Potter Wasps (Hymenoptera: Vespidae: Eumeninae) of India

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Indian Insects

Indian Insects
Diversity and Science
A Festschri for Professor C. A. Viraktamath’s 75th Birthday
Edited by
S. Ramani
Prashanth Mohanraj
H. M. Yeshwanth

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“My work enraptures but utterly exhausts me….To know that no one before you has seen an
organ you are examining, to trace relationships that have occurred to no one before, to immerse
yourself in the wondrous crystalline world of the microscope, where silence reigns, circumscribed
by its own horizon, a blindingly white arena—all this is so enticing that I cannot describe it.”
Letter to his sister Elena
Vladimir Nabakov (1945)
“Systematists will have only to decide (not that this will be easy) whether any form be
sufciently constant and distinct from other forms, to be capable of denition, and if
denable whether the difference be sufciently important to deserve a specic name.”
Origin of Species
Charles Darwin (1858)
Alice explained, “…I can tell you the names of some of them.”
“Of course they answer to their names?” the Gnat remarked carelessly.
“I never knew them do it.”
“What’s the use of their having names” the Gnat said, “if they won’t answer to them?”
“No use to them,” said Alice; “but it’s useful to the people who namethem,
I suppose. If not, why do things have names at all?”
Through the Looking Glass
Lewis Carroll (1871)

vii
Contents
Preface..........................................................................................................................................................................................ix
Acknowledgments ........................................................................................................................................................................ xi
Editors ....................................................................................................................................................................................... xiii
Chandrashekaraswami Adiveyya Viraktamath: ThePrince of Indian Taxonomists ................................................................. xv
Contributors ............................................................................................................................................................................... xxi
Chapter 1 Why Are Insects Abundant? Chance or Design? .................................................................................................... 1
K. N. Ganeshaiah
Chapter 2 Mayies (Insecta: Ephemeroptera) of India ............................................................................................................ 7
C. Selvakumar, K. A. Subramanian, and K. G. Sivaramakrishnan
Chapter 3 Dragonies and Damselies (Insecta: Odonata) of India ..................................................................................... 29
K. A. Subramanian and R. Babu
Chapter 4 Stoneies (Insecta: Plecoptera) of India ................................................................................................................ 47
R. Babu, K. G. Sivaramakrishnan, and K. A. Subramanian
Chapter 5 Taxonomy of Orthoptera with Emphasis on Acrididae ......................................................................................... 57
Rajamani Swaminathan and Tatiana Swaminathan
Chapter 6 Taxonomy and Biodiversity of Soft Scales (Hemiptera: Coccidae) ...................................................................... 69
Sunil Joshi
Chapter 7 Whiteies (Hemiptera: Aleyrodidae) of India .................................................................................................... 103
R. Sundararaj, K. Selvaraj, D. Vimala, and T. Venkatesan
Chapter 8 Pentatomidae (Hemiptera: Heteroptera: Pentatomoidea) of India ...................................................................... 121
S. Salini
Chapter 9 Indian Mymaridae (Hymenoptera:Chalcidoidea) .............................................................................................. 147
S. Manickavasagam and S. Palanivel
Chapter 10 Scelionidae and Platygastridae (Hymenoptera: Platygastroidea) of India .......................................................... 159
K. Rajmohana and Sunita Patra
Chapter 11 Bumble Bees (Hymenoptera: Apidae: Apinae: Bombini) of India ..................................................................... 173
Martin Streinzer
Chapter 12 Potter Wasps (Hymenoptera: Vespidae: Eumeninae) of India ............................................................................ 187
P. Girish Kumar, Arati Pannure, and James M. Carpenter

viii Contents
Chapter 13 Tiger Beetles (Coleoptera: Cicindelidae): Their History and Future in Indian Biological Studies .................... 201
David L. Pearson
Chapter 14 Coccinellidae of the Indian Subcontinent ........................................................................................................... 223
J. Poorani
Chapter 15 Flea Beetles of South India (Coleoptera: Chrysomelidae: Galerucinae: Alticini) .............................................. 247
K. D. Prathapan
Chapter 16 Taxonomy, Systematics, and Biology of Indian Butteries in the 21st Century ................................................. 275
Krushnamegh Kunte, Dipendra Nath Basu, and G. S. Girish Kumar
Chapter 17 Taxonomy and Diversity of Indian Fruit Flies (Diptera: Tephritidae) ................................................................ 305
K. J. David, S. Ramani, and S. K. Singh
Chapter 18 Hover-Flies (Diptera: Syrphidae) Recorded from “Dravidia,” or Central and Peninsular India
andSriLanka: An Annotated Checklist and Bibliography ................................................................................ 325
Kumar Ghorpadé
Chapter 19 AComparative Study of Antennal Mechanosensors in Insects .......................................................................... 389
Harshada H. Sant and Sanjay P. Sane
Chapter 20 Cross-Kingdom Interactions in Natural Microcosms: TheWorlds Within Fig Syconia
and Ant-PlantDomatia ........................................................................................................................................ 401
Renee M. Borges, Joyshree Chanam, Mahua Ghara, Anusha Krishnan, Yuvaraj Ranganathan,
MeghaShenoy, Vignesh Venkateswaran, and Pratibha Yadav
Annexure I: Revisions and Reviews of Taxa by Professor C. A. Viraktamath ....................................................................... 41 5
Annexure II: New Tribe and Genera Described by Professor C. A. Viraktamath ................................................................. 417
Annexure III: New Species Described by Professor C. A. Viraktamath ............................................................................... 41 9
Annexure IV: Research Publications of Professor C. A. Viraktamath ................................................................................... 427
Annexure V: Taxa Named in Honour of Professor C. A. Viraktamath .................................................................................. 437
Annexure VI: Courses Offered by Professor C. A. Viraktamath ........................................................................................... 439
Annexure VII: Theses Submitted Under the Guidance of Professor C. A. Viraktamath (1980–2019) .................................. 441
Annexure VIII: Research Projects Operated by Professor C. A. Viraktamath ...................................................................... 443
Index ......................................................................................................................................................................................... 445

ix
Preface
Inthe Agricultural Colleges the aim is to train farmers and
fruit growers and notentomologists….. …they should have
just a general knowledge of the external anatomy of insects
so as to be able to place the insects at least in their orders.
Thisamount of systematic work is quite sufcient for them.
C. C. Ghosh, Asst. to the Imperial Entomologist,
Pusa(1919)
…During this meeting we have been constantly reminded of
the very great difculty of getting our specimens identied.
… We do want … an institute such as the British Museum
(Natural History) … where specialists can work and our
insects be identied.…. Without pure science we cannot go
very far with our applied science.
M. Afzal Husain, Supernumerary Entomologist,
Pusa(1919)
The last subject taken up for discussion at the Third
Entomological Meeting at Pusa in February, 1919 was
“The Organization of Entomological Work in India.” When
initiating the discussion on this subject, T. B. Fletcher, the
then Imperial Entomologist to the Government of India, told
the gathering of entomologists from across the country that
after considerable thought, he felt that “entomological work,
particularly entomological research had to be improved in the
country.” The antithetical views appearing as the epigraph
above formed part of this discussion. Since then, Indian ento-
mology has been perpetually plagued by this Janus-faced
approach to insect taxonomy and systematics, the reverbera-
tions of which continue to haunt us to this day.
Linnaean taxonomy was introduced during Linnaeus’s
time to India by J. G. Koenig, a private student of his, who
came to India in 1767. Hewas followed by I. K. Daldorff, a
pupil of Fabricius, who came to India in 1790. Thisearly start,
however, failed to lend the necessary llip to the development
of indigenous expertise in insect taxonomy on Linnaean prin-
ciples in India. Unfortunately, this situation did notimprove
either during the period of British rule or even after the coun-
try gained political independence in the mid twentieth century.
Ironically, while a number of British entomologists worked
on the taxonomy of Indian insects, only a handful of Indian
entomologists developed the necessary expertise to work
independently on taxonomy. Thisexpertise was not, however,
passed on and did notfoster succeeding generations of home
grown entomologists. Insect taxonomy remained “provincial”
and dependent on metropolitan science. TheBritish Museum
(Natural History), London, was the main institution on which
Indian entomologists largely depended for the determination
of the taxonomic identity of insects. This situation was to
change dramatically in the late twentieth century as the BM
(NH), nowthe Natural History Museum, at London, changing
policy, began to charge (exorbitantly for a developing econ-
omy) for their identication services. If the country was notto
lose scarce foreign exchange reserves for the identication of
large numbers of insect specimens, the necessary capabilities
had to be built locally.
With the singular motive of serving this end, an ardent
entomologist, Professor C. A. Viraktamath, with great fore-
sight, had already set in motion a new approach to taxonomic
pedagogy. Theconcept of a “practical record” (and all that
it entailed) in taxonomy courses—till then the staple in any
practical class—was dispensed with. Heintroduced students
to taxonomic keys and trained them in the use of these keys to
determine the identity of specimens. More students were allot-
ted topics on taxonomy for their theses as part of the require-
ment for the completion of their post graduate programmes.
As students thus became more independent and competent in
the principles and practices of insect taxonomy, they contin-
ued to pursue this study in their places of employment. Thus
developed centres of taxonomic excellence on selected insect
taxa in various parts of the country.
Inthis festschrift, we have drawn together entomological
expertise from those who have been associated with Professor
Viraktamath and have drawn sustenance from his entomolog-
ical knowledge. Insect taxonomy has of course been the focus
of our attention as there is still no single book on the taxon-
omy of Indian insects that serious students of entomology can
turn to. As Eliot Zimmermann, the American entomologist,
put it “those who know most about this subject, know best
how little they know.” Acknowledging this sentiment to the
hilt, we venture into the shallows knowing full well that one
day the deep too will be conquered.
Thisbook makes no pretensions to being a textbook on
Indian insect taxonomy. It, however, takes the decisive pre-
liminary step of being the nucleus of such a book. We have
notbeen able to draw on the expertise of all the competent
taxonomists for various reasons. Of the 20 chapters com-
prising this book, 17 deal with the taxonomy of different
insect taxa. While four groups of insects (Ephemeroptera,
Odonata, Plecoptera, and Orthoptera) have been dealt with
at the order level, ve others (Hemiptera, Hymenoptera,
Coleoptera, Lepidoptera, and Diptera), which include the
mega orders, have been treated in parts at various ranks
below the order. A major endeavour has been to pro-
vide identication keys with appropriate illustrations in
many cases, with the focus squarely on the Indian fauna.
Depending on the nature of expertise of our contributors,
these keys serve to identify Indian insects at different levels
in the taxonomic hierarchy. Thechapters focus on the pres-
ent status and could form a starting point for students and
young researchers to leverage further work on these groups.
Fully aware of the incompleteness of a “book of even this
limited scope,” we hope that one that includes all Indian
taxa will soon be forthcoming.
The three remaining chapters explore different facets of
insect life by investigators who have been closely associated

xPreface
with Professor Viraktamath. Therst is a novel explanation
for why insects are so abundant, the other, an anatomical
study comparing mechanosensors in the antennae of insects,
and the third explores the exciting world of natural micro-
cosms inhabited by insects.
Idiosyncrasies in presentation are a necessary concomi-
tant of a compendium of this kind. While the authors were
assigned taxa or topics of their expertise, they were given the
leeway to present and organize the content of their chapters in
any manner they chose to with the delity of the content being
their sole responsibility.
Vive la Professor Viraktamath and may his illustrious
legacy ll in the long standing requirement for a book on the
taxonomy of all orders of Indian insects.
Editors

xi
Acknowledgments
First and foremost, our greatest debt is to Professor Viraktamath,
who instilled the love for entomology in each of us and taught
us to observe and bring order into the bewildering diversity
of insect life teeming around us. Healso permitted us to peer
unabashedly into and query his past, one that had undoubtedly
dimmed with the steady, relentless passage of time.
Itwas with trepidation that the six of us—Ramani, Yeshwanth,
Belavadi, Chandrashekar, Mallik, and Prashanth—embarked on
this venture of lling a lacuna in the orbit of Indianentomologi-
cal pedagogy. That there was a genuine requirement was never
in doubt. But our concern of whether we could we pull it off,
constrained by the bounds of a festschrift, was genuine too.
Fortunately, the circle of those who were Professor Viraktamath’s
students or had been associated with him was so wide that we
had little difculty in getting together a band of the majority of
the most respected taxonomists as well as those whose research
depended on his taxonomic expertise in the country.
We thank all the contributors to this volume for having
accepted our invitation to write with authority on taxa of their
interest and take responsibility for the contents of their respec-
tive chapters. We also thank others who submitted chapters
which could notbe included in this book for a variety of reasons.
H. V. Ghate, A. K. Dubey, A. Ramesh Kumar, V. V. Belavadi,
S. S. Anooj, and Professor C. A. Viraktamath graciously gave
of their time to review the manuscripts given to them. We are
grateful to all of them for this kind gesture.
Ms. Renu Upadhyay, assistant commissioning editor,
Ms. Shikha Garg, and Ms. Jyotsna Jangra editorial assis-
tants of the CRC Press, Taylor & Francis Group, eased
the pressures of working on the manuscript of such an
undertaking with great forbearance and understand-
ing. Fully appreciative of the rationale of our request to
increase the size of the book, they enabled it with alac-
rity. We remain indebted to them for answering our queries
without rancour and for their near stoic patience in waiting
for us to submit the completed manuscript even after it had
crossed the deadline.
We wish to acknowledge Ms. Rachael Panthier, Production
Editor, Taylor & Francis as well as the able team at Lumina
Datamatics, Puducherry led by Mr. Sundaramoorthy
Balasubramani for bringing in their expertise to ensure high
production standards.
Editors

xiii
Editors
Dr. S. Ramani joined the
Agricultural Research
Service of the ICAR, New
Delhi, India, in 1985 and
after serving in different
ICAR institutes retired as
Project Coordinator, ICAR-
AICRP on Honey Bees &
Pollinators in 2011. His
areas of specialization are
pollination, biological con-
trol, and taxonomy. He has
published more than 45 papers in peer-reviewed journals and
several popular articles on insects. His special area of interest
is taxonomy of Tephritidae. Hecollects stamps on insects as a
hobby.
Dr. Prashanth Mohanraj
retired as a principal scientist
from the ICAR-National
Bureau of Agricultural
Insect Resources, Bengaluru,
India, in 2017. He joined
the Agricultural Research
Service of ICAR and worked
at the ICAR-Central Islandm
Agricultural Research Insti-
tute, Port Blair, Andaman&
Nicobar Islands, for 14years.
He co-authored a book Butteries of Andaman & Nicobar
Islands, the Hindi translation of which won the best book
award from the ICAR. His areas of specialization are biologi-
cal control, natural history of insects, and taxonomy. Hehas
specialized on the taxonomy of Trichogrammatidae. Hehas
published over 65 research papers in peer-reviewed journals
and more than a dozen popular articles on insects.
Dr. H. M. Yeshwanth is an
assistant professor in the
Department of Entomology
at University of Agricultura l
Sciences, Bengaluru, India.
His research interests are in
the area of taxonomy, bio-
diversity, and natural his-
tory of insects. His special
area of interest is taxonomy
of Hemiptera, especially
Miridae, and he has
described about 25 new species of mirid bugs. Hehas pub-
lished over a dozen papers in highly rated international
journals. Hehas developed excellent techniques for photo-
graphing insects both in nature and from museum
specimens.

xv
Chandrashekaraswami Adiveyya Viraktamath
ThePrince of Indian Taxonomists
CAV, as professor C. A. Viraktamath is known to those in the
country familiar with his work or have sought his advice or
assistance on matters of insect taxonomy, are three letters that
are as recognizable as if he were a brand like BMW, BEL, or
HAL. Vakma, Mutt, and Chandra (with Viraktamath becom-
ing Vakma or Mutt and Chandrashekaraswami—Chandra)
are other sobriquets that he is known by to those who nd
his 38 letters long name—Chandrashekaraswami Adiveyya
Viraktamath—containing over half the letters of the alpha-
bet, more than a mouthful.
Seventy-ve years ago, on January 31, 1944, when their
second child, a boy was born to Adiveyya and Neelambika
in Byadagi—in the heart of present-day Karnataka, famous
for its deep red, spicy chillies—little did they imagine that he
would turn out to be one of the most respected taxonomists
notonly in the country, but also internationally.
His father who worked in the Revenue Department was
constantly transferred as part of government policy. Thisperi-
patetic existence saw CAV studying in different schools in the
different places at which his father was posted. After school,
he joined the College of Agriculture, Dharwad to pursue a
BSc degree in Agriculture. As at school, he continued to be
a brilliant student at college too and won the ASPEE Gold
Medal for the highest marks in Plant Pathology. He then
gained admission at the University of Agricultural Sciences,
Bangalore for the MSc (Agricultural Entomology) program.
Here, he did notfail to impress Dr. H. M. Harris (a taxono-
mist specializing on Nabidae, Hemiptera) and Dr. J. H. Lilly,
visiting professors under the Ford Foundation program, who
taught him insect systematics and insect physiology, respec-
tively. It was at this time that the ragi (Eleusine coracana)
crop was severely affected by the mosaic and streak dis-
eases. Leafhoppers were known to transmit these diseases,
but no one in India could identify the leafhoppers that were
their vectors. The expertise of taxonomists in the USAand
the UK had to be sought. Atthis juncture, Harris convinced
“Vakma” (for this was how he abbreviated the, to him, unpro-
nounceable name—Viraktamath) to take up the taxonomic
study of leafhoppers. Recommended by Harris and Lilly,
CAV gained admission to Oregon State University (OSU) as a
Ford Foundation scholar to study leafhopper taxonomy under
Dr.Paul W. Oman (PWO), the then most eminent leafhopper
taxonomist in the world.
H. M. Harris, Visiting Professor, Ford Foundation at University
of Agricultural Sciences, Bangalore
AtOSU, he was a man in a hurry. Hehad to pick up every-
thing from his mentor in a mere year’s time. To do this, he
reduced his course load (he’d complete the remaining courses
on his return to UAS, Bangalore). Here, he had to devote
all the available time to examining the large number of Old
World agalliine leafhoppers (the group assigned to him for his
PhD thesis) in Oman’s laboratory as well as all the specimens
arriving from museums and from PWO’s fellow entomolo-
gists from all over the world. His day began early and went on
into the wee hours of the night as he peered endlessly into the
microscope to dissect and draw the intricate structures that
served to distinguish one species of leafhopper from another.
Most nights he’d be the last to leave the laboratory. Working at
this frenetic, febrile pace, he notonly studied the specimens,
especially the types borrowed from different museums, but
also found the time to travel to the California Academy of
Sciences and sort their unsorted leafhopper material from the
Old World so that he could borrow the agalliine leafhoppers
from the collection for his research.
While G. P. Channabasavanna, the Head of the Department
of Entomology, UAS, Bangalore, had taught him the ne art
of camera lucida drawing, and Harris had introduced him to
the rudiments of leafhopper taxonomy, he still had to learn
the ner details under Oman, who patiently taught him the
intricacies of the genital architecture of male leafhoppers for

xvi Chandrashekaraswami Adiveyya Viraktamath
the rst time and even made dissecting needles with minutens
specically for Chandra’s (for that was the name he had been
given in Oman’s lab, and that was how he’d be addressed by
his peers from Europe and the USA from nowon) use. Oman
also introduced him to the techniques of collecting leafhop-
pers in the eld. Chandra could only look in wide-eyed disbe-
lief as Oman took him to various locales and unerringly told
him which species could be found on a specic plant or in a
particular area. Itwas here that he picked up the importance
of collecting a large series of specimens from the complete
range of occurrence of each species.
An early set of diagrams of a deltocephaline leafhopper by
C. A. Viraktamath during his Dharwad days, identies by
Dr. H.M. Harris
After this short stint of a year and a half at OSU, Chandra
returned a edgling taxonomist to the College of Agriculture,
Dharwad. Now began the trying task of locating for study
the types of agalliine leafhoppers of the Indian subcontinent
deposited in different museums, while simultaneously mak-
ing fresh collections from all over India. These studies had
to go hand in hand with the courses that he had to offer at
the college. Afew years after his return from the USA he got
married to Lalitha (nee Shashikala Koranmath) at Dharwad
on May27, 1973. Persistent requests from him soon lead to
his transfer to Bangalore. By then he had been promoted as an
assistant professor.
Itwas in Bangalore that he began his research work in ear-
nest. Thishad to be done in addition to teaching, which, as
will be seen later, was a taxing schedule. To begin, he had to
make his own collection, as leafhopper collections in Indian
museums and colleges were modest or non-existent. Inthose
days, it was notuncommon to see him wielding the outsized
5.5 feet long leafhopper net, designed by Oman—one that was
over twice as long and as many times as heavy as the insect
nets in common use—collecting leafhoppers on the univer-
sity campus and elsewhere in the wilds of India. Since his the-
sis was to include the entire Old World Agalliinae, he had to
source specimens of these leafhoppers from different muse-
ums and make collections, especially from the type localities
from across the country. Lalitha, his wife, was notin the least
resentful of his frequent absences from home in pursuit of
leafhoppers. On the other hand, she cheerfully assisted him
in processing specimens from the substantial catches from
these expeditions. Till Sreedevi, their rst daughter was born
in 1976, they would walk each evening to the laboratory from
home and she would lighten the hours by assisting him in the
tedious task of sorting, mounting, and pinning leafhoppers.
But when Smita, their second daughter was born a few years
later, in 1983, she had no time at all for entomological work.
His rst paper on the taxonomy of a leafhopper was pub-
lished in 1972 (years before he completed his PhD), describing
a new species of Austroagallia from the Galapagos. When he
submitted his thesis in March 1982 titled “AGeneric Revision
of the Old World Agalliinae (Homoptera: Cicadellidae),” sig-
nalling the completion of his PhD, he had published 15 papers
on leafhoppers. Recognition from fellow taxonomists came
fairly early to him when he was invited to present a paper and
chair the last session of the First International Workshop on
Biotaxonomy, Classication and Biology of Leafhoppers and
Planthoppers (Auchenorrhyncha) of Economic Importance
held between 4th and 7th October 1982 under the auspices of
the Commonwealth Institute of Entomology, Commonwealth
Agricultural Bureau International held at London. His paper
titled “Genera to be revised on a priority basis. Theneed for
keys and illustrations of economic species of leafhoppers and
preservation of voucher specimens in recognised institutions”
raised issues of serious taxonomic concern particularly in the
developing world.

xviiChandrashekaraswami Adiveyya Viraktamath
Subsequent studies on the taxonomy of leafhoppers saw him
making repeated visits to the Natural H istory Museum, London
and the National Museum of Natural History, Washington,DC.
Some of these visits were at his own expense as ofcial sources
of funding for taxonomic studies are hard to come by.
The 114 papers on Cicadellidae so far published by
him, some in association with national and international
associates, include 46 revisionary studies and 11 reviews
(Annexure I) of various taxa in addition to describing
a new tribe, and many new genera and species. These
also include a few papers on the biology and host plants
of cicadellids as well as their role as vectors of plant
pathogens. In all, 56 new genera (Annexure II) and 452
new species (Annexure III) have been described by him
and his collaborators. Though he continued to teach and
guide students after his retirement in 2004, he could from
then onwards devote most of his time to taxonomic work.
Thisis reected in the spurt in the number of taxa described
by him and the number of papers published since then.
Over 50 percent of both genera and species were described
in the 15years after his retiring from the university than
in the 32 years prior to that. His publications include
209 research papers and 6 edited books (Annexure IV).
Recognition for his work came in many forms, but the most
cherished for any taxonomist is recognition from his peers.
That CAV got this in no small measure can be gauged
from the number of species that have been named after
him. Twenty-nine species of leafhoppers were named in
his honour by 25 leafhopper taxonomists from laboratories
across the world. Both admiration for his work and gratitude
for his taxonomic assistance prompted 14 taxonomists,
largely from India, to name new Lepidoptera (1 species),
Hymenoptera (2species), and Coleoptera (6 species) after
him (AnnexureV). Inrecognition of his pre-eminence as a
leafhopper taxonomist, he was invited by Dr. Zhang Yalin
of the Northwest A & F University, Yangling, Shaanxi
Province, China, to assist post graduate students in their
studies on the taxonomy of Cicadellidae. All four of these
visits were funded by that university.
Hehad taught entomology for a few years at the under-
graduate level at the College of Agriculture, Dharwad. Itwas
at Bangalore, however, that he came into his own as a teacher.
Here, he began by teaching the “Introductory Entomology”
course to undergraduates in July 1974. While he continued to
offer a few courses in entomology to the undergraduates till
1995, he was soon entrusted the more onerous task of teaching
post graduate students. During the course of the next 30years,
he would offer a range of much sought after courses to those
pursuing their MSc and PhD programmes. These courses
included Insect Morphology, Internal Anatomy, Systematic
Entomology I and II, Immature Insects, Bioecology of Crop
Pests, Literature and Techniques in Entomology, and Insect
Vectors of Plant Pathogens (some of them even for 12 years
subsequent to his retirement) (Annexure VI). Themost sought
after, however, were the courses in systematics. Over the span
of 35years he offered over 90 courses.
Participants of the First International Workshop on Leafhoppers and Planthoppers of Economic Importance held between 4th–7th
October, 1982 at Royal Entomological Society, London (Professor C. A. Viraktamath, standing fth from right)

xviii Chandrashekaraswami Adiveyya Viraktamath
Hewas among the most feared and revered members on the
teaching faculty. His exacting standards and stern demeanour
infused a sense of purpose and the determination to outper-
form oneself in every student. Over the years, however, he
mellowed in his dealings with students with severity giving
way to tact.
While one could complete most other courses by merely
listening to the professor in class, he did notconne his teach-
ing solely to lectures, but gave original scientic papers and
chapters from the classics of taxonomy and evolutionary biol-
ogy as reading assignments. Theworks of Borror and DeLong
were required reading, so were those of R. F. Chapman, Ernst
Mayr, Alvah Peterson, Theodosius Dobzhansky, S. L. Tuxen,
Howard E. Hinton, Willi Hennig, and many others based on
the course being offered. TheInternational Code of Zoological
Nomenclature had to be studied very closely. Practical classes
also changed character under him. Having prevailed over a
very reluctant Head of the Department, he dispensed with
the need for the maintenance of practical record books inthe
course on Insect Systematics. Notonly were these books the
distinguishing feature of any practical class, they were incon-
ceivable without them. Forthe rst time at UAS—and per-
haps in the country—students of insect systematics had to use
keys and identify specimens during their practical classes.
Hebegan assigning “special topics” to motivate students to
collect and study specied insect taxa. Heremoved the limit
of 50 specimens that students were expected to collect for sub-
mission as part of the requirement for the course in Insect
Systematics. Themore diverse and the larger the collection
submitted, the more marks were you likely to get. One more
change he effected was in the pins used in pinning the insects
collected. Insect pins were notgiven to students, as they had
to be imported and were too expensive. The short stubby
ofce pins then in use were replaced by ne sewing needles
with plastic beads afxed to the blunt ends to serve as heads
for ease of handling. Thisminimized damage to specimens,
enabling the retrieval of valuable ones for retention in the
departmental collection.
Itwas under him that the chaotic collection of insects at
the Department of Entomology developed into a well-curated
modern museum. Thecollection was so dear to him that once
while a bomb scare made all others vacate the college build-
ing, he refused to move saying, “If there really is a bomb I
would prefer to die along with the entire insect collection.”
Over the years, synoptic collections of taxa notrepresented in
the collection were obtained from reputed taxonomists from
across the country. Thetotal number of insect specimens in
the collection is currently estimated at about 350,000. Theold-
est specimen, recovered and preserved from the decrepit old
collection at the department, is an avare weevil collected on
October29, 1908, by Leslie C. Coleman, the rst director of
agriculture, Mysore state. The collection also includes type
specimens of all insects described by those working in the
department. He set in place a free identication service for
farmers, students, amateurs, and others desirous of ascertain-
ing the identity of insects of their interest, which he contin-
ues to do even today. Thisgained greater signicance when
the Natural History Museum, London began levying a steep
charge for the identication of insects sent to them, a service
they had hitherto rendered free. Over 275 institutions from
across the country, in addition to a very large number of inter-
ested individuals, have so far availed this facility.
Professor C. A. Viraktamath in the Insect Museum
Hedidn’t have to wait too long after coming to Bangalore
to be made a Major Advisor entrusted with guiding post
graduate students (rst MSc student allotted in 1975) for the
completion of their research topics, a mandatory require-
ment for the completion of their MSc and PhD programmes.
Forty-four students submitted their theses under his guidance
(30MSc students and the remaining, PhD students). Twenty-
eight of these were on the taxonomy of different insect taxa,
eight of which were on leafhoppers. Over half the remaining
theses submitted under his guidance were on the bioecology
and management of insect pests of crops and others in the areas
of pollination biology, insecticide toxicology, studies on light
traps, biological control, and forensic entomology (Annexure
VII). Hedid notshy away from making his students take up
taxonomic studies of taxa other than leafhoppers. Some of
his students who have gone down these alternative paths have
blossomed to become taxonomists of note in taxa as diverse as
Orthoptera, Formicidae, Arctiidae, Chrysomelidae, Carabidae,
Scarabaeidae, Cerambycidae, Coccinellidae, Tephritidae,
Delphacidae, Pentatomidae, Miridae, Reduviidae, Aphididae,
and Apidae.
In addition to three projects on insect taxonomy,
he handled six others (with collaborators) on various
aspects of economic entomology from “assessing the
status of the invasive American leaf miner, Liriomyza
trifolii” and the “utilization of non-edible oils in pest

xixChandrashekaraswami Adiveyya Viraktamath
management,” to the “management of insect pests of gher-
kins” (the last at the instance of the Gherkin Growers’
Association). Thepest management strategy for gherkins
that he developed under this project continues to be in
vogue. Theresults of the project on the “pest status of
the Mexican bean beetle Zygogramma bicolorata, intro-
duced for the biological control of Parthenium hysteroph-
orus L.,” helped stave off a major setback to the weed
biocontrol programme in the country, as opinion in the
higher echelons of the agricultural fraternity was danger-
ously veering towards a blanket ban on the import of all
weed biocontrol agents. The“Network Project on Insect
Biosystematics” which he handled with elan at one of the
Bangalore centres enabled the growth of the departmen-
tal museum to become one of the finest in the country
(Annexure VIII).
Though aficted, since 2014, by Age-related Macular
Degeneration (an afiction of the eyes that blurs vision), he
continues to work unfazed. Anyone stepping into his labora-
tory these days is likely to nd him with a hand lens clenched
tightly in his right st, poring intently, in Holmesian fashion,
over a manuscript or scrutinizing a leafhopper from his col-
lection. Inspite of this impediment, however, there has been
no let-up in the pace of his work. Inthe 4 years since the
diagnosis of this ailment, he has published 17 papers on the
taxonomy of leafhoppers, with at least four of them being
30pages or more in length.
Professor C. A. Viraktamath Examining Leafhoppers
Hazarding the risk of being accused of hagiography, we
assert that CAV remains an enviable brand in the international
taxonomic landscape. Like any enduring brand, he remains
unique and authentic, well known to the people who desire
his services with no doubt having ever been cast on the con-
sistency in the quality of his work. His brand equity and brand
value can only wax in the years to come.
Editors

xxi
Contributors
R. Babu
Southern Regional Centre
Zoological Survey of India
Chennai, India
Dipendra Nath Basu
National Centre for Biological Sciences
Bengaluru, India
Renee M. Borges
Centre for Ecological Sciences
Indian Institute of Science
Bengaluru, India
James M. Carpenter
Division of Invertebrate Zoology
American Museum of Natural History
NewYork, NewYork
Joyshree Chanam
Centre for Ecological Sciences
Indian Institute of Science
Bengaluru, India
K. J. David
Division of Germplasm Collection and Characterisation
ICAR-National Bureau of Agricultural Insect Resources
Bengaluru, India
K. N. Ganeshaiah
School of Ecology and Conservation
University of Agricultural Sciences
Bengaluru, India
Mahua Ghara
Centre for Ecological Sciences
Indian Institute of Science
Bengaluru, India
Kumar Ghorpadé
Department of Entomology
University of Agricultural Sciences
Dharwad, India
G. S. Girish Kumar
Mangalore University
Mangalore, India
P. Girish Kumar
Western Ghats Regional Centre
Zoological Survey of India
Kozhikode, India
Sunil Joshi
Division of Germplasm Collection and Characterisation
ICAR-National Bureau of Agricultural Insect Resources
Bengaluru, India
Anusha Krishnan
Centre for Ecological Sciences
Indian Institute of Science
Bengaluru, India
Krushnamegh Kunte
National Centre for Biological Sciences
Bengaluru, India
S. Manickavasagam
Department of Entomology
Faculty of Agriculture
Annamalai University
Chidambaram, India
S. Palanivel
Department of Entomology
Faculty of Agriculture
Annamalai University
Chidambaram, India
Arati Pannure
Department of Entomology
College of Sericulture
University of Agricultural Sciences
Bengaluru, India
Sunita Patra
Zoological Survey of India
Kolkata, India
David L. Pearson
School of Life Sciences
Arizona State University
Tem p e, A r iz o na
J. Poorani
ICAR-National Research Centre for Banana
Trichy, India
K. D. Prathapan
Department of Agricultural Entomology
Kerala Agricultural University
Trivandrum, India

xxii Contributors
K. Rajmohana
Zoological Survey of India
Kolkata, India
Yuvaraj Ranganathan
Centre for Ecological Sciences
Indian Institute of Science
Bengaluru, India
S. Ramani
ICAR, Department of Entomology
University of Agricultural Sciences
Bengaluru, India
S. Salini
Division of Germplasm Collection and Characterisation
ICAR-National Bureau of Agricultural Insect Resources
Bengaluru, India
Sanjay P. Sane
National Centre for Biological Sciences
Bengaluru, India
Harshada H. Sant
National Centre for Biological Sciences
Bengaluru, India
C. Selvakumar
Department of Zoology
TheMadura College (Autonomous)
Madurai, India
K. Selvaraj
Division of Germplasm Conservation and Utilisation
ICAR-National Bureau of Agricultural Insect Resources
Bengaluru, India
Megha Shenoy
Centre for Ecological Sciences
Indian Institute of Science
Bengaluru, India
S. K. Singh
National Museum of Natural History
New Delhi, India
K. G. Sivaramakrishnan
Zoological Survey of India
Chennai, India
K. A. Subramanian
Zoological Survey of India
Southern Regional Centre
Chennai, India
R. Sundararaj
Forest and Wood Protection Division
Institute of Wood Science and Technology
Bengaluru, India
Martin Streinzer
Department of Neurobiology
Faculty of Life Sciences
University of Vienna
Vienna, Austria
Rajamani Swaminathan
Department of Entomology
Rajasthan College of Agriculture
Maharana Pratap University of Agriculture and Technology
Udaipur, India
Tatiana Swaminathan
Department of Entomology
Rajasthan College of Agriculture
Maharana Pratap University of Agriculture
andTechnology
Udaipur, India
T. Venkatesan
Division of Genomic Resources
ICAR-National Bureau of Agricultural Insect Resources
Bengaluru, India
Vignesh Venkateswaran
Centre for Ecological Sciences
Indian Institute of Science
Bengaluru, India
D. Vimala
Zoological Survey of India
Southern Regional Centre
Chennai, India
Pratibha Ya dav
Centre for Ecological Sciences
Indian Institute of Science
Bengaluru, India

187
12 Potter Wasps (Hymenoptera:
Vespidae: Eumeninae) of India
P. Girish Kumar, Arati Pannure, and James M. Carpenter
INTRODUCTION
Thesubfamily Eumeninae, commonly called potter wasps, is
the most species rich subfamily among the Vespidae. Thiscos-
mopolitan subfamily consists of about 3,794described species
in 205genera (Pannure etal. 2016; Tan etal. 2018). Theyare
usually solitary and rarely sub-social (Ducke 1914; Bohart
and Stange 1965). Adults are small to large and compact
to elongate with a sessile to strongly petiolate metasoma.
The taxonomy and other aspects of natural history of the
Eumeninae are poorly studied in India. Formost species noth-
ing is known about their biology, behaviour, prey associations,
and hence conservation status. Theneed for taxonomic work
on Eumeninae in India is underlined by the lack of adequate
and well-illustrated keys, both at generic and species level
(Pannure etal. 2016). Thischapter aims to bring together all
the fragmentary literature on the subfamily, thereby providing
a review that could serve as a probable basis for increasing the
knowledge of the group.
SUBFAMILY DIAGNOSIS AND KEY CHARACTERS
Subfamily Diagnosis: Eumeninae can be easily distin-
guished from other hymenopterans by a combination
of following morphological features: usually claws
bid (Figure12.1, 1); head with emarginate eyes (2);
middle tibia usually with one spur (3) except in a few
genera; tegula emarginate on inner side to receive the
parategula (4); mandibles long, usually crossing each
other at rest (5); hind wing with anal lobe (6); hind
coxa with a longitudinal dorsal carina (7); pronotum
extending back to tegula (8); pronotal lobe usually sep-
arated from tegula by a distance equal to or less than
length of lobe (9), but distance rarely greater; posterior
lingual plate longer than wide; fore wing longitudi-
nally folded at rest (10), usually reaching at least the
posterior margin of metasomal segment 4; solitary or
sub-social; making earthen pot-like nests, or nests in
burrows or in wood cavities; predatory in nature.
CONTENTS
Introduction ............................................................................................................................................................................... 187
Subfamily Diagnosis and Key Characters.................................................................................................................................187
Key to Indian Genera of Eumeninae .................................................................................................................................... 189
Major Work on Indian Fauna .................................................................................................................................................... 191
Biodiversity and Species Richness ...........................................................................................................................................191
Distribution Patterns in the Indian Context ...............................................................................................................................192
Immature Taxonomy ................................................................................................................................................................. 193
Molecular Characterization and Phylogeny/Barcoding ............................................................................................................ 193
Taxonomic Problems................................................................................................................................................................. 193
Biology ...................................................................................................................................................................................... 193
Economic Importance ............................................................................................................................................................... 194
Collection and Preservation ...................................................................................................................................................... 194
Useful Websites ......................................................................................................................................................................... 195
Conclusion ................................................................................................................................................................................ 195
Acknowledgements ................................................................................................................................................................... 195
References ................................................................................................................................................................................. 195

188 Indian Insects
Subfamily Key Characters: Mesoscutum with parat-
egula; posterior lingual plate longer than wide; tarsal
claws usually bid; solitary or sub-social.
Tribes/Genera: Uncertainties in the tribal and generic
classication of the subfamily Eumeninae in the past
are partly a result of the morphological intricacy of
the group (Carpenter and Cumming 1985; Carpenter
and Garcete-Barrett 2002; Hermes et al. 2014).
Therst generic classication was given by Latreille
(1802), who divided species into three genera, na mely,
Eumenes, Odynerus, and Synagris based on charac-
ters of mandibles, labial and maxillary palpi, and
shape of metasoma. Currently, 205genera are rec-
ognized in the world. Hermes etal. (2014) proposed
the rst tribal division of the subfamily based on
cladistic methods, which include Zethini, Eumenini
FIGURE12.1 Morphology of Eumeninae (Delta pyriforme pyriforme) 1. Tarsal claws in apical tarsal segment 2. Ocular sinus in frontal
view 3. Mid tibial spur 4. Dorsal view of mesosoma showing tegula and parategula 5. Mandibles 6. Anal lobe in hind wing 7. Carina in hind
coxa 8. Pronotal lobe extension in dorsal view 9. Lateral view of pronotum 10. Longitudinal folding in fore wing.

189Potter Wasps (Hymenoptera: Vespidae: Eumeninae) of India
(=Eumenes sensu lato), and Odynerini (=Odynerus
sensu lato), and this is presently accepted as the clas-
sication of the subfamily into tribes. From India,
189species in 47 genera under all the three tribes
have been reported (Table12.1).
Key tO indian genera OF eumeninae
(modied based on keys of Pannure etal. 2016 and Girish
Kumar and Sharma 2015b)
(Themetasomal terga1, 2are denoted as T1, T2,respec-
tively; metasternum 1 denoted as S1; SMC: sub-marginal cell
of fore wing)
1. Metasoma petiolate, T1in dorsal view usually longer,
less than half as wide as T2in dorsal view ................ 2
- Metasoma notpetiolate, T1in dorsal view usually broader,
more than half as wide as T2in dorsal view ..................17
2. Mid tibia with two spurs ............................................. 3
- Mid tibia with one spur ............................................... 5
3. Propodeal orice narrow dorsally; sub-marginal carina
strongly produced, propodeal valvula elongate and free
from sub-marginal carina posteriorly .... Zethus Fabricius
- Propodeal orice broad and rounded dorsally; sub-
marginal carina weakly produced, propodeal valvula
and sub-marginal carina notfree posteriorly .............. 4
4. TI more than twice as long as wide, dorsal surface with
longitudinal striae; labial palpus with three palpomeres
............................................... Calligaster de Saussure
- TI less than twice as long as wide, dorsal surface with-
out longitudinal striae; labial palpus with four pal-
pomeres ........................................ Discoelius Latreille
5. Mesosoma globular, as wide as high .......................... 6
- Mesosoma more or less attened dorso-ventrally, dis-
tinctly longer than high ............................................. 11
6. Clypeus bluntly angular (female) or atly convex
(male) at apex; temple in dorsal view as long as eye
............................................Katamenes Meade-Waldo
- Clypeus weekly to strongly emarginate (female) at
apex; temple in dorsal view shorter than eye .............. 7
7. Pronotum without pretegular carina; T2 with apical
lamella ......................................................................... 8
- Pronotum with pretegular carina; T2 without apical
lamella ......................................................................... 9
8. Mesepisternum with epicnemial carina present;
T2apical lamella strongly and broadly concave medi-
ally ..................................Omicroides Giordani Soika
- Mesepisternum with epicnemial carina absent; T2 api-
cal lamella notconcave medially ..... Eumenes Latreille
9. T1 (petiole length) less than 1.25×length of mesosoma,
never shorter than mesosoma .......... Delta de Saussure
- T1 (petiole length) 1.25×or more than length of meso-
soma .......................................................................... 10
10. Fore wing with prestigma longer than pterostigma;
F11of male long and hooked.................................................
............................................Phimenes GiordaniSoika
- Fore wing with prestigma shor ter or equals pterostigma;
F11of male short, nothooked...........................................
................................. Oreumenoides (Giordani Soika)
11. Axillary fossa oval; propodeal valvula not fused to
sub-marginal carina, sub-marginal carina produced as
a pointed process above valvula; SMC 2basally trun-
cate; body size smaller than 10mm .......................... 12
TABLE 12.1
Tribes and Genera of Eumeninae from India
Tribe Genus No. of Species
Eumenini Delta de Saussure, 1855 7
Eumenes Latreille, 1802 18
Katamenes Meade-Waldo, 1910 2
Omicroides Giordani Soika, 1934 1
Oreumenoides Giordani Soika, 1961 1
Phimenes Giordani Soika, 1992 4
Odynerini Alastor Lepeletier de Saint Fargeau, 1841 4
Allodynerus Blüthgen, 1937 1
Allorhynchium van der Vecht, 1963 5
Ancistrocerus Wesmael, 1836 8
Antepipona de Saussure, 1855 20
Anterhynchium de Saussure, 1863 6
Antodynerus de Saussure, 1855 3
Apodynerus Giordani Soika, 1993 3
Chlorodynerus Blüthgen, 1951 3
Coeleumenes van der Vecht, 1963 3
Cyrtolabulus van der Vecht, 1969 3
Ectopioglossa Perkins, 1912 2
Epsilon de Saussure, 1855 4
Euodynerus Dalla Torre, 1904 4
Gribodia Zavattari, 1912 1
Indodynerus Gusenleitner, 2008 1
Knemodynerus Blüthgen, 1940 4
Labus de Saussure, 1867 4
Leptochilus de Saussure, 1853 2
Lissodynerus Giordani Soika, 1993 3
Malayepipona Giordani Soika, 1993 2
Orancistrocerus van der Vecht, 1963 2
Paraleptomenes Giordani Soika, 1970 6
Parancistrocerus Bequaert, 1925 11
Pararrhynchium de Saussure, 1855 2
Pareumenes de Saussure, 1855 3
Pseudepipona de Saussure, 1856 1
Pseudonortonia Giordani Soika, 1936 4
Pseudozumia de Saussure, 1875 2
Pseumenes Giordani Soika, 1935 1
Rhynchium Spinola, 1806 4
Stenancistrocerus de Saussure, 1863 1
Stenodyneriellus Giordani Soika, 1961 6
Stenodynerus de Saussure, 1863 2
Subancistrocerus de Saussure, 1855 4
Symmorphus Wesmael, 1836 9
Tropidodynerus Blüthgen, 1939 2
Xenorhynchium van der Vecht, 1963 1
Zethini Calligaster de Saussure, 1852 1
Discoelius Latreille, 1809 2
Zethus Fabricius, 1804 6
Total species 189

190 Indian Insects
- Axillary fossa slit-like; propodeal valvula fused to sub-
marginal carina, sub-marginal carina notproduced as
a pointed process above valvula; SMC 2basally acute;
body size more than 10mm ...................................... 13
12. Female with a distinct subcircular fovea below median
ocellus; metanotum unidentate; propodeum not pro-
duced; T1in dorsal view conspicuously swollen in api-
cal half .......................................... Labus de Saussure
- Female without fovea below median ocellus; metanotum
not unidentate, precipitous; propodeum with extensive
horizontal portion, somewhat narrowed apically, behind
the postscutellum, abruptly sloping posteriorly; T1 in
dorsal view notconspicuously swollen in apical half
..................................Cyrtolabulusvan der Vecht
13. Mesepisternum with epicnemial carina present ........ 14
- Mesepisternum with epicnemial carina absent ............16
14. TI basally with transverse carina; ventral margins of
TI touching each other except for posterior diverging
part, thus SI visible only in posterior triangular part
..................................................Ectopioglossa Perkins
- TI without transverse carina; ventral margins of TI
basally close to each other, but nottouching each other
................................................................................... 15
15. Mesoscutum without notauli; S1transversely striate
posteriorly; fore wing with parastigma shorter than
pterostigma ..................... Coeleumenes van der Vecht
- Mesoscutum with notauli; S1irregularly rugose posteri-
orly; fore wing with parastigma longer than pterostigma
........................................... Pseudozumia de Saussure
16. Fore wing with prestigma longer than pterostigma;
female with cephalic foveae .......................................
.......................................... PareumenesdeSaussure
- Fore wing with prestigma shorter than pterostigma;
female without cephalic foveae .....................................
......................................... PseumenesGiordaniSoika
17. SMC 2 petiolate anteriorly; propodeal orice narrow
dorsally ............. Alastor Lepeletier de Saint Fargeau
- SMC 2not petiolate anteriorly; propodeal orice broad
and rounded dorsally ................................................. 18
18. Anterior face of pronotum with two small, deeply
impressed medial pits or foveae; which may be sparse,
contiguous, or faint in some species ......................... 19
- Anterior face of pronotum without medial pits or foveae
...................................................................................24
19. T1with transverse carina .......................................... 20
- T1without transverse carina ...................................... 23
20. T1with two transverse carinae close together at crest
of declivity; T1wider than long in dorsal view............
.................................... Subancistrocerus de Saussure
- T1 with one transverse carina; T1 variable in dorsal
view ........................................................................... 21
21. Anterior face of pronotum with foveae separated; T2
smooth basally, forming acarinarium beneath apex of T1
that is often full of mites .......ParancistrocerusBequaert
- Anterior face of pronotum with foveae coalesced
(foveae contiguous); T2without acarinarium ........... 22
22. T1in dorsal view subsessile, longer than wide, T2much
wider than T1 .......... Pseudonortonia Giordani Soika
- T1sessile; about as wide as T2 .................................
................................StenancistrocerusdeSaussure
23. Anterior face of pronotum with foveae coalesced, punc-
tate laterally, T1subsessile, in dorsal view usually nar-
rower than T2 ...........Paraleptomenes Giordani Soika
- Anterior face of pronotum usually with foveae sepa-
rated, smooth laterally; T1 in dorsal view about as
broad as T2........................Stenodynerus de Saussure
24. T1with transverse carina .......................................... 25
- T1without transverse carina .....................................29
25. TI with broad longitudinal median furrow posterior
to transverse carina; male antenna simple apically;
notauli present; female cephalic foveae well separated,
located midway between posterior ocelli and occipital
margin .....................................Symmorphus Wesmael
- TI without broad longitudinal furrow/groove; mese-
pisternum without epicnemial carina; male antenna
hooked; other characters variable; female cephalic
foveae closely spaced, nearer occipital margin than
posterior ocelli.................. .......................................... 26
26. Axillary fossa oval, broader than long; tegula exceed-
ing parategula ........................ Ancistrocerus Wesmael
- Axillary fossa narrower than long, slit-like; tegula
notexceeding parategula .......................................... 27
27. Propodeal dorsum forming shelf-like area behind meta-
notum; fore wing with prestigma less than half as long as
pterostigma ........................ Pararrhynchium de Saussure
- Propodeal dorsum below plane of metanotum, sloping
posteroventrally; prestigma more than half as long as
pterostigma, measured along posterior part ............. 28
28. T2 with well-developed apical lamella...........................
.................................... LissodynerusGiordani Soika
- T2without apical lamella ..............................................
....................................OrancistrocerusvanderVecht
29. Tegula evenly rounded posteriorly, not emarginate
adjoining parategula and usually notreaching apex of
latter ................................... Tropidodynerus Blüthgen
- Tegula not evenly rounded posteriorly, emarginate
adjoining parategula and often reaching or surpassing
apex of latter .............................................................30
30. T1with transparent or translucent apical border ......... 31
- T1without transparent or translucent apical border.....
......................................................................... 34
31. Parategula notvisible from above; tegulae posteriorly
bent inwards ........................ Knemodynerus Blüthgen
- Parategula visible from above ................................... 32
32. Metanotum semicircular in shape from above, carinate
posteriorly .......................... Antodynerus de Saussure
- Metanotum notsemicircular in shape from above ... 33
33. Pronotum with anterior face densely punctate laterally,
without dorsal carina ...........Chlorodynerus Blüthgen
- Pronotum with anterior face notdensely punctate, with
dorsal carina .........................Euodynerus Dalla Torre
34. Metanotum bidentate ................................................ 35

191Potter Wasps (Hymenoptera: Vespidae: Eumeninae) of India
- Metanotum notdentate ............................................. 36
35. Clypeus higher than wide; mid-anterior face of prono-
tum smooth and with short transverse rugae; T1dis-
tinctly narrower than T2; male terminal antennal
segment small ................. Apodynerus Giordani Soika
- Clypeus wider than high; mid-anterior face of prono-
tum usually densely punctate and with an upper trace
of transverse carina; T1 slightly narrower than T2;
male term inal anten nal segment relatively large.............
.............................................Antepipona de Saussure
36. T2with apical lamella ............................................... 37
- T2without lamella .................................................... 39
37. T1depressed subapically, gradually widened with lat-
eral sides divergent in dorsal view; propodeum with
sub-marginal carina projecting as rounded lobe above
valvula, bilamellate; epicnemial carina absent; axil-
lary fossa oval, broader than long..................................
........................................... LeptochilusdeSaussure
- T1not depressed subapically, usually with lateral sides
roughly parallel in dorsal view; propodeum with sub-
marginal carina notdifferentiated from valvula, mono-
lamellate (except Epsilon); epicnemial carina present;
axillary fossa narrower than long, slit-like ............... 38
38. Palpal formula 5:3; male vertex sometimes with large
and deep depression; propodeum with well developed
lateral carinae; T1-5 each with apical lamella.......
..................................................... Gribodia Zavattari
- Palpal formula 6:4; male vertex without large and
deep depression; propodeum without well developed
lateral carina; only T2with apical lamella..................
................................................ EpsilondeSaussure
39. Tegula never exceeds parategula posteriorly; axillary
fossa much narrower than long, sometimes slit-like
................................................................................40
- Tegula usually exceeds parategula posteriorly, or at
least equalling it; axillary fossa oval, at least as wide as
long ............................................................................ 44
40. Fore wing with prestigma half or less than the length of
the pterostigma ........... StenodyneriellusGiordaniSoika
- Fore wing with prestigma more than half the length of
the pterostigma, often nearly equal .......................... 41
41. SMC 3separated from the apex of marginal cell by
distance shorter than its minimum width.....................
...................................Allorhynchium van der Vecht
- SMC 3 separated from the apex of marginal cell by
distance longer than its minimum width .................. 42
42. Scutum posteriorly and scutellum anteriorly smooth
and impunctate ............................. Rhynchium Spinola
- Scutum and scutellum punctate ................................ 43
43. Mesepisternum without epicnemial carina......................
...........................................Indodynerus Gusenleitner
- Mesepisternum with epicnemial carina present..............
....................................... Anterhynchium de Saussure
44. T1subsessile, in dorsal view narrower than T2, wider
apically than basally.......Malayepipona Giordani Soika
- T1sessile, in dorsal view about as wide as T2 ......... 45
45. Metanotum projecting over propodeum; propodeum
with sclerotized dorsolateral projections; propodeum
with sub-marginal carina not differentiated from
valvula; SMC 2with second recurrent vein far from
SMC3 ........................Xenorhynchium van der Vecht
- Metanotum not projecting over propodeum; propo-
deum without dorsolateral projections; propodeum
with sub-marginal carina projecting as rounded lobe
above valvula; propodeal valvula free from sub-
marginal carina posteriorly ....................................... 46
46. Tegula narrower and longer, surpassing parategula pos-
teriorly.[female vertex with reniform fovea, about as
wide as ocellar triangle; hind coxa with ventral lobes]
.................................................. Allodynerus Blüthgen
- Tegula broad, equal to parategula posteriorly.........
.......................................... Pseudepipona de Saussure
MAJOR WORK ON INDIAN FAUNA
TheIndian Eumeninae was rst comprehensively studied by
Bingham (1897). Many taxonomic changes were made after
his studies especially at generic level. Most of the earlier
genera like Eumenes, Odynerus, etc. have been split into
different genera like Eumenes into Coeleumenes, Delta,
Eumenes s. str., Oreumenoides, Pareumenes, Phimenes,
Pseumenes, etc. and Odymerus into Ancistrocerus,
Antepipona, Antodynerus, Apodynerus, Epsilon, Odynerus
s. str., Paraleptomenes, Subancistrocerus, etc. After
Bingham (1897), only scattered papers were published.
Cameron (1897, 1902, 1903a, 1903b, 1904, 1907, 1908,
1909, 1913), Nurse (1903, 1914), Rothney (1903), Paiva
(1907), Meade-Waldo (1910a, 1910b), Dover (1921, 1925),
Dover and Rao (1922), van der Vecht (1937, 1959a, 1959b,
1961, 1963, 1969, 1981), Giordani Soika (1941, 1960, 1966,
1982), Wain (1956), Gusenleitner (1988, 1996, 1998, 2001,
2006, 2007, 2008), Giordani Soika and Khan (1991), and
Krombein (1978, 1991) are some of the important work
published through the twentieth century containing descrip-
tions of Eumeninae. During the last two decades, Lambert
and Narendran (2002), Lambert (2004), Lambert et al.
(2007, 2008), Srinivasan and Girish Kumar (2010, 2013),
Girish Kumar (2011, 2012a, 2012b, 2012c, 2013a, 2013b,
2013c), Girish Kumar and Lambert (2011, 2012), Girish
Kumar and Sharma (2012, 2013, 2014, 2015a, 2015b), Girish
Kumar and Carpenter (2013, 2015a, 2015b), Girish Kumar
et al. (2013a, 2013b, 2013c, 2013d, 2014a, 2014b, 2014c,
2015a, 2015b, 2016a, 2016b, 2016c, 2016d, 2017a, 2017b,
2017c), Mohammed Shareef etal. (2013), Girish Kumar and
Sureshan (2016), Pannure etal. (2016, 2017, 2018), and Selis
(2018) have made contributions to the Indian eumenid fauna.
BIODIVERSITY AND SPECIES RICHNESS
India has four biodiversity hotspots, the Western Ghats, the
Himalayas, the Indo-Burma region, and Sundaland (which
includes the Nicobar group of islands). Most of the faunal

192 Indian Insects
elements in India are very signicant and peculiar with many
endemic species. The diversity and species richness of pot-
ter wasps is fairly high in India with 189species (about 4.8%
of the world species) and many additional subspecies under
47 genera in 3 tribes (Table 12.1). Several workers during
the past decade have been exploring the potter wasp fauna
of the Indian subcontinent mainly in the north and north-
eastern regions and have described a number of new species
and found new records from this region (Girish Kumar 2011,
2012a, 2012c; Girish Kumar and Carpenter 2013, 2015a,
2015b; Girish Kumar and Lambert 2011; Girish Kumar etal.
2013a, 2013b, 2013c, 2013d, 2014a, 2014b, 2015, 2016a, 2016b,
2016c, 2016d, 2017a, 2017b) (Table12.2). Ina series of studies,
17species in 12genera have been reported from Rajasthan
(Girish Kumar and Sharma 2014), 19species in 11genera from
Chhattisgarh (Girish Kumar and Sharma 2015b), and 11spe-
cies from Arunachal Pradesh (Srinivasan and Girish Kumar
2010). Girish Kumar (2012) recorded the genus Omicroides
for the rst time from NE India. Girish Kumar etal. (2017c)
recorded an unreported genus Pseudepipona from India and
the species Pseudepipona (Pseudepipona) vicina Gusenleitner,
1973 from the northern Himalaya. North and northeastern
regions seem to be more diverse than the south Indian region.
Atotal of 72species and subspecies in 31genera have been
reported from South India, and it appears like the number of
species will denitely see an increase (Pannure et al. 2016).
Eumenine fauna from most Indian states such as Andhra
Pradesh, Bihar, Delhi, Goa, Gujarat, Haryana, Jammu and
Kashmir, Jharkhand, Lakshadweep Islands, Madhya Pradesh,
Maharashtra, Manipur, Mizoram Nagaland, Odisha, Punjab,
Uttar Pradesh, Telangana, and Tripura is very poorly and frag-
mentarily known. Considering the fact that Western Ghats and
Eastern Ghats and Andaman and Lakshadweep Islands are
still untapped regions, diversity of potter wasp species in the
region should denitely be higher.
DISTRIBUTION PATTERNS IN
THE INDIAN CONTEXT
Accounts of the distribution patter ns of eumenine wasps within
India is lacking. From the geological standpoint, the Indian
fauna is comprised of two components, namely, the major
Oriental component (represents most of India) and the minor
Palearctic component (mainly northern Himalayas). Some
genera such as Allorhynchium, Antepipona, Anterhynchium,
Antodynerus, Delta, Eumenes, Paraleptomenes, Phimenes,
Rhynchium, Stenodyneriellus, and Subancistrocerus are
widely distributed within India (Srinivasan and Girish
Kumar 2010; Pannure etal. 2016). Genera such as Alastor,
Cyrtolabulus, Epsilon, Euodynerus, Knemodynerus, Labus,
Oreumenoides, and Pareumenes are usually not com-
mon, but occur in different parts of India (Pannure et al.
2016). The genus Xenorhynchium is widely distributed, but
not recorded from northeastern India. Many genera such
as Apodynerus, Coeleumenes, Discoelius, Ectopioglossa,
Parancistrocerus, Pseudozumia, Pseumenes, and Zethus
show a peculiar distribution pattern with presence in the
southern part of India and the north-eastern part of India.
Thenorthern part of India has rich biodiversity and Palearctic
elements. Many genera such as Allodynerus, Calligaster,
Gribodia, Malayepipona, Omicroides, Orancistrocerus, and
Pararrhynchium are restricted to north-east India. Thegenus
Indodynerus generally occurs in the southern part of India
(Girish Kumar et al. 2013a). The genus Lissodynerus has
so far been recorded from the Andamans, South India and
Sikkim (Girish Kumar etal. 2015b; Selis 2018). Thegenera
Ancistrocerus, Katamenes, and Symmorphus mainly occur in
the Himalayas. Thegenera Chlorodynerus, Leptochilus, and
Stenancistrocerus have been recorded from western India
only. Thegenus Tropidodynerus has been recorded from west,
north, and north-eastern India (Girish Kumar et al. 2013d).
TABLE 12.2
Recent Records of Eumeninae from India
Species Year Distribution Reference
1Subancistrocerus venkataramani Girish Kumar 2013 Jharkhand Girish Kumar (2013a)
2Epsilon manasicum Girish Kumar& Carpenter 2014 Assam Girish Kumar etal. (2014a)
3Paraleptomenes darugiriensis Girish Kumar, Carpenter& Sharma 2014 Arunachal Pradesh, Assam,
Meghalaya, Sikkim, West Bengal
Girish Kumar etal. (2014b)
4Alastor (Alastor) venkataramani Girish Kumar & Carpenter 2015 Telangana Girish Kumar and Carpenter (2015a)
5Leptochilus (Neoleptochilus) hassani Girish Kumar & Carpenter 2015 Maharashtra Girish Kumar and Carpenter (2015b)
6Lissodynerus rutlandicus Girish Kumar, Srinivasan& Carpenter 2015 South Andaman (Rutland Island) Girish Kumar etal. (2015b)
7Allorhynchium tuberculatum Girish Kumar & Carpenter 2016 Kerala Girish Kumar etal. (2016a)
8Parancistrocerus jaferpaloti Girish Kumar & Carpenter 2016 Kerala Girish Kumar etal. (2016c)
9Parancistrocerus loharbandensis Girish Kumar & Carpenter 2016 Assam
10 Parancistrocerus turensis Girish Kumar & Carpenter 2016 Meghalaya
11 Discoelius vasukii Pannure& Carpenter 2017 Tamil Nadu Pannure etal. (2017)
12 Pararrhynchium venkataramani Girish Kumar& Carpenter 2017 Arunachal Pradesh, Meghalaya Girish Kumar etal. (2017b)
13 Antepipona tricolorata Selis 2018 Sikkim Selis (2018)
14 Lissodynerus unicus Selis 2018 Sikkim
15 Symmorphus (Symmorphus) incisus Selis 2018 Sikkim

193Potter Wasps (Hymenoptera: Vespidae: Eumeninae) of India
Thegenus Pseudonortonia has been recorded from western,
southern, and north-eastern India. The genus Stenodynerus
has been recorded from south, north, and north-eastern
India. The genus Pseudepipona has been recorded from the
northern Himalayas. The genus Xenorhynchium is endemic
to the Indian subcontinent (Girish Kumar and Kishore 2011;
Pannure etal. 2016).
IMMATURE TAXONOMY
Thetaxonomy of eumenid wasps rests largely on the exter-
nal morphology of the adults. There is hardly any informa-
tion available for the immature stages, even though they could
be useful. Thereare only about 2%–3% of known species for
which larval descriptions are available. Themost important
descriptions of immature stages, mainly larvae of Palearctic
Eumeninae species, are provided by Tormos et al. (1997a,
1997b, 1997c, 1998, 2005, 2008). Identication of Eumeninae
at the generic and specic levels using larval characters is
difcult (Grandi 1961; Evans 1977; Tormos etal. 1998). Itis
therefore imperative to conduct detailed descriptive stud-
ies that will help to know characters that dene the taxa at
genus and species level. Itis believed that such studies will
contribute greatly to the clarication of the phylogeny and,
hence, the systematics of Eumeninae (Tormos etal. 2008).
The morphology of immature stages of Oriental species is
largely unknown.
MOLECULAR CHARACTERIZATION
AND PHYLOGENY/BARCODING
The efforts made to achieve a robust established classica-
tion that reects evolutionary relationships in Eumeninae
has been very limited. Hermes et al. (2014) proposed the
rst formal tribal division of the subfamily based on cladis-
tic methods into Zethini, Eumenini (=Eumenes sensu lato),
and Odynerini (=Odynerus sensu lato). Inconict with this
tribal subdivision of Eumeninae, Bank et al. (2017), based
solely on a molecular approach, proposed subfamily ranks
for the two major clades of “Zethini”: Raphiglossinae and
Zethinae. Eumeninae, a primary monophyletic lineage of
the Vespidae, turned out to be a paraphyletic group based on
their phylogenomic approach (Bank etal. 2017), which, how-
ever, was based on a limited taxonomic sample. Thephyloge-
netic relationships among the major lineages of Eumeninae
are little investigated for two main reasons: few taxonomists
are working on the group compared to those who are work-
ing on social subfamilies, Polistinae and Vespinae; and the
remarkable diversity found among the Eumeninae wasps,
which has resulted in a troubled taxonomic history (Hermes
etal. 2014). In general, comprehensive investigations of the
phylogenetic relationships of Eumeninae are very scarce for
the Oriental fauna, compared to the Nearctic (Carpenter and
Cumming 1985), Neotropical (Hermes and de Oliveira 2016),
and Palearctic Region (Vernier 1997) fauna. Arecent study by
Hermes etal. (2014) included 18species of the Oriental fauna.
Another effort was made by Nugroho (2015) to study the potter
wasps with a petiolate metasoma (Vespidae, Eumeninae) in
the Indonesian Archipelago based on a phylogenetic analysis
at the generic level. But for these two studies, no efforts have
been made to provide a natural classication/phylogenetic
analysis of Oriental Eumeninae and nothing at all with refer-
ence to the Indian fauna at the generic level. Comprehensive
work is required, supplementing the previous work in other
regions with an analysis of the Indian fauna.
TAXONOMIC PROBLEMS
Taxonomic studies on Eumeninae have been limited with no
intensive research on this group. No revisionary work is avail-
able on this subfamily from India, and even faunal studies in
the Indian region have received little attention. Major contri-
butions on Indian fauna were done by Giordani Soika (1941,
1960, 1966, 1982, 1991) and Gusenleitner (1988, 1996, 1998,
2001, 2006, 2007, 2008). No updated key to the genera and
species of Indian Eumeninae is available. Recently, Srinivasan
and Girish Kumar (2010), Girish Kumar and Sharma (2015b),
and Pannure etal. (2016) have published generic keys to the
fauna of Arunachal Pradesh, Chhattisgarh, and South India,
respectively.
BIOLOGY
Nesting: An amazing plasticity of nesting habits is shown
by Eumeninae. Theyowe the common name “potter
wasps” to the fact that some of these species build more
or less free-standing mud nests looking like earthen
pots. Free-standing mud nests may be unicellular or
multicellular (Isely 1914). Inaddition to species that
construct their nests with mud or masticated plant
material, there are species of Eumeninae that exca-
vate the soil or occupy and modify pre-existing cavi-
ties (Iwata 1976; Evans 1966; Cowan 1991; Nugroho
etal. 2016). Infact, the primitive and the most com-
mon nesting type of wasp is the “renting”, nesting in
pre-existing cavities. Themost commonly used cavi-
ties are borings found on decaying wood material,
but some species also use other structures such as
hollow plant twigs and stems, articial cavities on
man-made structures and ceilings, old mud nests,
small cavities on rocks, walls, or concrete slabs,
and even in deserted ground burrows of other acu-
leates (Carpenter and Cumming 1985; Cowan1991).
They easily accept articially prepared trap nests
such as burrows drilled into the wood blocks or
bundles of cut hollow reed or plastic tubes, greatly
facilitating the study of their biology (Medler and
Fye 1956; Parker and Bohart 1966; Krombein 1967;
Jayasingh and Freeman 1980; Budrienè etal. 2004;
Buschini and Buss 2010; Fateryga 2013a). Several
species dig their own ground burrows, and of these
some construct a mud tube or “turret” over the nest
entrance (e.g., Evans 1956). A few species build
free-standing mud nests on the surfaces of rocks or

194 Indian Insects
plant twigs (e.g., Evans 1977). Based on the nest-
ing habits, they may be classied into three types:
excavators, renters, and builders (Maindron 1882;
Iwata 1976). Theconstruction material and form of
nests are inuenced by the availability of nest sites
and construction materials, as well as the ability of
particular designs to thwart nest parasites and preda-
tors (Cowan 1991; Hermes etal. 2013). Forexample,
the turret of burrowing species functions to pro-
tect against ants (Fateryga 2013b). A few species
are even polymorphic in nest construction (Cooper
1979; Hermes etal. 2015). Nesting behaviour of most
Indian species have notbeen studied. Jayakar (1963)
and Jayakar and Spurway (1967) observed the nest
building of Delta esuriens (Fabricius). Jayakar and
Spurway (1965) observed the nesting behaviour of
Delta conoideum (Gmelin). Jayakar and Spurway
(1971) reported the nesting of Pareumenes brevi-
rostratus (de Saussure), involving a primitive form
of cooperation. Srinivasan and Girish Kumar (2009)
described the nesting behaviour of Xenorhynchium
nitidulum (Fabricius).
Mass Provisioning vs Progressive Provisioning:
Most of the Eumeninae wasps provision their nests
with caterpillars or coleopteran larvae. Each cell
within a nest will be provisioned with several prey
items before the egg hatches (“mass provisioning”)
so that the hatchlings will be provided with ready-
made food in their nest itself. Cells giving rise to
female wasps usually receive a greater amount of
food than the cells containing males, which usu-
ally are smaller (Buck etal. 2008). However, some
wasps show primitive signs of social behaviour
(pre-social), such as Xenorhynchium nitidulum,
Paraleptomenes miniatus, Eumenes pomiformis,
etc. (Deleurance 1946; Jayakar and Spurway 1966;
Krombein 1978; West-Eberhard 1987), which prac-
tice progressive provisioning, especially when there
is scarcity of prey (Cowan1991). Ina series of stud-
ies, Jayakar and co-workers have studied nesting
biology and behaviour of a few Indian eumenine
wasps (Jayakar 1963, 1966; Jayakar and Spurway
1964, 1965, 1966, 1967, 1968, 1971). Srinivasan and
Girish Kumar (2009) also studied some behavioural
aspects, such as provisioning of the nest, of the pot-
ter wasp Xenorhynchium nitidulum (Fabricius).
Sex Determination: Like other hymenopterans, sex
determination in Eumeninae is through haplodip-
loidy. The males are smaller than females with a
shorter larval/pupal development time than the
females (Buck et al. 2008). Inmixed-sex nests of
renting wasps, females develop in the inner cells of
the nest and emerge after the males, which develop
in the outer cells (Buck et al. 2008). Most of the
species spend more than one generation per nest.
Usually there is only one generation, with the nest
then abandoned after emergence. Only where there
is progressive provisioning is there overlap of gen-
erations (Field 2005).
Mite Association: Association of Eumeninae with
some mite species is an interesting phenomenon
observed in certain genera (Krombein 1961; Cooper
1954). Insome genera, both male and female wasps
carry the mites in a specialized region called the
acarinarium. In the genus Parancistrocerus, this
mite chamber (acarinarium) is formed by the trans-
versely depressed base of second tergum which is
usually covered by the apical portion of rst tergum.
These have long been considered as morphologi-
cal adaptations to securely transfer benecial mites
into nests, and thus are thought to be the product of
a mutualistic relationship where mites protect potter
wasps against natural enemies (Okabe and Makino
2008, 2011). However, detailed investigations have
notbeen made to understand the association of mites
with potter wasps.
Strepsipteran Association: An interesting phenom-
enon of parasitic association of strepsipteran insects
under the metasomal tergum is observed in many
species of Eumeninae (e.g., Salt and Bequaert 1929;
Girish Kumar and Carpenter 2013).
Prey Associations: Larvae of Coleoptera and
Lepidoptera form the majority of the prey of sev-
eral Eumeninae which provision their nests (Melo
etal. 2011), but a few species like Paragymnomerus
signaticollis tauricus (Kostylev) take sawies
(Fateryga 2018). The knowledge of prey-predator
associations in Eumeninae is scanty and fragmentary
(Krombein 1967; Itino 1992; Callan 1993).
ECONOMIC IMPORTANCE
They are chiey predators of many insect larvae of
Lepidoptera (Geometridae, Tortricidae) or Coleoptera
(Chrysomelidae and Curculionidae), some of which are
pests of agricultural importance (Cowan 1991). They can
be potentially useful biological control agents for some her-
bivorous insects such as the spruce budworm (Jennings and
Houseweart 1984), other caterpillars (Fye 1965), larvae of
alfalfa weevils (Bohart et al. 1982), or leaf beetles (Sears
et al. 2001). They may also play a signicant role as pol-
linators in the environment, as their ower preferences are
marked over a season (e.g., Cooper 1953). Trap-nesting potter
wasps can be used as bioindicators of environmental change
(Tscharntke etal. 1998).
COLLECTION AND PRESERVATION
Specimen Collection: Adult specimens for taxonomic
studies can be collected by using different methods
such as sweep nets, aspirators, malaise traps, yellow
pan traps, rearing of adults from the nests, etc. Adult
insects can also be collected from the nests by hand
picking or using vials.

195Potter Wasps (Hymenoptera: Vespidae: Eumeninae) of India
Preservation: Collected specimens can be put directly
from the net into vials of 70% alcohol. Long term
preservation in alcohol can be improved by freez-
ing. The alcohol should be changed periodically so
as to prevent damage. Theadults from alcohol can be
mounted or pinned whenever convenient. Usually very
small specimens are mounted on cards in such a way
that all characters are visible easily. Larger specimens
are pinned by using standard entomological pins, pass-
ing through the mesosoma from dorsal side. Thestruc-
tures like genitalia are mounted on micro slides.
Dissecting and studying genitalia can be performed by allow-
ing preserved specimens in a relaxing chamber for a few hours
till they become exible for genitalia extraction. Thegenital
capsule can be taken off with a forceps and entomology pins,
then immersed in 10% potassium hydroxide, and heated at
medium temperature (50°C) for clearing. The genitalia can
then be immersed in acetic acid to neutralize the bleaching
and either temporary or permanent slides prepared for study.
USEFUL WEBSITES
1. http://www.zobodat.at (Gusenleitner 2015). Thissite
is prepared and maintained by Landesmuseum,
Zurich and Biologiezentrum, Linz. This is a very
useful site for getting information regarding species
of Eumeninae.
2. http://data.gbif.org/search/taxa/ The Global
Biodiversity Information Facility (GBIF) is an inter-
national network aimed at providing open access
to data about all types of life on earth. Thissite is
coordinated through its secretariat in Copenhagen,
and India is an associate participant from Asia since
2003. Thisis also a very useful site for information
on species of Eumeninae.
3. http://www.eol/pages/Encyclopedia of Life is another
very useful site for getting information about life on
eart h. Thiswebsite is maintai ned by the Field Museum,
Harvard University, MacArthur Foundation, Marine
Biological Laboratory, Missouri Botanical Garden,
Sloan Foundation, and Smithsonian Institution.
Information about species of Eumeninae is available.
CONCLUSION
Taxonomy and natural history studies on Indian Eumeninae
have notbeen undertaken for many genera, and a comprehen-
sive revision is needed in order to clarify the status of prob-
lematic species and subspecies. Forexample, Allorhynchium
argentatum (Fabricius, 1804) and Allorhynchium metallicum
(deSaussure, 1852) are very similar except some minor dif-
ferences in the intensity of punctures on abdominal tergum.
Several subspecies are required to be studied in great detail
to conrm the taxonomic status and determine their actual
status as subspecies/species. For example, Anterhynchium
(Anterhynchium) abdominale abdominale (Illiger, 1802) and
Anterhynchium (Anterhynchium) abdominale bengalense (de
Saussure, 1855) show noticeable difference in colour patterns
on their abdominal tergum, but intermediate forms are also
available. Studies on the taxonomy of potter wasps in India
have been very scanty and fragmentary compared to other
Asian countries, but it has progressed in very recent years.
Earlier workers like Cameron (1897, 1902, 1903a, 1903b,
1904, 1907, 1908, 1909, 1913), Nurse (1903, 1914), Rothney
(1903), Paiva (1907), Meade-Waldo (1910ab), Dover (1921,
1925), and Dover and Rao (1922) have published accounts of
some species. Thestudies on the Eumeninae in Asian coun-
tries like China and Japan have been impressive as compared
to Indian fauna (Yamane 1990; Zhou et al. 2011). Studies
on the taxonomy of Eumeninae in India have made prog-
ress in the last few years. Many common genera have been
reviewed (Girish Kumar 2013a, 2013b, 2013c; Girish Kumar
and Sharma 2012, 2013, 2015a; Girish Kumar and Carpenter
2013; Girish Kumar etal. 2014a, 2014b, 2015a, 2016b, 2016c,
2017a, 2017b; Pannure etal. 2016). However, all the regions
of India have notbeen thoroughly surveyed, and such sur-
veys might help in unravelling new distributional records for
several species. Thereis a need for furthering investigations
regarding diversity, distribution, zoogeography, ecology, and
biology of Indian Eumeninae.
ACKNOWLEDGEMENTS
The authors are grateful to Dr. Kailash Chandra, Director,
Zoological Survey of India, Kolkata and to Dr. P. M.
Sureshan, Ofcer-in-Charge, Western Ghat Regional Centre
of Zoological Survey of India, Kozhikode, for providing
facilities and encouragement. Thesecond author (AP) grate-
fully acknowledges the University of Agricultural Sciences,
Bengaluru for providing facilities and encouragement.
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