger Beetle, Hypaetha quadrilineata Coasts of India west to Iran and east to Malaysia. Head and thorax copper with some green reflections. Elytra dark greenish bronze with two wide, yellow-white bands running length of each elytron. (p. 203) 2 Asian Coastal Tiger Beetle, Hypaetha biramosa Coasts of India, Andaman and Nicobar Islands east to Indonesia. Dark bronze or bronze-green, ivory white border along outer edges of elytra bulging inward at middle and at rear. (p. 204) 3 Sandspit Tiger Beetle, Hypaetha copulata Northwestern India to Saudi Arabia. Shiny bronze with green reflections towards sides. Broad whitish band along entire outer margin of elytra. (p. 204)

Thick-thighed Tiger Beetles, Callytron 4 Side-banded Tiger Beetle, Callytron gyllenhalii Western India to Pakistan. Shiny greenishbronze. Elytra with or without a thin white border running along entire outer edge. Female elytra ovalshaped. (p. 205) 4a C. g. gyllenhalii -- thin white border around outer edges of elytra. Northwestern India. 4b C. g. immarginatum -- no white border around outer edges of elytra. Unknown locality but perhaps central west coast of India. 5 Dull Tiger Beetle, Callytron limosum East coast of India to Thailand. Body dull green with copper reflections. Elytra with narrow, white band along entire outer edge. (p. 206) 6 Spotless Tiger Beetle, Callytron malabaricum West coast of India to Pakistan. Head and thorax bronze above with blue-green sides. Elytra dull bronze almost black with no light markings. (p. 206)

Enigmatic Tiger Beetle, Apteroessa 7 Enigmatic Tiger Beetle, Apteroessa grossa Highly localized and endemic to southern India. Black above on all surfaces. Head and thorax large and about same width. Eyes small. Elytra oval, their surfaces covered with hair-like setae; three large yellow spots on each elytron. (p. 207)

266

Plate 30

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Plate 31.1 Iridescent Tiger Beetle (Cosmodela

Plate 31.2 Cross section of vertical larval

duponti) exhibiting shiny iridescent colours.

tiger beetle tunnel with larva at the bottom. Photograph by H. Ashida.

Photograph by Kesavamurthy, Birdwing

Plate 31.3 Robber fly (Family Asilidae) with tiger beetle prey after it was caught in flight. Photograph by E. S. Ross.

268

Mate 32.1 Montane Glossy Tiger Beetle (Pronyssa Plate 32.2 Black-headed Tiger Beetle (Calochroc nontanea) on leaf after landing on undergrowth fabriciana) on orange substrate that matches )ush to escape danger. Photograph by Vivek orange spots on its elytra and which disrup >arkar.

its body shape to confuse visual predators Photograph by Nagaraj Shastri.

Ecology and Behaviour

Physical Habitats Despite the large diversity of tiger beetle species occurring in India, there are only about 10 distinctive habitat types into which all the species can be categorised based on where adults and larvae exist. A few examples of these types include muddy pond edges, ocean beaches, moist forest floors, and boulder faces. Most tiger beetle species are limited to only one of these habitat categories. However, adults of a few species, such as the broad-ranging Melancholic Tiger Beetle (Myriochila melancholica), occur in as many as four or five different habitat categories. Some species broaden their use of habitats by temporarily expanding into different adjacent ones during the height of the rainy season. Some subspecies, such as those of the wide spread Yellow-legged Tiger Beetle (Myriochila fastidiosa), each occupy distinctly different types of habitat. This could

either indicate a broad ecological role for the species (niche) or that it subspecies are in reality separate species, even though inseparable on the basis of body features, a phenomenon called cryptic species. Larvae tend to be even more restricted to habitat types than are the adults of

the same species. For both adults and larvae, however, the habitat type occupied by each is influenced by numerous components. These include physical, chemical and

climatic qualities, such as soil composition, moisture, temperature, and chemistry, vegetation cover, seasonality, as well as, food supply. In addition, other habitat characteristics are important for providing mating and oviposition sites, and hiding places from parasites and predators. The ensemble of special adaptations that each species exhibits controls the unique range of habitat types it can occupy, thereby restricting where that species occurs within any given area to those sites providing for its survival and reproduction.

Temperature is a critical and general part of the habitat for tiger beetles. Tiger beetles are largely dependent on external sources of temperature (ectothermic)

to maintain their internal body temperatures so that they can be active. Because adult tiger beetles engage in much running and flying, they maintain internal body temperatures that are just below their lethal limit of 39°C. High internal temperatures below that limit allow for maximum speed and movement. With lower internal temperatures, adults become sluggish and are less likely to escape enemies,

chase down mates, or capture prey. On the other hand, internal temperatures that are too high cause water balance problems, reduce gamete production, and affect general metabolism. We know, for instance, that different species of tiger beetles in the same locality can be active at and tolerate a different range of maximum

h Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

and minimum temperatures. These different physiological capabilities in the same

microhabitat may force species to be active at different times of the day and thus avoid competitors and predators whose temperature regimes are not synchronized with those of the tiger beetle. The influence of temperature on timing of adult tiger beetle activity may also explain some of the differences in geographical range for some species.

Although physiology and structural features such as body size and body colour can be important for adjusting internal temperatures, various adaptations in behaviour

also have an influence on thermoregulation. To regulate internal temperatures, adult tiger beetles assume different and predictable positions with

their bodies. Early in the day as temperatures warm, adults extend their long legs (Fig. 7.1) (stilting), placing their body above the thin layer of hotter

Figure 7.1 Transverse-banded Tiger Beetle (Cicindela calligramma) with legs extended in a stilting behaviour to raise its body above the hot substrate. Photograph by Mohith K. Udupi.

air right next to the soil surface. As temperatures continue to warm, this boundary layer of hot air expands and the beetles combine stilting with an inclined orientation of their bodies toward the sun

(sun facing) that exposes only the front of the head to the sun's direct rays. When temperatures raise enough to make even these behaviours ineffective for controlling

internal temperatures, the beetles take advantage of evaporative cooling by seeking out wet substrates, or digging burrows in cooler substrates, or by simply becoming

inactive in shaded places. Running in and out of shaded areas (shuttling) is another common temperature-controlling behaviour. To regulate body temperatures on cool mornings, some species will crouch down to contact the warmer soil surface

with the underside of their body.

For larvae, placement of their burrows in the shade of vegetation, within stems of bushes, or raised on chimney-like turrets (Fig. 5.4) also helps regulate internal temperatures. Extra deep burrows make it possible for some high-elevation larvae to pass the winter below the frost line (Plate 31.2).

Most adult tiger beetles are active during the day, but some species of desert and tropical habitats are active at night, presumably to help escape extreme daytime temperatures. Other species forage actively during the daylight hours but mate or deposit eggs at night. Adults of some species that are normally active during the day are regularly attracted to lights at night.

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Ecology and Behaviour

Flooding is another important habitat factor, especially for the many species found in various water edge habitats. On the one hand, these habitats attract abundant prey for both adults and larvae, but just as importantly, they usually provide suitable egg-laying sites (oviposition). However, frequent or extreme flooding may harm the relatively immobile larvae if they are flushed from their burrows or their habitat is adversely altered or even destroyed. Larvae of some shoreline species can

survive when kept under water for as long as 6-12 days. Preliminary laboratory tests show that they are able to reduce their metabolism by as much as 90% by cutting back on cellular biochemical reactions that use oxygen. They may also be able to breathe air trapped in their closed tunnels. Even with these adaptations for surviving flooding, however, the larvae will have lost valuable development time that could adversely influence their survival in the short term in that habitat.

Seasonal Cycles Adult tiger beetles generally are active neither in cold weather nor hot, dry weather. They need moisture and warmth to support their own physiology as well as have

enough prey to survive and reproduce. For parts of the year, much of India is either

too cold or too hot and dry for adult tiger beetles to be active. In late April to July, the prevailing winds over the subcontinent switch to blow from the southeast off the Bay of Bengal. This moisture laden air meets the hot air over the land and rises to produce intermittently heavy rains throughout the summer into early fall. The majority of tiger beetle species in India become active within days after the first significant summer monsoon rainfall. By September the prevailing winds shift to come out of the northwest and bring

moisture from the Arabian Sea. This causes an October-December rainy season for

the extreme southeast of the Deccan (including the Madras coast), which is still warm

enough to force up the incoming moist air so that it condenses. Numerous species

of tiger beetles endemic to this part of India are only active during the fall monsoon.

The major exceptions to this dependence on monsoonal rainfall for seasonal activity are found among species that occur on coastal beaches, such as the Brackish Tiger Beetle (Callytron limosum) and the Four-lined Tiger Beetle (Hypaetha quadrilineata). Presumably because moisture conditions, temperatures, and

prey availability are more constant throughout the year regardless of rainfall patterns, the activity of these species is not so restricted to any particular season with abundant rainfall. The Melancholic Tiger Beetle (Myriochila melancholica) provides another example of a species whose adult activity is not tied to any particular season. It is one of the most widespread species of tiger beetle in India

and tolerates a wide range of habitats regardless of rainfall. Its broad adaptations to habitat types is probably also involved in its large and expansive distribution

across Asia to southern Europe.

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Mating Behaviour and Egg Laying Adults of some species begin reproduction

soon

after

emergence from their pupal stages, but adults of some species delay sexual activity until the following summer.

A few observations suggest

that males of some species emerge before the females. To begin the mating process, a

male approaches a female in

intermittent sprints similar to

Figure 7.2 Male and female Brush Tiger Beetle (Neocollyris)

those used in pursuing prey.

in coitus. Photograph by V. C. Balakrishnan.

When he gets close enough,

he leaps onto the back of the female and grasps her elytra with his middle and front legs. For many species, the male also grasps the female thorax with his mandibles, but in some species such as the Brush Tiger Beetles (Neocollyris), their body anatomy and small mandibles makes this impossible (Fig. 7.2). The male will often struggle with his target as she attempts to throw him off her back. Only if a male can successfully hold on, does he have a chance of successfully fertilizing the female's eggs. Tiger beetles have several specialized structures to aid in successful mating. Males of almost all species of tiger

beetles have the bottom surfaces of several tarsal segments of the front legs thickly covered with pads of hair-like setae (Fig. 2.3) that apparently aid in grasping and

holding the female. His hind legs remain on the substrate, and he uses them to walk along as the female moves

Figure 7.3 Pair of Purple-sided Tethys Tiger Beetles (Jansenia azureocincta) with male riding female in mate guarding behaviour. Photograph by Milind Joshi,

around. Females have species specific grooves and indentations (coupling sulcus) in the rear part of the sides of the thorax. One theory is that these grooves provide purchase or a hold on the female thorax for the mandibles of a male of the same species to tightly and advantageously grasp the female. After mating, the male may hold on with his mandibles and legs and continue to ride the female for as long as several hours (mate guarding) (Fig. 7.3). This behaviour serves to exclude other males from fertilizing her eggs. 274

Ecology and Behaviour

Most of the time tiger beetles only mate successfully and have offspring with members of their own species. However, in some instances closely related species may overlap in geographic range and mate with each other. These matings can result in hybrid offspring. Occasionally males of one species will attempt to mate

with a female of a distantly related species, but this union would be unlikely to result in genetically viable offspring. After the egg is fertilised, the female injects it into the soil (oviposition). To test a site before depositing an egg, the female uses her antennae and mandibles. If conditions are appropriate in terms of soil texture, salinity, slope, moisture, and

temperature, she extends her ovipositor and then turns her body almost vertical. With abdominal thrusts and a digging-cutting action of the end of the ovipositor, she inserts her ovipositor from several millimeters to more than a centimeter below the surface and injects a single ovoid-shaped egg. She then removes the ovipositor

and covers the hole with the surface substrate so that no disturbance is obvious. Recent studies have found that females of some species oviposit while they are in adult burrows. She can oviposit 10 to 20 eggs in a day in captivity, but few counts of actual numbers of eggs laid in the wild have been made. The emerging larvae construct their burrows at the site chosen by the female for oviposition. The larvae go through three developmental stages (instars) in which they

molt into successively larger bodies. The larval life ends when the third instar larva

enlarges its burrow and turns into a transformational stage (pupa). In this stage, the adult body structures grow and develop. Finally, the fully formed adult beetle

emerges from the pupa to start the life-cycle over again. The length of time from egg to adult is variable and depends on food availability and weather. Most species have a two-year life cycle, but some species develop in one year while others require

three or more years.

Predation and Parasitism For adult tiger beetles the

most important predators are robber flies (Plate 31.3), lizards,

and birds. Some birds (kestrels and flycatchers) and robber flies

generally

catch

adult

tiger beetles in the air during flight. Lizards and some bird predators like shrikes catch tiger beetles on the ground

before they can fly away. In some areas, spiders, scorpions, other arachnids and predatory

bugs

can

be

Figure 7.4 Adult tiger beetle with ant head attached onto antenna. Photograph by C. R. Brown.

important 275

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

predators on adult tiger beetles. Adult tiger beetles commonly have attached to their legs and antennae the heads of ants (Fig. 7.4), which may indicate that groups of attacking ants regularly overpower and kill tiger beetles. Enemies of larval tiger beetles include ground-foraging birds, ants, and provisioning wasps, but most important are attacks by parasitoid wasps and flies. While predators capture, kill and eat their prey, parasitoids instead lay their eggs on

or near the prey, and the larval parasitoids then eat the prey item when they emerge from their eggs. Many species of the worldwide genera of ant-like wasps Methoca and Pteromborus (Tiphiidae) specialized locating ground-dwelling tiger beetle

larvae in their tunnels. These small, female wasps sting and paralyze the tiger beetle

larva.Then the female deposits her egg on the immobilized larva, plugs the chamber and fills in the top part of the tunnel with soil. The larval wasp hatches in 4-5 days, consumes the tiger beetle larva, and emerges as an adult wasp.

Another major parasitoid of larval tiger beetles are bee-flies of the genus Anthrax (Bombyliidae). Females of these flies hover over the tunnel entrance or settle on the ground nearby, but just out of reach of the larva's mandibles. The fly turns her abdomen under and flips eggs at the tunnel entrance.

The eggs that fall into the tunnel then roll to the bottom, where the fly larva

(maggot) soon hatches out. The parasitoid crawls onto the larval tiger beetle, attaches itself to the underside or top of the abdomen or thorax (Fig.

7.5), and stays there until the

beetle reaches its pupal stage. During this stage, the tiger beetle is defenceless, and the fly larva consumes it.

Figure 7.5 Third instar tiger beetle larva with larval bee fly parasitoids {Anthrax sp.) attached to its back. Photograph by S. Spomer.

Other non-lethal parasites that use adult tiger beetles for transportation to new sites (phoretic) include mites of the family Trombidiidae. These small arachnids are

typically bright red and immobile and can be seen attached to the legs or thorax of a tiger beetle.They are most common in the warm tropics.

Antipredator Defences Excellent vision to detect danger coupled with quick escape flights and fast running are the primary defences for most adult tiger beetles. Many forest floor species when disturbed or threatened fly up to land on leaves of undergrowth bushes to escape danger (Plate 32.1). Defensive chemicals, such as benzaldehyde and cyanide, are released against some predators, such as robber flies. Larger body size deters smaller

predators. Adults of most diurnal species take refuge at night protected in burrows 276

Ecology and Behaviour

or under detritus and vegetation. Most nocturnal species spend the day resting in these same types of refuges. An interesting behavioural pattern of mixed species communal aggregation

(roosting) was discovered among 10 tiger beetle species in riparian areas of the Shivalik Hills of north western India. Up to 5000 individuals crowded together on a few undergrowth plants along river banks (Plate 32.3).The most common participants in these communal roosts involved several closely related species, the Combed Tiger Beetle (Calomera plumigera), the Sand Tiger Beetle (Calomera angulata), and the Waterfront Tiger Beetle (Calomera chloris). This communal roosting behaviour

was seen primarily in the early monsoon season and associated with overcast conditions, decreased temperatures, and reduced precipitation. The function of this behaviour most likely protects the adults from predation, but perhaps influences thermoregulation as well.

Sounds also play a role in tiger beetle behaviour. Many tiger beetles produce high frequency sounds by rubbing body and wing parts together (stridulation) in order to distract or alarm predators. Sounds produced by pursuing predators, such as high-pitched ultra-sounds used by bats for echo locating flying prey, can be perceived by the sensitive ears of tiger beetles to avoid them. These ears (tympana) are rare among other beetles. On tiger beetles they are located on the top of the abdomen and under the base of the elytra.

Camouflage is another widely observed anti-predator mechanism. Elytral colour patterns of most adult tiger beetle species closely match the colour of the soil substrate on which that species characteristically occurs. In addition,

the pattern of lines and spots (maculations) on the elytra can serve to disrupt the perceived shape and form of the tiger beetle (Plate 32.2). Other species have colour patterns that look like inanimate objects, such as pebbles or wood detritus, common in their habitat. Mimicry

is

an

evolved

superficial resemblance, usually between two unrelated species. Often, mimicry

functions to protect a species from predators by protecting a harmless species (mimic) that looks like and is mistaken

for a noxious species (model). Different tiger beetle species apparently serve as models and mimics. In parts of Southeast Asia, a young katydid (Order Orthoptera) (Fig.7.6) closely resembles an adult Arboreal

Figure 7.6 Late immature stages of the arboreal grasshopper (Orthoptera: Tettigonidae) Leptoderes omatipennis that closely resembles adults of the Arboreal Tiger Beetles (Tricondyla). Photograph by E. S. Ross.

Tiger Beetle (Tricondyla). 277

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

The

Flightless

adult

Tethys Tiger Beetle (Jansenia westermanni) of south India

(Fig. 7.7) has an elytral pattern that is remarkably similar to

that of a distinctly patterned female

velvet

ant

(family

Mutillidae) species with a potent stinger. Not only do they look similar, but they

both run on soil substrates in similar stops and sprints. Some of these velvet ants even produce stridulation sounds

Figure 7.7 Flightless Tethys Tiger Beetle (Jansenia westermanni) resembling in shape, colour, and running behaviour a female Velvet Ant (Hymenoptera: Mutillidae). Photograph by Nagraj Veeraswami.

and defense chemicals similar to their counterpart tiger beetles.

Larval tiger beetles rely primarily on a few behavioural responses to escape or avoid their enemies. Most commonly they quickly retreat from the top of the tunnel to hide at the bottom. In addition, if a predator is able to grab a larva in

its tunnel, the hooks on the back of back of the larva (Fig. 5.1) dig into the side of the tunnel and make it difficult for a predator to physically pull the larva out. The depth of the tunnel, curves in its course, and hard clay, sandy or rocky soil can

hinder digging attempts by predators. As a last recourse, a larva will abandon a tunnel to escape an enemy, scuttle across the soil surface using its short legs and undulating body movements to find a suitable site where it quickly excavates

another tunnel.

Larvae detect danger primarily through vision. Studies have revealed they possess fine focusing visual acuity. The larval eyes are important for both locating potential prey items and accurately detecting danger. The larvae may also use vibrations through the ground to detect larger predators.

Competition Competition occurs when one individual uses so much of a critical resource that another individual gets much less than it needs to survive. The critical or limiting resource can be space, egg-laying site, water, food, mates, or any number of other factors. Individuals engaging in competition can be of the same species (intraspecific) or of different species (interspecific). Numerous studies around the world show that food is a common resource for which tiger beetles compete, and thus feeding behaviour can be critical to understanding which tiger beetles can

survive together in the same habitat.

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Ecology and Behaviour

Feeding behaviour Tiger beetle larvae wait at the opening of their tunnels for arthropod prey to

approach within striking distance (attack zone). To capture the prey item, a larva rears backward up to half of its length out of the tunnel and grasps the prey with its mandibles (Fig. 5.3). Larvae of some species dig cone-shaped depressions in this

attack zone, apparently to increase the probability of capturing prey struggling to get out of the depression. Others construct turrets above the tunnel opening that function, at least partly, to provide an attractive landing spot for potential prey. When the prey item is successfully captured, the larva pulls it to the bottom of the tunnel and eats the digestible portions. Larvae attack and eat most types of prey except those with noxious chemicals and those too large to be quickly subdued and dragged to the bottom of the tunnel. After eating the digestible parts of the prey,

the larva carries the indigestible remains to the mouth of the tunnel and throws them backward away from the tunnel opening. Adults are visual predators and capture a wide variety of living arthropods.Though they have excellent sight, when they chase prey, they run so fast they can no longer see where they are going. Thus, adult tiger beetles use a pursuit pattern that involves active running interspersed with a pause-and-look behaviour. Alternatively, the tiger

beetle waits in a shaded area and ambushes approaching prey. The tiger beetle then grabs the prey item with its mandibles. Other adult tiger beetles frequently scavenge dead organisms, fallen fruits, and occasionally organic waste, such as, noodles.

Recently researchers discovered that, unlike most other insects that wave their

antenna around as feelers to acquire information, tiger beetles hold their antennae

rigidly and directly in front of them to mechanically sense their environments and avoid obstacles while running.

Adaptations reducing competition If populations of tiger beetle two species compete for the same food, one early outcome results in the better competitor being so successful it captures most of

the available prey. The second species of tiger beetle can no longer continue in the

area and its population is forced to move to a new region or it goes extinct. This reaction of the poorer competitor population to that of the dominant competitor (competitive exclusion) is difficult to observe in the field. In most cases, we see only the result and rarely witness the actual process of exclusion itself. For species that occur together in the same microhabitat and at the same time, their competition may also become reduced by dividing up the prey resource. The basis for how tiger beetle species divide the prey resource in their habitat has been studied in many locations with the same results; adult mandible length has been found to play a prominent role. Mandibles of different tiger beetle species can vary in number of teeth, curvature, and length (Fig. 7.8). Of these characters, however, mandible length seems to be the most important factor. 279

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

The

mandibles

of adult tiger beetles are highly correlated

with the size of prey captured and eaten. Small mandibles are not effective in grabbing

or subduing large prey items. Large mandibles, in contrast, can handle large prey items, but they are clumsier at processing

Figure 7.8 Representative mandibles (at same scale) of a large and a small tiger beetle species differing in number, curvature, and lengths of their teeth. A, Cicindela sp., B, Cylindera sp.

small prey. Thus, several tiger beetle species occurring

together in the same microhabitat at the same time of year and hour of the day could still reduce the impact of competition for food if each species had different­ sized mandibles to help maintain the associated differences in prey preference among the species. Indeed, in most habitat types where food is not abundant, tiger beetle species occurring together tend to have different-sized mandibles.

A typical example of mandibles influencing tiger beetle species diversity can be found in south India. Along river banks there it is usual to find together four common species, the Lattice Tiger Beetle (Lophyra cancellata), the Riverbank Tiger Beetle (Calomera cardoni), the Little Tiger Beetle (Cylindera minuta) and the Sand Tiger Beetle (Calomera angulata). These four species apparently reduce their interspecific competition for food both by having different sized mandibles to capture prey and

by using different microhabitats along the river banks.

However, there are also examples of similar-sized tiger beetle species that use the same food and occupy the same microhabitat, but they usually are active at

different times of the day. Nocturnally active species may thus potentially reduce competition from diurnally active species in the same habitat. Larvae of different species in the same area sometimes separate activity by season, thereby, perhaps reducing competition.

280

Conservation and Economic Impact

8

Because their taxonomy, morphology, physiology, behaviour, and distribution are

so well known in even remote parts of the world, tiger beetles lend themselves both to basic natural history studies, as well as, to applied scientific areas, such as, crop

protection and conservation efforts. This combination of utility, access, and deep background data have made tiger beetles a valuable tool for investigating and

ultimately understanding a broad range of basic and applied biological questions

in India.

Conservation Largely because of the cooperative efforts between passionate amateur collectors and dedicated professional taxonomists over the past two centuries, a more complete understanding of tiger beetle classification has developed and remains relatively stable. The vast majority of species encountered worldwide, even in such remote parts of the world as Sulawesi, New Guinea, and the Sudan, can be identified. These days, it is relatively easy for inexperienced helpers and students to recognize tiger beetles to species in a given area compared to learning most other insect groups in the same area. These amateurs have no trouble applying that knowledge

to reliably census the tiger beetle species diversity within any designated area.

Moreover, the work itself has proven to be faster because students of tiger beetles can quite easily census an area during the season of adult activity and reliably find most of the species within a short time, even in such complex and species-rich

habitats as tropical forests. An example of this increased efficiency in a species diversity census across several major faunal groups has been documented at one

particular site. Ornithologists took almost five years of intensive work to document 90 percent of the bird species occurring there, while in the same area butterfly and dragonfly workers took two or three years to arrive at this level of knowledge for their respective taxa. By contrast, those of us looking for tiger beetles found 90 percent of the fauna within the first fifty-five hours of searching. Their presence and level of abundance become quickly and reliably apparent. Around the world tiger beetles are among the few insect groups for which endangered species can be declared with certainty, and therefore, meaningfully placed on national red lists. In the United States, Israel, Bolivia, Spain, and Sweden,

entire populations of some tiger beetle species have officially been declared threatened, primarily due to suitable habitat destruction. Those of us working in conservation cannot afford the time and expense of defending false claims of rarity. The reliability and accurately of tiger beetle species diversity censuses have produced

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

favorable results by minimizing the uncertainty about detectability that haunt conservationists studying many other taxa that have proven difficult to observe. By protecting threatened populations of tiger beetles, we also secure habitat for many

other species, often called the umbrella effect. We have strong evidence that, at large spatial scales, the species richness of tiger beetles is a good predictor of the species richness of other, harder-to-census taxa, such as butterflies and birds. And because the number of species in a given locale can be so quickly determined, we can census hundreds of hectares for tiger beetles in the time it would take to census one hectare for birds or butterflies. Tiger beetles thus make excellent bioindicators to monitor diversity. Bioindicators also play a role in the early detection of habitat degradation and

monitoring restoration progress. Because most tiger beetle adults and larvae are specialised in habitat use, they tend to be highly sensitive to minor changes.They can function as barometers of degradation that might imperil them and their habitats. They can also illustrate the step by step success of a habitat during its recovery.

Collections made long ago provide valuable evidence for comparing the historic distributions of tiger beetle species with their current geographic ranges; tiger beetle records on museum specimens accumulated over the last century and

a half and documented in publications can established habitat changes that might not otherwise have been obvious.

Agricultural Pest Control In search of an economically viable alternative to expensive and often dangerous pesticides, several studies in India have investigated the presence of predators like

tiger beetles as a natural pest control agent in such crops as cauliflower, sugarcane, tea, and rice. Most of these studies, however, only assumed that the presence of predatory insects in these crop fields would have some impact on controlling pests. To date only a few studies in India have attempted to measure how much the presence of predatory insects actually reduces the pest populations. Another study also tested the influence of pesticide residues on Indian tiger beetles and other

predatory insects.

Promoting Scientific Contributions Our greatest hope in publishing this field guide has been to generate increased

public interest and appreciation about tiger beetles in India. By promoting public awareness of the important role tiger beetles have in the biological world, we are

confident that interest can be transformed into proper maintenance of their natural habitats. However, this increased interest may produce unintended and deleterious consequences under some circumstances in the form of increased collecting, trading and selling of tiger beetles, especially the rarer ones. While there is little evidence to

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Conservation and Economic Impact

date that over collecting has seriously impacted any tiger beetle species, increased collecting of rare and localized species may put these forms at greater risk. For that reason we urge tiger beetle enthusiasts to limit numbers of specimens they collect. Workers can also help in the conservation of these and many other insect species by recording new distribution data, monitoring sites to assess any decline in species diversity and population numbers therein, and observing and recording ecological

and habitat conditions that may limit survival of adults and larvae. All this biological

information should be shared with appropriate workers, NGO's, land managers, or government agencies.

Biogeography

9

Biogeography is the study of the geographic distribution of plants and animals.

It involves patterns of habitat use in the short term and geological time affecting distribution. Tiger beetle species are not evenly distributed across India, and therefore, the range maps for no two species in India are identical. Each species

has a distinctive range whether measured in total area, shape of the boundaries, or

latitude and altitude. As is typical of most plants and animals, more species of tiger beetles occur in some parts of India than others (Fig. 9.1). Why is this?

Figure 9.1 Isoclines connecting squares in a grid with the same number of tiger beetle species (each square 3° latitude and 3° longitude) on the Indian Subcontinent (Reprinted from Pearson and Cassola 1992, Conservation Biology volume 6: 376-391. Used by permission from the publisher, Wiley-Blackwell).

One obvious explanation is that where the environment is less extreme and

resources like food are reliable, more tiger beetle species can survive. Annual rainfall amounts have been shown to affect plant productivity in a habitat which in turn influences the richness of the animal species living there. This correlation is evident when comparing the rainfall amounts and tiger beetle species richness across the Indian Subcontinent (Fig. 9.2). The harsh dry deserts of northwest India support few species compared with the more benign high rainfall areas in northeast and southwest India where the highest numbers of species are found.

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

180

Another explanation involves historical movements (dispersal) of tiger beetle

species within the Indian subcontinent, together with movements from other areas

(migration) onto the subcontinent. Geologically, the Indian Subcontinent is made up of a single tectonic plate. During the Mesozoic in the early Cretaceous, about 140 million years ago, this Indian Plate was connected to Africa, Australia, Antarctica, and South America as the supercontinent Gondwana. In the late Cretaceous, roughly

Figure 9.3

Movement of the Greater Indian Plate from the rest of Gondwanaland towards the Eurasian land mass during the Palaeocene 35 - 55 million years B.P. (Reprinted from Pearson and Ghorpade 1989, Journal of Biogeography 16:333344. Used by permission from the publisher, Wiley-Blackwell).

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Biogeography

100 million years ago, the Indian Plate split from Africa and began moving north (Fig. 9.3) across the ancient Tethys Sea. In the middle Tertiary of the Cenozic, about

35 to 55 million years ago, the Indian Plate collided along the southern coast of Asia (Eurasian Plate). With a combination of folding and pushing below the Eurasian Plate (subduction), this collision also began to push up the Himalayas and other mountain ranges (Fig. 9.4).

' v’tG hl;A

Nd S

/'

99 k9.4 Present-day mountain chains and highlands of the Indian subcontinent (l=Garo Rajmahal Gap; 2=Palghat Gap) (from Pearson and Ghorpade 1989, Journal of Biogeography 16:333-344. (Reprinted from Pearson and Ghorpade 1989, Journal of Biogeography 16:333-344. Used by permission from the publisher, Wiley-Blackwell). Figure

Molecular and morphological evidence point to the migration of the ancestors of our present day Indian tiger beetle species onto the Indian Plate once

it contacted the Laurasian land mass (Fig. 9.5). Here they establish themselves as parts of the Indian fauna, and in many cases they evolved into new species

(speciation). Following their arrival onto the Indian Plate, tiger beetle species dispersed by various routes. Fairly direct routes became available for coastal species and those

occupying other continuous habitats, such as, river banks were not likely to have been unimpeded when dispersing throughout the Indian Plate. But movements of those species occupying discontinuous habitats were also inevitable. A possible explanation for such discontinuous dispersal involved strong storms transporting flying adults from one suitable habitat to another some distance away.

287

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Figure 9.5

Theorized dispersal routes used by tiger beetles species moving onto and within the Indian Subcontinent. Purported ancestors dispersed from Africa (route 1), southwestern Asia (route 2), northern Asia (route 3), and southeastern Asia (route 4); within the subcontinent they moved via forested mountain chains (routes 5a and 5b), land bridges between Sri Lanka and the southern tip of India (route 6), and sandy ocean beaches (routes 7 and 8) (Reprinted from Pearson and Ghorpade 1989, Journal of Biogeography 16:333-344. Used by permission from the publisher, Wiley-Blackwell).

When tiger beetle species poured over the Himalayas and associated mountain ranges from southeastern, southwestern, and northern Asia, they likely encounter a

climate across India that was wetter than it is today. The forests of the low internal mountain ranges, the Western and Eastern Ghats, at that time were connected and apparently could serve as a continuous belt of habitat along which forest-adapted tiger beetle species could disperse freely. As the climate dried out, the forest was disrupted, and many of these tiger beetles became isolated in higher elevation remnants of forest along the earlier belt (Fig. 9.5). Contrary to these explanations about migration for tiger beetles into India,

the origin of one group of species, the Tethys Tiger Beetles (genus Jansenia), is

especially perplexing. It may have involved their ancestors drifting on the ancient Indian tectonic plate as it moved across the Tethys Sea from Gondwana to crash into Eurasia millions of years ago. This geological explanation for a restricted historical

presence would help explain why the Tethys Tiger Beetles have their centre of species richness in South India with only a few species occurring elsewhere in India. Why so few species of this genus occur in northern India may be best explained by the influx of many superior competitors from the Eurasian Plate. Of the few species of Jansenia known from northern India, all of them are restricted to extreme habitats like boulder and cliff faces. Apparently, these habitats were so unsuitable to invading Asian species that they avoided them. Consequently, as such habitats 288

Biogeography

became available, the northern Tethys Tiger Beetles utilized them in order to escape

the dominant Eurasian competitors'adverse impacts.

Spatial patterns of diversity are often based on observations of animals and plants in relation to latitudinal changes across large land masses; more and more species typically are encountered towards the equator. The patterns of tiger beetle species diversity on the Indian subcontinent, however, are considerably more complicated (Fig. 9.1), such that the concept of latitudinal gradient in diversity

explains little of this pattern. A better explanation of this complex spatial pattern of diversity includes both habitat severity and biogeographical dispersal routes. Higher rainfall areas with their accompanying more benign habitats may have allowed greater access to the Indian

Subcontinent from the Eurasian mainland. These favourable continental contact areas, especially in northeastern India, funneled dispersal routes from several parts

of Asia onto the Indian subcontinent. Here immigrants from various parts of Eurasia overlapped to produce a crossroads effect, and many species from numerous origins accumulated randomly.

Local and regional studies of tiger beetle species within India have concluded that development of endemism provides another factor contributing to the overall

high diversity of tiger beetles. Of the 241 tiger beetle species recorded for India, 119

(49%) are endemic to the country. Sixty-six (56%) of these endemic species belong to two genera, Neocollyris and Jansenia. Understanding exactly what historical and ecological factors stimulated the evolution of so many new species on the Indian

subcontinent will be revealed only with more data on distribution, natural history

and molecular evidence. Tiger beetles species and their evolutionary branches follow a pattern of

distribution common to many plants and animals in India. Almost all of them are restricted to one of six distinct biogeographic zones: 1) Himalayas, 2) Western/ Eastern Ghats, 3) Coastal, 4) Gangetic Plains, 5) Northeastern Tropics, and 6) Andaman/Nicobar Islands. A final complicating factor is that the vast majority of observations and

collections of tiger beetles have been from the northeastern, southern, and northern parts of India. Other regions, such as, central and northwestern India with only a few

field studies, are underrepresented in this Field Guide. So, are there fewer species in these regions simply because few people have looked there or do fewer species

really exist there? Work in those regions of India are likely to provide excellent opportunities for amateurs to make significant contributions to tiger beetle studies

with the likelihood of revealing new biological information and perhaps discovering new species.

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Observing and Studying Tiger Beetles

10

Permits and regulations If you are planning to conduct a research program on Indian tiger beetles, and if you will be collecting specimens, understand that government permits are required. Studies and collections done without prior permission, especially in protected areas, can result in heavy fines.

As per the Wildlife (Protection) Act 1972, and Biological Diversity Act 2002, collection of any biological material is regulated. Scientific study on flora and fauna

species can be permitted with prior permission from Chief Wildlife Warden of the concerned states (for National Parks and Sanctuaries) and State Biodiversity Boards for sites outside protected areas. Biological material can only be sent outside of the country for scientific research and identification with prior permission from the National Biodiversity Authority and State Biodiversity Boards. No commercial sale of any specimens is permitted. If you are a professional photographer intending to sell commercial photos of tiger beetles from protected areas, a permit is required from the Chief Wildlife

Warden of respective states. Fee and rules has been defined by the Government. Currently amateur photography and observations do not require a permit.

Field observations If you can learn to become motionless or make very slow, smooth movements, it is amazing how close tiger beetles will allow you to observe their normal behaviour. Watch out for your shadow though, as its movement can easily frighten the wary

beetle. Close-focusing binoculars can help in observing and recording detailed behaviour. Hand nets can be used to capture, mark, and release individuals for further observation. Larval tiger beetles quickly become used to nearby observers, even to the point of accepting prey items offered them with forceps.

If you want to measure the available prey in an area, sticky traps and pitfall traps are the most useful. Nightlights, ultraviolet or full spectrum, attract many but not all species of adult tiger beetles, even diurnally active ones. Some species, especially

arboreal forms in dense undergrowth, are best found by beating the foliage with a stick into a portable sheet or inverted umbrella. Flight interruption traps, such as Malaise traps, are effective in confined habitats along streambeds, paths through forests, or at the base of cliffs. Learn to identify the round larval holes in the soil surface that then open down into their vertical tunnels. You can collect individual larvae by inserting a grass stem

A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

or other thin, flexible object down the length of the larval tunnel. Then dig carefully

down the length of the inserted object until the larva is exposed. Alternatively, larvae will sometimes grasp the intruding grass stem inserted into the tunnel, and they can be carefully extracted as they cling to the object with their mandibles.

Captive studies The adults and larvae of most tiger beetle species will adjust to life in a covered terrarium with native soil from the site where they were collected, appropriate light and moisture. Adult feeding, mating behaviour, and oviposition can be easily observed, photographed and studied. Larvae can be raised to test such factors as moisture, temperature, and food level influence on larval development, survival, and their fecundity as adults. Some species are more aggressive than others toward each

other, however, and several adult individuals in the same container can be deadly.

Molecular studies Molecular analysis and DNA studies are quickly becoming the preferred mode of study, and many museums and institutions are now developing frozen tissue collections. Alternatively, long-term storage (10-20 years) in alcohol at -20° C maintains DNA quality adequately for many analyses. The cold temperature slows down the DNA degradation. However, many chemicals can degrade DNA, so specimens captured in killing jars and those cleaned in solvents often are not useful for these molecular studies.

DNA and other molecules can be collected from long-term dried specimens that have not been treated with chemicals. Often a single leg part is enough for molecular analysis and does not destroy the rest of the specimen.

Photography Just as for birds, butterflies, and dragonflies, the collection of tiger beetle specimens is becoming less common. Most collecting is now reserved for specific types of

studies in which tissues, cells, and structures can only be observed and studied using specimens. For studies of behaviour, ecology, and range extensions (and contractions), or for the pure pleasure of recording the beauty and intricacies of tiger beetle form, colour, and pattern, photographs are more and more the preferred method.

If you wish to send a photographic record of a tiger beetle to an expert for identification, make sure the image is as sharp as possible.Take multiple photos from

above, from the side (including the legs), and the front of the head. Additionally, it is important to add information about the body length and the location (region, city, geographical coordinates). Body length can be determined with a reference

measurement in the actual image, such a scale on a ruler, or the known length of an 292

Observing and Studying Tiger Beetles

object like a blade of grass or width of a small stone. Almost all Indian tiger beetles

can be identified to species or even subspecies with this information. However, some species of Brush Tiger Beetles (Neocollyris) and many of the small species of

Rounded Thorax Tiger Beetles (Cylindera) are notoriously difficult to distinguish even with excellent photos. Collecting a specimen of these species may be the only way to study them so microscopic species characters can be compared. Becoming a successful tiger beetle photographer means developing skills not only in the techniques of camera use, but also in nature study. Patience, and more patience, is often the difference between an adequate shot and a spectacular

photograph. Learn all you can about your subject and its environment before you

try taking pictures. How prone to fly is the species? If it does fly, will its flight path

take it to a predictable part of the environment? How can you best stalk your subject, bearing in mind that this will include arriving at the best spot that will allow you to take advantage of the shallow depth of field needed for close-up photography? In extreme cases, non-purists will even capture the insect, cool it down in a refrigerator or with ice from a field pack for 15 minutes, and then release it into a controlled photographic situation, where the beetle's slowed activity allows more critical focusing and sharper images.

Type of camera and lens, lens aperture, ISO rating, shutter speed, the use of one or more flashes, recording format (typically RAW vs JPEG), and many other

alternatives comprise only part of the array of decisions you will need to make. Whatever your decisions, there will be one constant element in tiger beetle photography: it will eventually involve a lot of crawling and lying on the ground. You

will soon be selecting equipment that adapts well to this kind of behaviour. Whatever medium you choose to capture your tiger beetle images, your knowledge and appreciation of these marvelous creatures will be immensely

enhanced as a result. Post your photos on line for others to see and admire. One of these exciting sites, Insectlndia, is on Facebook (https://www.facebook.com/groups/ lnsectlndia/?ref=group_header). Here hundreds of enthusiastic Indian amateur entomologists and photographers share their photos for fun and for science. We used many of these photos in this field guide, and their data were useful in updating

regional distributions of several Indian tiger beetles. Also for examples of spectacular beetle photography, look at the web site"Carabidae of the World", www.carabidae.org, which has more than 2000 photos of tiger beetles from around the world.

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Fowler, W. W. (1912) Fauna of British India including Ceylon and Burma (Coleóptera General Introduction and Cicindelidae and Paussidae). Today and Tomorrow's Printers and Publishers, New Delhi, 529 pp.

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A Field Guide to the Tiger Beetles of India: Identification and Biology of the Cicindelidae

Horn, W. (1905) Systematischer Index der Cicindeliden. Deutsche Entomologische Zeitschrift 2:1-56. Horn, W. (1915) Coleóptera Adephaga (Family Carabidae, Subfamily Cicindelinae). In: R Wytsman, (Ed.), Genera Insectorum (Fascicles 82 A-C), Brussels.

López-López, A., Aziz, A. A. and Galián, J. (2015) Molecular phylogeny and divergence time estimation of Cosmodela (Coleóptera: Carabidae: Cicindelinae) tiger beetle species from Southeast Asia. Zoológica Scripta doi:10.1111/zsc.12113. Matalin, A.V. and A. V. Anichtchenko (2012) New records of tiger beetles (Coleóptera, Cicindelidae) from India and Sri Lanka with description of a new subspecies. Eurasian Entomological Journal 11:151-156.

Mawdsley, J. R. (2010) Cladistic analysis of Cicindela Linnaeus 1758, subgenus Pancallia Rivalier 1961, a lineage of tiger beetles from southern India (Coleóptera Cicindelidae). Tropical Zoology 23:195-203.

Mittal, O. R, Gill, T.K. and Chugh, S. (1989) Chromosome studies on three species of Indian cicindelids (Adephaga: Coleóptera) Caryologia 42:115-120. Naviaux, R. (1994) Les Collyris (Coleóptera, Cicindelidae): révison de genres et description de nouveaux taxons. Société linnéenne de Lyon 63:106-290. Naviaux, R. (2002) Les Tricondylina (Coleóptera, Cicindelidae): révison de genres Tricondyla Latreille et Derocrania Chaudoir et descriptions de nouveaux taxons. Annales de la Société entomologique de France 5:1-106. Pajni H. R. and Bedi, S. S. (1973) Preliminary survey of the cicindelid fauna of Chandigarh, Punjab, India. Cicindela 5:41 -54.

Pearson, D. L. (1994) Selecting indicator taxa for the quantitative assessment of biodiversity. Philosophical Transactions of the Royal Society B: Biological Sciences 345: 75-79. Pearson, D. L. (2019) Tiger beetles (Coleóptera: Cicindelidae): Their history and future in Indian biological studies. In: Belavadi, V. V., Mallik, B., Ramani, S. and Yeshwanth, H. M. (Eds.), Festschrift for C. S.Viraktamath, Diversity of Insects in India. Taylor and Francis, UK, pp. 201-222.

Pearson, D. L. and Carroll, S. S. (1998) Global patterns of species richness: Spatial models for conservation planning using bioindicator and precipitation data. Conservation Biology 12:809-821. Pearson, D. L. and Ghorpade, K. D. (1989) Geographical distribution and ecological history of tiger beetles (Coleóptera: Cicindelidae) of the Indian subcontinent. Journal of Biogeography 16:333-344. Pearson, D. L., Knisley, C. B., Duran, D. P. and Kazilek, C. J.. (2015) A Field Guide to the Tiger Beetles of the United States and Canada: Identification, Natural History, and Distribution of the Cicindelinae, 2nd edition. Oxford University Press, New York, 251 pp. Pearson, D. L. and Vogler, A. P. (2001) Tiger Beetles: the Evolution, Ecology, and Diversity of the Cicindelids. Cornell University Press, Ithaca, 333 pp. 296

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Rivalier, E. (1971) Remarques sur la tribu de Cidndelini (Col. Cicindelidae) et sa subdivision en soustribus. Nouvelle Revue d'Entomologie 1:135-143. Roza,A.S.andMermudes,J.R.M.(2017)Tigerbeetles'(Coleoptera:Carabidae,Cicindelinae) pupal stage: current state of knowledge and future perspectives. Zootaxa 4266: 348 -358.

Sawada, H. and Wiesner, J. (2006) Records of tiger beetles collected in North India II (Coleóptera: Cicindelidae). 97. Contribution towards the knowledge of Cicindelidae. Entomologische Zeitschrift 116:127-134. Shivashankar, T. and Pearson, D. L. 1994. A Comparison of mate guarding among five syntopic tiger beetle species from peninsular India (Coleóptera: Cicindelidae). Biotropica 26:436-442. Shivashankar, T. and Veeresh, G. K. (1987) Impact of differential feeding on the reproduction of tiger beetle Cicindela cancellata DeJean (Cicindelidae: Coleóptera). Proceedings Indian Academy of Sciences (Animal Science) 96:317-321. Shivashankar, T., Kumar, A. R. V., Veeresh, G. K. and Pearson, D. L. (1988) Angular turret building behaviour in a larval tiger beetle from South India (Coleóptera: Cicindelidae) Coleopterists Bulletin 42:63-68. Sinu, P. A., Nasser, M. and Rajan, P. D. (2006) Feeding fauna and foraging habits of tiger beetles found in agro-ecosystems in Western Ghats, India. Biotropica 38:500-507. Uniyal, V. P. and Bhargav, V. (2007) Tiger beetles: A field study in the Shivaliks of Himachal Pradesh. Wildlife Institute of India, Dehradun. http://oldwww.wii.gov.in/ faculty/vpuniyal/tiger_beetle. Uniyal, V. P. and Mathur, P. K. (2000) Altitudinal distribution of tiger beetles (Cicindelidae: Coleóptera) in Great Himalayan National Park Conservation Area, Western Himalaya. Indian Forester 126:1141-1143.

Vogler, A. P. and Pearson, D. L. (1996) A molecular phylogeny of the tiger beetles (Cicindelidae): Congruence of mitochondrial and nuclear rDNA data sets. Molecular Phylogenetics and Evolution 6:321-338.

Wiesner, J. (1988) Die Gattung Therates Latr. und Ihre Arten (15. Beitrag zur Kenntnis der Cicindelidae, Coleóptera). Mitteilungen der Münchner Entomologischen Gesellschaft 78:5-107. Wiesner, J. (1992) Checklist of the Tiger Beetles of the World (Coleóptera: Cicindelidae). Verlag Erna Bauer, Keltern, Germany, 364 pp. Yadav, J. S., Burra, M. R. and Dange, M. P. (1989) Chromosome number and sex­ determining mechanism in 32 species of Indian Coleóptera (Insecta). National Academy of Sciences and Letters 12:93-97.

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