D P Abrol, R K Thakur, H D Kaushik and Sunita Yadav

'i/J ALL INDIA COORDINATED RESEARCH PROJECT ON HONEY BEES AND POLLINATORS

Published:June,2013

Technical Bulletin NON-AP/S BEE POLLINATORS by D PAbrol, R K Thakur, H D Kaushik & Sun ita Yadav

Printed at Dorex Offset Printers, D.N. College Road, Hisar- 125 001

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D P Abrol R K Thakur H D Kaushik Sunita Yadav

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. ALL INDIA COORDINATED RESEARCH PROJECT ON I I .

. <·l1' •, '·,/I\\· HONEI BEES AND POLLINATORS

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Dr. S. AYYAPPAN SECRETARY & DIRECTOR GENERAL

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GOVERNMENT OF INDIA DEPARTMENT OF AGRICULTURAL RESEARCH & EDUCATION

AND INDIAN COUNCIL OF AGRICULTURAL RESEARCH

MINISTRY OF AGRICULTURE, KRISHI BHAVAN, NEW DELHI-110 114 Tel.· 23382629; 23386711 Fax 91-11-23384nJ

E-mai : dg.icar@nic.tn

The imp01tance of pollination in agriculture has been recognized for millennia. However, pollinator decline is being observed due to changing environmental conditions worldwide resulting in pollination crisis thus raising a serious concern to agricultural production and conservation and maintenance ofbiodiversity.

Possible drivers for the decline of insect pollinators include habitat loss, intensive land use, pollution including pesticides, loss of the honeybee's genetic diversity, and detrimental beekeeping practices and climate change. This demands a response from land managers, conservationists and political decision makers to the impending 'global pol linator crisis'. Understanding the causes of pollination fai lure in plants can aid the successful conservation and recovery of rare plants, maintenance of crop yie lds, and sustainable use of wild plant resources such as forest timber. Feasible conservation strategies involve making efforts to protect or restore p lant resources and native pollinators, and the creation of new protected natural areas, which ensures food provision, mating and nesting sites for pollinators.

Pollinator diversity is immense. There are more than 20,000 pollinating bee species in the world, as well as numerous other insect and ve1tebrate pollinators. Despite being represented in large numbers their role as pollinators has little been understood. Where there are ve1y specific niche requirements for the plants and their pollinators, loss of the pollinator can have cascading effects across the ecosystem. Honey bees continue to provide generally satisfactory and frequently excellent pollination of most of our introduced flowering crops. A few crops exhibiting obviously specialized pol linator needs, such as Lucerne and perhaps red clover, have been catered for by the introduction of specialist bees. Where numbers of these bees have been adequate, crop yields have increased. For agricultural as a whole, the diversification of pollination assemblages for crops is clearly important.

Wild and domesticated non-Apis bees effectively complement honey bee pollination in many crops. I am happy to learn that efforts have been made to provide information on the management of various non-Apis bee pol linators in the fonn of a technical bu I let in Non- A pis Bee Pollinators. Project Coordinator AICRP (HB&P) and his team deserve appreciations for their efforts to bring out this comprehensive publication from A ll Indi a Coordinated Research Project on honey Bees and Poll ina tors.

Dated the I Oth June, 20 13 New Delhi

1k: (S. Ayyappan)

J

Prof. Swapan Kumar Datta Deputy Director General (Crop Science)

Indian Council of Agricultural Research Krishi Bhavan, Dr. Rajendra Prasad

Road, New Delhi-1 1 0001

For any nation's progress and prosperity, five factors are most important i.e. land, \ovater, environment forest and biodiversity. Many ecosystems, including agro-ecosystems, depend on pollinator diversity to maintain overall biological diversity. Survival of many plant species is dependent on pollinators. In their absence, it would not have been possible for us to enjoy so many types of food grains, fruits, vegetables and other materials produced by plants. Most of the pollinators are wild bees belonging to different genera. Tt1 India, much ofth work has been undertaken on various aspects of honey bees (Apis spp) alone, however, in nature there a lso exist a large number of solita1y bees such as mining bee, leafcutter bee, alkali bee, carpenter bee, etc also which play an impmtant role in pollination of various crops and flowering plants growing in the wi ld.

WORLD WlTHOUTBEES .......... ..

"If the bee disappeared offthc surface of the globe, then man would only have four years of life left No more pollination; no more plants. no more animals. no more man."

-Albert Einstein

Poll.inators provide pollination services that are crucial for the productivity of agricultural and natural ecosystems. lt has been estimated that over three quarters of the world's crops and over 80% of all poll inators to agricultural crops has been estimated at about USD 200 billion. However, pollinators arc currently under threat with declines in pollinator populations and diversity occurring worldwide. This loss of pollinator habitats and modern agricultural practices, which are dominated by the excessive and ind iscriminate use of pesticides and other agrochemicals. There is global concern that if the decline continues it could have an adverse impact on sustainable agricultural production. Warnings have been insecticides, monoculhire, changes in cropping patterns and pollution, resulting in to depletion of population of useful poLlinating insects, is threatening to reduce our total supply offood by 1/3.

Realizing the importance of poll inators, the present effo1ts put by the authors to compile information with regard to nest architechJre and rearing technologies of various non-Apis bee poll inators is worth appreciati.ng and compliment the excellent efforts made by A1CRP on Honey bees and Pollinators in g iving a right direction for the movement of research on pollinators. This technical bulletin on "Non-A pis Bee Pollinators encompasses the theory and practice of diversity of pollinators in India, value of wild bees as pollinators. their foraging ecology. habitat diversity. domestication, mass rearing and conservation strategies. etc. It is expected that the bulletin will serve as a scientific guide fo r the scientists, teachers, field staff and beekeepers to understand the ecological requirements and behavior of non-Apis bee pollinators so that their ro le could be recognized not only for increasing crop yields but for maintaining stabi lity of our ecosystem also.

f Ct :_D ~(fr-(Swapan K. Datta)

l -

. ~. ~ . ~ .. -

Dr. T.P. Rajendran Assistant Director General (Plant Protection)

mp~ I CAR

Indian Council of Agricultural Research 215, Krishi Bhavan, Dr. Rajendra Prasad

Road, New Delhi-11 0014

Pollinator bio-resource has been natural gift for production of valuable

commodities for human/animal life. Nah1ral pollinators fauna have been evolved

through specific ecological selection by each species of plants. In agriculture,

enhancement of crop production has been a major accent by mobilizing such inputs

that have low cost and availability in nature. By manipulating agro-ecologies and

agr·o-techniques service of pollinator fauna cou ld be enhanced to reap higher

harvest.

While various domesticated honeybees have been exploited in this regard,

attention has been focused in this All India Coordinated Research Project to study

non -Apis pollinators too for the optimization of potential pollination of target crop

species. Efforts are therefore needed to explore, manage, conserve and

inundate/ inoculate wherever required the alternative pollinators. The continuing

trend toward large scale monoculture, increased pesticide use, and the concomitant

loss of ideal natural habitat cause decline in the number and diversity of crop

poll inators. Wild and domesticated non-Apis bees can effectively complement

honey bee pollination in many crops.

The technical bulletin on Non-Apis pollinators contains the comprehensive

account of for rearing technologies and their uti lization for increasing crop

productivity. The effmis put forih in preparation of this bulletin by authors and

Project coordinator Dr. R. K Thakur (AICRP HB & P) in bringing out this timely

publication deserves appreciation.

~ (T. P. Rajendran)

~ - -- - --- ------'

Preface · ·

Pollinators and pollination are crucial in the functioning of almost all terrestrial ecosystems including those dominated by agriculture because they are in the front line of sustainable productivity through plant reproduction. Productivity of many crops benefits fro m the

presence of pollinating insects, so a decline in pollinator abundance should compromise global agricultural production. Approximately 70% of the tropical crop species depend on pollinators for optimum yields. The economic value of such pollinated crops to India is $726 million and India is the world's second largest vegetable producer. Pollinator decline will have serious socio­economic consequences for countries like India, which host a large population of small and marginal farms for whom falling yield level would be critical for subsistence. Pollinating insects are in decline worldwide resulting in pollination crisis, for (food) crops as well as wild plants and loss of natural biodiversity. Possible drivers for the decline of insect pollinators include habitat loss, intensive land use, globalization and introductions of foreign species, pollution including pesticides, worldwide presence of the invas ive parasitic mite Varroa destructor, introduction and spread of other (new) parasites, loss of the honeybee's genetic d iversity, and detrimental beekeeping practices and climate change. This demands a response from land managers, conservationists and political decision makers to the impending 'global pollinator crisis'. Understanding the causes of pollination failure in plants can aid the successful conservation and recovery of rare plants, maintenance of crop yields, and sustainable use of wild plant resources such as forest timber. Feasible conservation strategies involve making efforts to protect or restore plant resources and native pollinators, and the creation of new protected natural areas, which ensures food provision, mating and nesting sites for poll inators.

Most plants use animals to move their pollen from the male to the female parts of the flower. In the wild, seed production is often pollination-limited, suggesting that pollinators can strongly affect plant fitness. Within the agricultural context, artificial selection for ease of culture has only partially reduced plants' dependence on pollinators. Pollination by animals, primarily bees, remains an essential step in the production of many crops, including melons, squash, apples, berries, and almonds. The main insect group involved in managed pollination are the bees, and in patticular the honey bee (genus Apis). The European honey bee is, by far, the most widespread domesticated bee. Despite their impmtance as honey producers, the role as a pollinator remains the most important economic contribution, outweighing the importance of all the other hive products together. Honey bees can be easily kept, and are capable of pollinating a wide spectrum of plants. It has been estimated that, worldwide, close to 100 crops are pollinated by honey bees. Other bees that are widely used for pollination in agriculture are bumblebees (Bombus spp.), leafcutter bees (Megachile spp.) and mason bees (Osmia spp.). Bumble bees are used, for instance, for tomatoes, eggplants, peppers, melons, raspberries and blackberries. In total, more than 300,000 colonies are repmted to be in use in greenhouses in Europe and North America. Leafcutter bees are used for specific crops such as legumes and, especially, alfalfa, in pa1ticular in the United States .. The solita1y mason bees are effective pollinators at low temperatures(<

l2°C) when honey bees are not yet active, and they are used for early-blooming fruits like apples and pears. In addition to pollination from managed bee populations, the importance of wild insect populations for pollination is becoming increasingly clear.

According to an estimate the world production value for crops used for human food was €1618 trillion, and the total value of the 46 insect pollinated direct crops was €625 billion, that is 39% of the world production value during 2005. The economic value of insect pollination was €153 billion. The rate of vulnerability of the world agricultural production used for human food in the face of total pollinator loss was 9 .5%. Furthermore, some regions are specialized in the production of some pollinator-dependent crop category, the vulnerability of the world production for these categories is much higher than the overall worldwide value. The decline in pollinator population and diversity presents a serious threat to agricultural production and conservation and maintenance of biodiversity in many parts of the country. One indicator of the decline in natural insect pollinators is decreasing crop yields and quality despite necessary agronomic inputs.

Honey bees continue to provide generally satisfactmy and frequently excellent pollination of most of our introduced flowering crops. A few crops exhibiting obviously specialized pollinator needs, such as Lucerne and perhaps red clover, have been catered for by the introduction of specialist bees. Where numbers of these bees have been adequate, crop yields have increased. For agricultural as a whole the diversification of pollinaton assemblages for crops is clearly important. Wild and domesticated non-Apis bees effectively complement honey bee pollination in many crops. In this book an attempt has been made to provide information on the management of various non-Apis bee pollinators.

D PAbrol RKThakur

HD Kaushik Sunita Yadav

Contents .

1.0. Introduction

1.1 Non - Apis bee pollinators-An overview

1.1.1 Short-tongued bees:

l. 1.2 Long-tongued bees:

1.2 Value of wild bees as pollinators

J .3 Diversity of bees in India

1.3.1 Family Stenotritidae Cockerell

1.3 .2 Family Colletidae Lepeletier

1.3.3 Family Andrenidae Latreille

1.3.4 Family Halictidae Thomson

1.3.5 Family Melittidae Schenck

1.3.6 Family Megachilidae Lah·ei lle

1.3.7 Family Apidae Latreille

1.4 Pollination potential ofwild bees

2.0 Carpenter Bees

2. 1 Genus Xylocopa Latreille

2.1.1 Genus Xylocopa Subgenus Biluna Ma, 1938

2. 1.2 GenusXylocopa Subgenus Copoxyla Maa, 1954

2. 1.3 Genus Xylocopa Subgenus Ctenoxylocopa Michener, 1942

2.1.4 Genus Xyfocopa Subgenus Koptortosoma Gribodo, 1894

2.1.5 Genus Xylocopa Subgenus Maaiana Minckley, 1998

2.1.6 Genus Xylocopa Subgenus Nodula Ma, 1938

2.1.7 GenusXylocopa Subgenus Pro:xylocopa Hedicke, 1938

2.1.8 Genus Xylocopa Subgenus Xylocopa Latreille, 1802

2.1.9 Genus Xylocopa Subgenus Zonohirsuta Ma, 1938

2.2 Biology and Life History

2.3 Social Organization

2.4 Foraging Ecology

2.4.1. Abiotic Requirements for Foraging

2.4.2 Water Balance

2.4.3 Nectar Robbing

2.4.4 Food Sources

2.5 Crop Plants Poll inated

2.6 Domestication and Mass Rearing

2.7 Future prospects

1-22

23-48

3.0 Bumble Bees 49-57 3.1 Distribution and diversity

3.2 Effectiveness as pollinator

3.3 Bumble bee foraging activity

3.4 Foraging activity on different crops grown in polyhouse

3.5 Status of pollination and domestication research in India

3.6 Domestication

3.7 Advantages

4.0 Leaf Cutting Bees 58-61 4.1 Nesting biology

4.2 Impact on seed setting

4.3 Conclusion

5.0 Alkali Bees 62-67 5.1 Important Species of Nomia

5.2 Life Cycle and Habits

5.3 Bee Forages and Feeding Characteristics

5.4 Nesting Sites or Beds

5.5 Qualities ofGood Nesting Sites

5.5. 1 Soil Moistme

5.5.2 Soil Composition and Texture

5.5.3 Vegetation

5.6 Establishing the Bees

5.7 Measurement ofBee Density

5.8 Limitations

6.0 Stingless Bees and l\leliponiculture 68-74 5.1 Forage sources for stingless bees

5.2 Pollination of non-crop species

5.3 Advantages of stingless bees over honey bees

5.4 Disadvantages of stingless bees

7.0 Management of Wild Bees 75-87 7.1 Protocols for study of non-Apis solitary bees

7.1.1 Pollinator diversity

7.1.2 Habitat diversity

7.2 Conservation strategies

7.3 Non- Apis bees and future prospects

8.0 Conclusions and Future Strategies 88-89 Suggested Readings 90-91 Acknowledgement 92

1.0 Introduction

I nsect pollination ofag1icultural crops is a critical ecosystem service. Fmit, vegetable or seed production from 87 of the 115 leading global food crops depends upon animal pollination (Klein eta/., 2007). The value of insect pollination for worldwide agricultural production is

estimated at 153 billion, which represents 9.5% of the value of the world agricultural production used for human food in 2005 (Gallai eta/., 2009). The area cultivated with pollinator-dependent crops has increased disproportionately over the last decades, suggesting that the need for pollination services will greatly increase in the near future. Apis mellifera has occupied dominating position in commercial pollination around world because this is highly social bee. But on the other hand wild bees are also valuable pollinators. Their contribution has always been under evaluated perhaps because of our limited insight into their behaviour and mechanism for nesting. The other reason may be that we rely more on the easily manageable honey bees which provide by products also. But today the modem beekeeping suffers from a magnitude of problems, including parasitic mites, various diseases, unability of honey bees to work at low temperah1re and adverse climatic conditions. These difficulties have threatened the honey bee's general utility as an agricultural pollinator (Torchia, 1990) This contributes to the concern to beekeepers, growers of insect-pollinated crops, and policy-makers over recent widespread declines in honey bee populations referred to as colony collapse disorder.

For agriculture as a whole the diversification of pollinaton assemblages for crops is clearly important. Wild and domesticated non-Apis bees effectively complement honey bee pollination in many crops. Examples of management of non-Apis species for agricultural pollination include the use ofbumble bees, primarily for the pollination of greenhouse tomatoes, the solitary bees Nomia and Osmia for the pollination of orchard crops, Megachile for alfalfa pollination, and social stingless bees to pollinate coffee and other crops. The value of the alfalafa leaf cutting bee M. rotundata (F.) as a better pollinator than honey bees for alfalfa has been clearly demonstrated by Richard(1987). He concluded that with the inh·oduction of Megachile bees alfalfa seed yield increased from 50 kg/ha to 350 kg/ha and with more careful handling it can be raised up to 1000 kg/ha.

1.1 Non -Apis Bee Pollinators-an Overview There are about 19,000 described species of bees in the world (Linsley 1958) and, with the exception of one species,Apis mell(fera L., the domestic honey bee, all of them are grouped under the genera I tenn"wild bees." These include:

1.1.1 Short-tongued Bees: Family Andrenidae

Colletidae

Halictidae

Melittidae

Important genera Andrena, Panwginus, Perdita, Pseudopanwginus

Colletes, Hylaeus

Agapostemon, Dufoumea, Halictus, Nomia Hesperapis, Melitta

1

1.1.2 Long-tongued Bees: Anthophoridae

Apidae

Megachilidae

Anthophora, Melissodes, Nomada, Xylocopa

Apis, Bombus. Euglossa, Melipona, Trigona

Anthidium. Lithurgus, Megachile, Osmia

The tenn "wild bee" is used commonly for all bees (solitary bees, bumble bees, carpenter

bees and stingless bees) except honey bees in the genus Apis. Bees generally are distinguished from other flying hymenopterous insects by their characteristic plumose body hairs. Bees are of many sizes, shapes, and colours. Some of the smallest bees, Perdita, are less than 3 tmn; whereas the largest leafcutter bee is over 80 mm. Almost the entire range of colours is found among the brightly marked bees, including many beautiful metallic species. One can easily observe many species ofbees actively visiting flowers for nectar and pollen or engaged in the processes of constructing nests. According to an estimate there are at least 30,000 species of bees in the world. This number of species is more than all the fish, bird, and reptile species combined (Bohart, 1972).

Most bee species construct either single or complex nests underground. Some make earthen, leaf, or resin nests on rocks and plants. Other bees make or utilize crevices in rocks or plant stems, insect borings, and plant galls for their nesting sites. Most bees live a solitary existence-each female after mating locates and builds her nest without the aid of other bees, and usually at a distance from her sister bees. However, some bees are quite gregarious and nest close to one another, sometimes in dense populations of up to a million nests in a few acres of soil. Some bees prefer to nest at the same site year after year, but others relocate their nests each season. A small percentage of wild bees are social or semisocial; that is, there is a division of labor among the bees occupying a single nest (Michener, 2000).

1.2 Value of Wild Bees as Pollinators

2

One cannot easily determine the figure on the value of wild pollinators, simply because total impact on the environment is not known. Studies on the impact of each pollinator species on fruit or seed production of our major crops is almost nonexistent. The reproduction of wild flowering plants is often taken for granted to aid in maintaining soil moisture and fertility, and to provide food not only for wild life but for our domestic livestock as well. How many billions of dollars are these benefits worth? It is easy to document the value of crop species visited by bees, but here again the importance of wi ld bees as crop pollinators has been neglected. It has long been the general consensus that honey bees adequately pollinate crops and there is little need for wild bees. Unfortunately, it is not true since adequate research on the economic benefits of wild poll ina tors has not been done. Interestingly, the research completed on the few wild pollinator species has revealed relatively higher returns compared with investment costs.

The dependence on one species for crop pollination sometimes creates problems. ft seems wise to make greater efforts to study, conserve, and try to manage as many species of wild bees as possible. There are several crops that are under pollinated by the honey bees, either

because the bees arc not physically adapted to pollinate them or the crops are not attractive to honey bees. Some of our most important crops, valued at billions of dollars, are in this category. These crops are alfalfa, soybeans, cotton, vegetable seed, and sunflowers, each of which is adapted to specific types of pollinators. Recent research on the uti lization of several species of wild bees as crop pollinators is just beginning to indicate some of their economic benefits, e.g the alfalfa leafcutter bee. The alkali bee was the first wild bee utilized as a crop pollinator in the United States beginning in the early I 950's. Since that time, the alfalfa leafcutter bee and the blue orchard bee have been domesticated as crop pollinators.

Friese ( 1923) estimated that out of 20,000 species of bees (Superfamily; Apoidea), only four species of honeybees (now nine) and 300 species of stingless bees (Family: Meliponinae) live in the permanent perennial colonies. The majority of the bees are solitaty where a female constructs a nest cons isting of one or more brood cells stocked with nectar or pollen that prov ide food for the larvae that will emerge from the eggs she deposits just before the sealing ofthe cel l. In general, two thirds of the bee fauna is comprised of the solitary bees (Michener, l 965; Linsley, l958; Bingham, l 897; Batra, l977). Michener (2000) apprehended 16,325 species of bees, grouped under 425 genera. The taxa found in whole of the world were reorganized under 7 families. Still much needs to be known from different regions of the world about the existence of different species of solitaty bees, more particularly, from the Oriental region.

Unl ike other commonly known insects, excluding honey bees that belong to genus Apis, bees have least attracted the attention oflndian taxonomists and biodiversity workers. No doubt various aspects on honey bees (Apis) such as their domestication, management and crop pollination have been considerably explored in India. However, this is not true for non-Apis bees. Most of them forage on wild flowerings located in the forests, buffer zones or more often that grow as weeds all along the cultivated crops. The wi ld flowerings infact constitute the primary resource for nectar and pollen for most of bees. Non-Apis solita1y bees also visit cultivated crops in good populations but, as an alternate to wild flowerings.

1.3 Diversity of Bees In India In North India, Batra (1977) recorded 89 species of solitaty bees out of the 97 species studied. The bees belonging to fam ily Megachilidae and Anthophoridae are most conunonly distributed throughout India. Since India is a vast subcontinent with marked topographical and climatic differences, the climatic and floristic conditions vary from tropical and subtropical to subtemperate and temperate conditions and this is the reason that bee fauna from one region differs from the other. Most bees are distributed from valleys through plains to seashores. The Indian species of Bombus is generally restricted to higher elevations especially in the Himalayan ranges. Bingham (1897) recorded 24 species of Bumble bees from higher elevations of Kashmir; Himachal through Sikkim and Assam. Mani ( 1962) reported four species ofbumble bees at elevations of over 4000 m in Himalaya. William ( 1991) has recorded 28 species ofbumble bees from Kashmir including areas under illegal control of Pakistan. Xylocopa species and Pithitis smaragdula in the nmih western states of India remained confmed to 914 m (Kapil and Dhaliwal, 1968) and

391 m (Kapil and Kumar 1969; Kapil et al., 1971) above the mean sea level. The other species are generally abundant in warmer, semi-arid areas, yet distributed in temperate and mountain hill ranges also. The number of species repmted in each genera holds the following numerical order: Halictus 52, Nomia 25,Bombus 28, Xylocopa 19, Megachile 44,and others are still less (Bingham, 1897; Batra, 1977).

The data in table 1 summarizes the occurrence of various bee fauna within the modern territorial limits of our country. Around 92% of known species were recorded from nmthern (Jammu and Kashmir, Punjab, Uttaranchal, Uttar Pradesh, Himachal Pradesh, Haryana) or westem part (Rajasthan, except extreme north and eastern green parts and Gujrat) and remaining 8% were described from rest of the Indian regions. In other words, maximum investigations on biosystematics and floral relationships of bees have been made from the northern territmies and a huge area of southern peninsula is still unexplored. The calculation concerning diversity of bees has revealed that a total of 633 species grouped under 60 genera are found in our country. Certainly, this is not a satisfying number for a huge area with enough of climatological variations, such as India besides, synonymies for many species is still pending to be worked out. Gupta(2003) in an overview of the bee fauna in India stated that the cmTent knowledge shall form a base for the Melittologists to fill the gaps in future. The recent most biosystematic information given below in Table 2 shall be further useful to the biodiversity, ecological and pollination-workers of the country.

Table 1. Diversity of bees in India (Gupta, 2003)

S.No Family Sub-Family Tribe Genus No. of

Broad Distribution in India Species

Colletidae Colletinae - 1. Co/letes Latreille, 005 Northern region 1. Lepeletier Lepeletier 1802

Hylaeinae - 2. Hylaeus Fabricius, 014 13 in Northern region and 2 in

Viereck 1793 Southern region

2. Andrenidae Andrenidae - 1. Andrena Fabricius, 034 Widely distributed in northern Latreille Latreille 1775 region

3. Halictidae Rophitinae - 1. Systropha llliger,

002 Punjab and northern Rajasthan Thomson Schenck 1806

Nomiinae - 2. Nomia Latreille, 067 Widely distributed throughout

Robertson 1804 India

- 3. Pseudapis Kirby, 002 Northern region 1900

4. Steganomus 003 Northern and western Ghat -

Ritsema, 1873 regions

Nomioidinae - 5. Ceylalictus Strand, Discontinuous: extreme Borner 1913 003 northern, coastal western and

south-central region

6. Nomioides Schenck Deserts of north-western region - 1867 003

4

S.No Family Sub-Family Tribe Genus No. of

Broad Distribution in India Species

Halictinae 7. Halictus Latreille, 47 in hilly region of north Thomson 1804 055 extending upto north east and 8 in

west, central and western ghats

8. Homalictus 006 North and north eastern region Cockerell, 1919

9. Sphecodes 11 in north and north eastern Latreille, 1804 017 region and 6 in south

Rajasthan and Gujarat

10. Thrinchosto-ma 002 Northeastern region Saussure, 1890

4. Melittidae Melittidae Melittini 1. Melitta Kirby, 1802 001 North and north eastern region Schenck Schenck

Fideliinae Paraho· 1. Pararhophites 002

1 in northwestern region and 1 in 5.

Cockerell phitini Friese, 1898 Gujrat

Discontinuous distribution: Lithur- 2. Lithurgus Berthold,

014 northern, western coastal, gini 1827 extreme southern and

northeastern region

Osmiini 3. Chelostoma Sikkim and allied hilly territories

Latreille, 1809 001 of north east, continue upto Myanmar

17 in Himachal Pradesh, 4. Heriades Spinola,

022 Uttaranchal and north-eastern 1808 hills and 5 in southern Rajasthan

andGujarat

5. Hoplitis Klug, 1807 010 8 in Northwestern region and 2 in western coastal region

6. Noteriades, 004 Himachal Pradesh

Cockerell, 1931

7. Osmia Panzer, 1806 4 in Himachal Pradesh , 005 Uttaranchal and 1 in eastern

coast of southern region

8. Protosmia Ducke, 001 Northern (medium hilly) region

1900

9. Pseudoheriades 005

Northwestern region (and Peters, 1970 eastern buffer zone of Thar)

S.No Family Sub-Family Tribe Genus

10. Wainia Tkalcu, 1980

Anthi- 11 . Acedanthi-dium Michener and

diini Griswold, 2000

12. Anthidiellum Cockerell, 1904

13. Anthidium Fabricius, 1804

14. Dianthidium Cockerell, 1900

15. Eoanthidium (Popov, 1950)

16. Euaspis Gerstaecker, 1857

17. Jcteranthidi-um Michener, 1948

18./ndanthidium Michener and Griswold, 1994

19. Pachyanthi-dium Friese, 1905

20. Trachusa Panzer, 1804

21. Trachusoides Michener and Griswold, 1994

22. Aglaoapis Cameron, 1901

Mega- 23. Coelioxys Latreille,

chilini 1809

24. Megachile Latreille, 1802

6. Apidae Xylocopinae Xyloco- 1. Xy/ocopa Latreille, Latreille Latreille pini 1802

Alloda- 2. Braunsapis pini Michener, 1969

6

No. of Species

002

001

001

004

004

002

002

002

001

003

004

001

001

032

105

036

006

Broad Distribution in India

So far recorded from Lonavala [Maharashtra only

Northern (Himachal Pradesh and Uttaranchal only)

Northeastern region

Northern and northwestern hilly and xeric regions

Northwestern xeric region

Northwestern xeric region

1 in northeastern region and 1 widely distributed in almost whole of India except, extreme southern region

Northwestern xeric area

So far recorded from Pune

Discountinuous: Kangra, Bengal, Kerala, Bombay and Panchmarhi

Discontinuous: northwestern region, Central region, Bombay

So far recorded from Appangala (Karnataka)

So far recorded from Bombay

Widely distributed all over country

Widely distributed all over country

Widely distributed all over country

Discontinuous: northern, parts of western and extreme southern region

]

l

Family Sub-Family Tribe Genus No. of

Broad Distribution in India S.No Species

Cerat- 3. Ceratina Latreille, 014 Almost throughout India inini 1802

Nomadinae Noma- 4. Nomada Scopoli, 013 Mostly confined to northern

Latreille dini 1770 regions

Epeo- 5. Epeolus Latreille, 003 2 in north and northeastern

lini 1802 region and 1 in Central region

Ammo- 6. Parammobato-des Distinct territory not known [label

batini Popov, 1931 001 indicates only India, perhaps extreme northern India]

7. Pasites Jurine, 1807 001 Southern region

Apinae Ancy- 8. Tarsalia Morawitz, 001 Western Ghats

Latreille lini 1895

Euce- 9. Eucera Scopoli, 003 Northwestern region rini 1770

10. Tetralonia Spinola, 015

13 in Northern region and 2 in 1839 Central region

11. Tetraloniella 002

1 in extremity of northeastern Ashmead, 1899 region and 1 in central region

An tho- 12. Amegilla Friese, 021

Widely distributed all over phorini 1897 country

13. Anthophora 010

Widely distributed all over Latreille, 1803 country

14. Elaphropoda 004 Mountains of extreme northeast

Lieftinck, 1966 region

15. Habropoda Smith, 014 Confined to northern to

1854 northeastern region

Melee- 16. Melecta Latreille, 002 Extreme northern region tini 1802

17. Tetralonioi-della Southern slopes of Himalaya

003 through northeastern region Strand, 1914 upto Indonesia

18. Thyreus Panzer, 006

Widely distributed all over 1806 country

19. Bombus Latreille, Exclusively along the high and Bambini

1802 026 medium range of mountains in Himalaya

Melipo- 20. Lisotrigona Moure, 001 Central region

nini 1961

21 . Trigona Jurine, 003

Arid zone in western region and 1807 Northeastern up to Indonesia

7

S.No Family Sub-Family Tribe Genus No. of

Broad Distribution in India Species

22. Apis Linnaeus, 2 Widely distributed, 1

Apini 005 introduced and 1 confined to 1758 very high altitudes in Himalaya

006 012 060 633

* Table contains the taxonomic categories found and known from India.

A detailed historical account, regarding discovery of different genera and species of non-Apis bees from south Asian counh·ies up to Indonesia has earlier been published by Gupta and co­workers during 2003.

According to him, the country can be subdivided under certain specific area based upon homogeneousness of ecological factors faci I i tati ng the presentation. The regions specified are:

a) Extreme northern region: It includes high altitude of Himalayas ranging between 6-7,000 feet and more in Jammu and Kashmir, Himachal Pradesh and Uttaranchal.

b) Northern region: This indicates the medium and low altitudes area and, plains ofTarai at and south of Himalayan range i.e. including southern Jammu and Kashmir, Punjab, Haryana, eastern Rajasthan adjacent to Uttar Pradesh, Uttar Pradesh and plains ofBihar.

c) Western region: Almost whole of Rajasthan (excluding eastern hilly-green region) and Gujrat.

d) Western Ghats: Western hilly Maharashtra, Goa, western Karnataka and, otherwise specified Lakshadweep and Mini coy Islands.

e) Central region: W11ole ofM. P., most of the Maharashtra, Chhattisgarh, Andhra Pradesh, excluding coastal area.

f) Northeastern region: T ncludes hills of eastem Himalayan range, all the seven states of the northeast and Dish·icts like Darjecling and Sihguri in W. Bengal.

g) Eastern region: Excluding hilly north, West Bengal, hilly South Bihar, Orissa and extreme coastal area ofAndhra Pradesh.

h) South-central: The region include Karnataka, and JiOiihern parts ofTamilnadu.

i) Extreme southern region: Specified for Pondicherry, coastal Tamilnadu and Kerala and, otherwise not specifiedAndaman and Nicobar Islands.

1.3.1 Family Stenotritidae Cockerell

8

The smallest fami ly of bees that consists of only two genera namely, Ctenocolletes Cockerell with ten species and, Stenotritus Smith with eleven species. Both are exclusively found in Australia and, so far no species has been recorded from anywhere else in the world.

l

l I 1

1.3.2 Family Colletidae Lepeletier

This is a medium s ize family subdivided into 5 subfamilies namely, Colletinac Lepeletier, Diphaglossinae Vachal, Xeromelissinae Cockerell, Hylaeinac Viereck and Elllyglossinae Michener. Among them, subfamily Colletinae and Hylaeinae are the lone representatives

of this family, known by one genus each, in India. The moderately populated genus Colletes, represented by 5 species, is restricted to northern region so as other identically populated genus Hylaeus with 15 species. It was interesting to note that 3 species of Co/fetes arc more or less confined to middle range height (around 5000 to 6000') in southern slopes of greater Himalaya in Uttaranchal and Sikkim. All of them are quite limited in their distribution. On the contrary, species found in plains and those reaching upto Rajasthan in south, are comparatively widely distributed. The area of species distribution may be apprehended as Jammu and Kashmir, Punjab, Haryana, Himachal

Pradesh, Uttaranchal, eastem Rajasthan and one species record exists from Sikkim and Pun e.

Probably, majori ty of species of this genus are also distributed in intermediate area namely, union territory of Delhi , northem Uttar Pradesh as well as the green northern

Rajasthan. Practically there exists no ecological barrier between the noted tenitmy and the probable area of dish·ibution. These areas require intensive surveys. Species of Colletes

have been collected digging deep bunows on sloppy surfaces of hard grounds in Sikkim. Grewal, eta!. (1970a) described the life histmy of C. nurse; fiom Punjab. They also collected many females in Punjab while busy excavating burrows in sandy, bare and dry crop fields. Batra ( 1977) made many ecological observations on nesting sites of Colletes. Batra (1968) and Gupta and Yadav (200 1) noted down different floral species visited by some species of this genus.

Genus Hylaeus in major contains those species which were earlier grouped under genus Prosopis . The name Hylaeus has priority thus accepted. Prosopis however, still is a subgenus ofreorganizedHy/aeus (Michener, 2000: p. 182). It is a worldwide genus having representatives on all continents. Most of the species collected from oriental region are yet

to be assigned a subgeneric categmy. Species of our destined area have been collected from Kashmir, Punjab, Rajasthan, northem pa11 ofW. Bengal and Sikkim. Two out of 15

species known from our country were collected from southern extremities probably reaching up to Sri Lanka (Wijesekara, 2001 ). An intensive survey to the nation waits so that exact picture of its distribution would be marked on the map of our country. Species of this genus are known to visit some culti vated crops in Punjab (Batra, 1968).

1.3.3 Family'Andrenidae Latreille

It is a large fam ily of bees subdivided under four subfamilies namely, Alocandreninae Michener, Andrcninae Latreille, Panurginae Leach and Oxaeinae Ashmead. However, in

g

our country this family is represented by subfamily Andreninae only having the single representative genus Andren a . Species of this genus can be marked all over northern India, more particularly plains of Punjab, Haryana, Himachal Pradesh, Rajasthan, Uttaranchal, Uttar Pradesh, Bihar and W. Bengal. Two species were collected one each from western coast of Maharashtra, the central Maharashtra and Assam. 34 species are known so far after judging synonymies. Major bulle of them show affinities for their western allies found in Pakistan, Afghanistan and, up to Turkey. On the contrary, eastern species are fairly distributed upto Myamnar and Malaysia. Precisely, most of species seem to have adapted for their specific eco-geographical environments in India but many more have their affiliates towards both neighbouring territories. An intensive survey of the remaining part ofthe counhy would yield several new species and their floral data, unknown so far.

Rahman (1940) briefly described a nest of Anclrena if erda, collected near a toria field. This is not yet known that species of Andren a excavate the tunnels themselves or they occupy pre-existing subtetTanean butTows. However, the highly branched, 50-60 em. deep burrows have at least one cell at the end. Wain ( 1968) also recorded nests ofA. bellidoides

in the hills of western ghats. They were constructed gregariously. Grewal et al ( 1970b) published the nesting behaviour of Andren a lea en a.

It has been observed that species of this genus bear a good affection for the flowerings of cruciferous crops. The oil seed producing region all along the Shivalik andAravali range harbours comparatively good population of many species. Mohammad (1935), Rahman (1940), Batra (1968), Atwal (1970), Bhalla eta!. ( 1983), Kumar et al (1988), Kumar et al.

(1994) and, Gupta and Yadav (200 l) have listed many species of Andren a collected on different cruciferous crops in the mentioned area. Abrol (1986, 1988, 1989) detailed the behavioural aspect of some Andren a species from Jammu and Kashmir.

1.3.4 Family Halictidae Thomson

10

This is a considerably large family having many semi social bee species, common I y known as sweat bees. It is further subdivided into four subfamilies namely, Rophitinae Schenk, Nomiinae Robertson, Nomioidinae Bomer and, Hahctinae Thomson. The group is represented by 160 species in our country. Subfamily Rophitinae and Nomioidinae consist of a limited number of species with a restricted distribution whereas, subfamily Nomiinae and Halictinae are widely distributed.

The single representative ofRophitinae, genus Systropha was reported with two species limited to western Punjab and nmthern Rajasthan. Batra and Michener (1966) described one of them alongwith its subsoil fom1ed nest and the larva. Identically, two representatives of Nomiinae, genus Pseudapis and Steganomus are confined to certain pocket area of the cOtmtry. The third representative, genus Nomia with 67 species, is widely distributed all over India. Subfamily Nomioidinae is known by genus Ceylalictus

and Nomioides both with three species each. Among them first one exhibits discontinuous distribution and the second is found in the arid region of north-west.

Subfamily Halictinae is ft.uther subdivided into tribe Augochlorini and Halictini. Among them Halictini has representation in our country and, of fairly wide occurrence. Two species of genus Thrinchostoma arc known exclusively from the northeastern region of the country. In major its species were recorded from the eastem countries next to India. Genus Hom a/ictus, found in similar territory, is comparatively widespread, reaching upto the north at the southem slopes of H imalayan all along the range from north to northeast.

Genus Sphecodes is also represented in the area inhabited by Homalictus but in addition to its eleven species, six are found in the southern hilly zone ofRajasthan and at the tail end of Araval i hi lis in Gu jrat. The species found in western zone have affinities with their western allies. Genus Halictus, represented with 55 species, is largely confmed to the northern tenitories and at low level hills of central and western patts of the country. The aspect of taxonomy of the fam ily needs intensive investigations for most of the area of our country besides, the bionomics and biology.

Works pertaining to the nesting biology of a few species of Lasioglossum, Nomioides, Nomia and Halictus appeared so far, were published during 1960s to 1980s. Among them significant contributions regarding social behaviour and nests of nomiine bees, specially observations made for several species of Lasioglossum and Halictus, were published by

Batra (1964, 1965, 1966, 1968, 1970, 1971 , 1995, 1997 etc.), Kukuk (1980), Batra and Bohart ( 1970). Same aspect was described for many other halictid bees of Indo-Malaysia

regions by Sakagami, Ebmer and Tadauchi (1996) and, Sakagami and Ebmer ( 1987). Most of the halictids nest gregariously in the moist subterranean soil (resembling Co/fetes). However, their burrows were comparatively shallower culminating into 5-6 collateral branches. Each of which further bear around 1-3 or even 9 cells at the end. Precisely, a nest is a cooperative effort of as many as 20-25 females, suggesting their gregarious or semisocial behaviour. Nest tmmels were lined by mandibular secretions that provided evidence that they were self-excavated by females (Rahman, 1940; Sakagami and Michener, 1962, 1963; Sakagami and Wain, 1966).

A comparative account of the properties of nest-building secretions of Nomia, Anthophora, Hylaeus and other bees was presented by Batra (1972). Batra ( 1978b) further presented aggression, territoriality, mating and nest aggregation of some solitaty bees that belong to Halictidae, Megachil idae, Colletidae and the earlier known fam ily Anthophoridae.

Kumar et al (1994) compared the pollination efficiency of bees by making comparison in the rate of visits made on to ria in Himachal Pradesh. They identified Halictus catullus and Hal ictus splendidulus alongwith some other bees, on this crop. The regional studies made with regard to identification of species of various genera pollinating some selected crops,

11

were published by Batra (J 968) and Gupta and Yadav (2001).

1.3.5 Family Melittidae Schenck

This fifth family ofbees is of rare occunence in India. Michener (2000) further classified it into three Subfami lies, Dasypodainae Borner, Meganomiinae Michener and Melittinac Schenck. Among them Melittinae, including tribe Melittini , with single Genus Melitta, known with single species [i\1. harrietae (Bingham, 1897)] reaches India. It is found in a very scanty population in the extreme north and notih eastern hilly regions at the middle range altih1de in Himalaya (Gupta, 2003 ). It has many allies in the hilly and atid countries,

west and east to India. The family has numerous representatives in Africa and countries in the middle east. Probably, high range ofHimalaya imposed a restrictive entry of erstwhile Palaearctic species into our country. Comments on nesting of a species of Ctenoplectra were recorded by Bingham ( 1897) from Tenasserim. This genus is now placed in tribe Ctenoplectrini, under subfamily Apinae of Apidae. So far nothing is known about the nesting behaviour and pollination aspects etc about J'vfelitta in India. Malyshev ( 1923) described them for some species known from Russian region.

1.3.6 Family Megachilidae Latreille

12

Until now it exists as the largest family of bees with regard to number of taxa known from

India. Gupta ( 1993) consolidated 161 species of bees included in this fami ly found in six states in nmihwest. It detailed taxonomy alongwith their flower records. Still work pertaining to synonymy for several taxa of the country, is pending besides, the multi­seasonal expeditions of the southern peninsular region. As a whole, 24 genera with 229 species are included in this family from India. The taxonomic categories of this famny have undergone a fair amount of shuffling by virtue of cladistic analysis made by Roig­Alsina and Michener (1993). This resulted into recognition of two subfamilies namely, Fideliinae and Megachilinae (Michener, 2000).

Subfamily Fideliinae is represented by a single tribe Pararhophitini in India, including genus Pararhophites with two species. The small bees of this genus love xeric conditions

and were largely la1own in the western area beyond Indian limits. Its two species were collected from area adjacent to Baluchistan [Ferozpur, at the western border of Punjab, reaching upto Rajasthan] and second was found in Junagarh in Gujrat. These bees were collected while pollinating flowers of Convolvulus at both vicinities. The nests of one species were found in good aggregation made in subterraneous bunows in Baluchistan (McGinley and Rozen, 1987).

Subfamily Megachilinae includes five tribes and all have good representation in our country. The first, Lithurgini represented by genus Lithwgus Be1thold (14 species), has discontinuous distribution. Species like L. atratus and L. dentipes have been of great concern with regard to their pollination of Cotton (Malvaceae) and, earlier many have

conunented upon their nests build in hollow sticks (Horne, 1870, 1872; Malyshev, 1930; Lieftinck, 1939).

Second tribe Osmiini includes numerous genera but most of them are known from the colder regions in the north and atid northwest. Genus Chelostoma, represented by a single

species is an exception. It was initially recorded from Sikkim and is found up to Myanmar. On the contrary, Genus Heriades (22 species) is widespread in Himachal Pradesh, adjacent Punjab and extends southward all along Aravali hills upto Gujrat, in a scattered pattern. These small black bees have great affection for Compositae flowers (Gupta and Yadav, 2001). Hoplitis (10 species) also has more or less similar distribution pattern however, its two species were recorded at northern tetTitmy of the western coast. Noteriades (four species) and Protosmia (one species) are restricted to Himalayan range. Fonner is found at medium altitudes in Himachal Pradesh and the later is exclusively confined to high mountains in the extreme north.

After making taxonomic revisions, genus Osmia is reduced to five species found in India. Most of them are restricted to Himachal Pradesh and Uttaranchal at medium range mountains. Its one species was exclusively recorded from Karaikal located at south­eastern coast. Genus Pseucloheriades is quite widely distributed in Rajasthan and coastal Gujrat. The nests and immatures of one of its species were described from 'Moonj straws' (Gupta and Sharma, 1995: misident. as of Heriacles) . Recently many more bees were

collected from western Rajasthan and Gujrat by this author [yet to be named]. More particularly they were collected while busy pollinating Coconut at Somnath, Veeraval and

Diu in Gujrat. Genus Wainia was exceptionally recorded from Lonavla in Maharashtra with two species.

Tribe Anthidiini has fair representatives in our country but mostly with a discontinuous distribution. Genus Aceclanthidium, Anthicliellum, Eoanthiclium, Jcteranthidium, lndanthiclium, Trachusoicles are known with merely one or two species. Euaspis (=Parevaspis), the cleptoparasite, is almost cosmopolitan throughout the counhy. Its one species Euaspis abdomina/is is found in the northeastern states extending upto Myanmar and Malaysia. The second, Euaspis carbonaria is found almost everywhere except the

extreme southern region of the country. Both species are well known cleptoparasites of MegachiLe. Species of Pachyanthidium, Trachusa and Dianthiclium are found in different pocket areas at different parts of the nation. However, most of the species are of prominent occurrence in arid zones of the northwest. A thorough expedition would yield many more species of this tribe.

Tribe Dioxyini, represented by a good number of taxa in the countries beyond the western limits of India, is known by a single genus Agiaoapis here. Its single species, Aglaoapis

brevipennis was collected from Bombay around a century ago and until now not found anywhere else.

13

TribeMegachilini, now consist of only two genera namely, lvfegachile that include the leaf

cutting and mason bees and, its well known cleptoparasitic genus Coelio.xys. Both arc

found almost everywhere all over the country (Gupta, 2003 ). Species of genus Megachile

(exceeding 130 in number), have attracted many authors with regard to their nesting and

pollination aspects.

The comments on nests of various leaf cutting bees were initially published by Horne

(1868, 1870 and 1872), Maxwell-Lefroy and Howlett (1909), Bingham (1897, 1 R98a,

1898b, 1908) and Dutt (1912). Unti l then, the interesting act of cutting leaf pieces with its

mandibles, followed by their transportation to the nesting s ite by a female megachi line bee

attracted enoug h attention all over the world. They use leaf cuttings, petals, mud, soil,

pebbles and plant resins etc. for the construction of their nest chambers in self excavated or

existing horizontal buiTows in muddy walls or in subterranean soil. The nesting biology

for a few Indian species was described by Pagden ( 1934 ), Malysbev ( 1930), Pie! ( 1930)

and, Wain ( 1968). Later Kapil, Grewal and Atwal ( 1970), Kapil et al ( 1975), Chaudhary

and Jain (1978), Gupta and Sharma (1995) and, Gupta, Naval and Charan (2003b)

included nesting behaviour, immaturcs and correlated aspects in their studies.

The family includes several good pollinators of Leguminosae, Compositae, Solanaceae

and, several fruit crops etc. Ralunan ( 1940) could identify very few species ofMegachilini

on Sarson and Tori a. Batra ( 1968) identified several megachilids pollinating many

cultivated and wild crops in Ludhiana. Mishra et al (1976) and Batra (1976a and 1979a)

presented some more works related to pollination of some cultivated crops from Himachal

Pradesh and Punjab, respectively. Abrol ( 1986a, 1988b, 1988c) notedMegachile ncma and

Megachile .flm·ipes (sic) on alfal fa and recorded their ceo-pollination behavioural

relationships. Kumar et al ( 1993) recorded around 9% of bees belonging to Megachi le and

28% belonging to Ceratina, on Cichorium intybus in Himachal Pradesh. Gupta ( 1993)

presented the complete flower record ofmegachilid bees collected from six states in the

northwest. Gupta and Yadav (200 1) recorded a good number of megachil ids from four

cultivated crops in eastern Rajasthan and adjacent Uttar Pradesh.

Some of the good and effective megachilid species recorded by them include: Megachile

bico/01; M Ia nata, M cephalotes, M. gathela, M alb((i·ons and M. ere usa for almost whole

of the northern half of the country.

1.3. 7 Family Apidae Latt·eille

14

Based upon the c ladistic analysis of Roig-Alsina and Michener (1993), the reorganized

Apidae consists of subfamilies Xylocopinae, Nomadinae and Apinae (Michener, 2000).

Table 1 lists the three named subfamilies, their 12 tribes and corresponding 22 genera

found in Ind ia. So far 192 species have been grouped under different generic categories.

Still works re lated to synonymies of several species is pending and many of them would

face taxonomic shuffling or would be synonymized. Species of Xylocopa (36 species and

subspecies) [just concluded taxonomic revision, and is being published separately in this volume of the book .. Author], Cera/ina (14 species),Amegilla (21 species), Anthophora ( 10 species) and, Thyreus (6 cleptoparasitic species), arc widely distributed.

On the contraty, Braunsapis (six species, having discontinuous distribution), Nomad a (13 species, mostly confined to no1ihern region, upto the area of westem arid Rajasthan), Tetralonia (15 species, out of them only two were recorded from M.P. and adjacent Maharashtra and rest are confined to north India), Hahropoda (14 species, all restricted to Himalayan range) and, Bombus (restricted around or above the altitude of 3000 feet in Himalaya), are region specific.

A few genera namely, Epeolus (3 species), Parammobatodes (one species with doubtful occun·encc in India), Pasites (one species), Eucera (3 species found in nmthwest), Tetraloniel/a (two uncommon species), Elphropoda (four species) andMelecta (with two species in extreme north, in Himalaya), Tetrafonioidella (3 species all found throughout

the Himalayan range up to east), Lisotrigona (one species known from M.P.) and, Tarsalia (one species from Poona) come under the category of lesser known genera in the country.

The social stingless bees belonging to genus Trigona (three species on record) were originally col lected from all over the Himalayan range however, during past two decades,

many more have been collected from several additional territories, by this author. They were Agra, Allahabad, Jamalpur (Bihar), Jabalpur and, Pachmarhi [M.P.] and, recently during the summers of2003, more specimens ofthis genus were collected at Dwarka and Somnath, i.e. at the costal Gujrat and, more as a surprise during September 2003, many more specimens of this genus have been collected on the flowers of Techtona stems (Bignonaceae) in Jodhpur.

Genus Apis, the honey bee, has five representative species in our country. Among them Apis mel/!fera the Italian honey bee, was introduced in India during sixties in the previous centt.uy. lt ultimately 'merged' with A pis indica and now 'becomes' a cosmopolitan species by virtt.1e of its adoption in artificial domestication programme by ICAR stations all over India. Engel (2000) bas recently published the information conceming honey bees of

India. Apis dorsal a (with three subspecies A. d. dorsa/a, A. d. laboriosa and A. d. bighami), Apis cerana, Apisflorea andApis andren(formis are the other species. Apis andren?formis and Apis dorsa/a binghami are known with limited distribution records. The first of these was recorded from Khasia hills and second was collected from Sikkim and Meghalaya. Apis dorsata laboriosa, the giant Himalayan honey bee is confined to the high altitudes in

northern region. Roubik et al ( 1985) noted distribution and nesting of A. d. laboriosa. Batra (1996) described its biology and declared it as a good pollinator of apple at high

altitudes in Garhwal (Uttaranchal). Qayyum andAlunad ( 1967) described the biology of A

dorsata. Thakar and Topani (1961, 1962) detailed the nesting behaviour of A. dorsata and

1!J

16

A. jl01·ea. Sham1a and Thakur ( 1999) put forward the morphometric characterization of A. dorsa fa and, Sharma et al (2000) detailed the diurnal activity of A. cerana andA. mell(fera on different flora.

Since early times the carpenter bees that belong to genus Xylocopa have been of great interest to mankind therefore, enough observations on nestings and immatures were made

on various species of this genus. Horne ( 1872), Perkins ( 1899), Max:well-Lefroy and Howlett (1909), Dover (1924), Dutt (1912), Kannan ( 1925), Iwata (1964), Ma (1938 and 1954) and, Beeson (1938) published comments on their nests and nesting behaviour. Sakagami and Yoshikawa (1961), Kapil and Dhaliwal ( 1968a, 1968b, 1969), Bhaskar and Gopinath ( 1975) further recorded their biology from woods, trees, logs and bamboos in northern India.

Green ( 1899) recorded interesting sleeping habits of Crocisa ramose. Malyshev ( 1925, 1936) described the nesting habits of several Anthophora and other solitary bee species. Schwarz ( 1939), detailed social species of Trigona found in Indo-Malayan region. Sakagami ( 1960, 1966) noted Ethobiological aspect of several Allodape and other genera of Apidae. Lieftinck (1955, 1957, 1962, 1964, 1972, and 1974) made remarks on

ethobiology of many species that belong to former Anthophoridae such as Crocisa. Xylocopa, Thyreus, Melecta, and Habropoda etc. Iwata ( 1964) referred the egg of Xylocopa as the largest among all insects. Kapil and Kumar (1969), Batra (1972, 1977, 1980), Pagden (1957), Batra, Sakagami and Maeta (1993), Reyes and Sakagami (1990), Reyes and Michener ( 1990), Batra and Norden (1996) and, Williams (1991, 1994, 1998) are the few references, dealt with nesting behaviour and immatures of some Indian species of Ceratina, Anthophora, Amegilla, Braunsapis, Bombus and Trigona. Identically, thousands of references regarding pollination are available for this family ofbees, from all over the world. However, when referring to Indian works, some of the important ones may be listed as follows:Howard et al (1920) first of all refened a number of pollinators, including honey bees, for several Indian crops. Work ofMohammad (1935) concerning

pollinators oftoria and sarson has been referred above. Similarly, role of Apis indica was studied by Latif et al (1960) on toria and sarson. Several notes on the pollinators of leguminous crops including alfalfa were put forward by Batra ( 1977, 1991, 1994, 1995). Sandhu et al ( 1976) used Cera tina (Pithitis) smaragdula in India on alfalfa. Rai and Gupta ( 1983) published a useful note on the role of honey bees on the pollination of apple and pear. Abrol ( 1989, 1990, and 1992) described the ecology and behaviour of pollinators of strawberry and apple, including honey bees in major. Shanna and Gupta (1993) listed the flowering plants visited by Apis mellifera and A pis cerana. 1l1ey concluded a total of 119

species of flowering plants in Solan (H.P.) among them 72 were visited regularly. Out of these 41 species provided both nectar and pollen, 22 nectar only and 9 species of plants were visited exclusively for pollen. Goyal and Gupta (1994) presented a detailed account

of beekeeping with Apis mellifera in India. Batra (1995) wrote on the use of solitary bees for blueberry pollination. Recently, Sharma and Gupta (200 1) published the impact of bee pollination on sustainable production in apple orchards. Kumar and Singh (200 1) recorded a preference for competing flora with sunflower for honey bees. Kumar et al (2002) made observations on different modes of honeybee pollination and its effect upon the oil content in seeds of sunflower.

1.4 Pollination Potential of Wild Bees

Inspite of being represented in large numbers, non-Apis bee pollinators have-received little attention. The probable reason seems their unpredictable seasonal availabil ity, lack of knowledge about their biology and host plant relationship. It was only in the mid sixties

that scientific interests were generated to study and understand their life processes in India (Atwal, 1970). Failure of honeybees in the pollination of alfalfa which requires tripping (Kapil et. al , 1977., Kapil and Jain 1979, 1980) dwindling of bee colonies due to acute floral dearth in heavy monsoon during June to September and similar effects during severe winter in Himalayan ranges from November to February and prevalent bee diseases has created the necessity for exploration of altemate yet suitable non-Apis bee pollinators to augment crop yields in India. in addition to alfa lfa, clovers and fruits like apple, pear, peach, almond and several other crops such as sunflower, hybrid tomato, cotton, onion,

carrot and cucurbits (Kapil and Dhaliwal, 1968) can be future potential crops requiring the serv ices of the non-Apis pollinators.

A wide variety ofnon-Apis bee pollinators are associated with variety of crops. The data presented in Table 2 summarises the association between non-Apis bee pollinators and different crops in India. Alfalfa (Medico go sativa) is a perennial herbaceous protein rich legume. It is one of the most important fodder crop throughout the world. It is a cross plant and its flowers being typically papilionaceous has stamina! column held inside keel. Tripping i.e. release of stamina! column is considered to be a prerequisite, of cross

pollination (Free. 1993 ). Most investigators have concluded that long tongued bees are not vety important in alfalfa seed production patiicularly in dry or arid conditions because of

the fact that the honey bee collect nectar from alfaalfa flowers from the side of the keel, thus avoid the mechanical shock of the stamina! column. Most of the workers has emphasized importance of ceriain non-Apis species in the pollination of its flowers. Among a variety of wild bees associated with alfalfa flowers, the investigations based in U.S.A. and Alberta (Canada) showed that leaf cutter bee (Megachila rotundata) and a

alkali bee (Nomia mellandari could be exploited and propagated as alfalfa pollinator (Bohart, 1972).

17

a) Megachile specie,· working on leguminous flO\\ crs b )Xyloc:opa{enestrata on cucurbits

18

A variety of non-Apis bees have been repmted to be associated w ith some other crops in tllis region (Kap il and Jain 1980). They reported megachilid species associated with

alfalfa under local conditions. The studies (S ihag, 1990) have suggested the inclusion of some of these bees M . .flavipas, AI.femomta. A! lanata and !vf. cephalates under the genus Chalcodoma. The major period of their activity for alfalfa pollination in the April-May months. Their broods however undergo dmmancy as mature larvae first in the months of June-July and then in winters. Unti l mid March, there arc many overlapping generations

in the months of September to November. High day temperature and dry condition favour their foraging activity, brood formation viz-a-viz alfalfa pollination. 01 impotiant non­Apis alfalfa pollinators are Mel/ita leporilla. Atlthoph quadrifasciata and few other species ofAI/drella andMegac!Jile.

The leaf cutter bee earlier called as Afegachile pac{/lca, is a fast nest gregarious bee and it nest in any such tunnel available in wood above the ground. It has been found to accept the man made art i licia l nesting devices including tunneled wooden block, corrugated boards or the plastic tubes of appropriate tunnel size i diameter. Its native region is South-West Asia and it spread to eastern USA around 1930. Today, it is successfully managed for alfalfa pollination in many states of USA and southern parts of Canada as well as many of the European countries. It is to, tenned as million dollars bee as it involve the inputs and

outputs of millions of dollars in making various mechanical devices and nesting materials and management techniques for alfal fa seed production.

The alkali bee is another fast nesting gregarious bee. lt however nest in big ali<alinc soil. Once established favourable site may produce as many as 2 lakh cells per acre. The best sites are bare or slightly vegetated tender silty loamy soils. Main adventage in its management is that its nests can't be easily removed, stored or transported. There have

been many refinements in developing artificial nesting sites. The most impotiant step has been to keep the site puffy a little moist by providing plastic films several feet below the

J

surface. Heated calli are sometimes employed to enhance pupation and emergence of adults to coincide with alfalfa blooming. In addition to rains moulded soi l, rats, skunks,

birds and variety of other parasites are its natural enemies. A temperature of30°C is most appropriate for rapid pupation of the diapausing larvae. Pithitus smaragdu/a, a green metallic bee is another important alfalfa pollinator. It is a small carpenter bee and it makes nesting tunnels in cut pithy stems such as common reeds (Eriant/ws mw?ia).

The wi ld bees are associated with many other crops also. The crops like clovers (Ladino clover, red clover, egyptian clover, white clover and sweet clover), soybea pigeon-pea

sunhemp, broad bean, mustard, rape, coffee, papaya, cotton, sunflower safflower and apples are few other important crops which could be benefited for cross pollination. A variety ofhumble bees belonging to the genus Bombus utilized for pollination of clovers. The bumble bees are semisocial bees as these initiate nesting as an individual female, but later establish a large size colony. Osmia coeru/escens, a leaf cutter bee has also be reported to increase seed yields in red clover in USA Osmia comi(fi-ons has been successfully managed for apple pollination in northern and central Japan. It nests in bamboo and hollow reed. The stingless bees commonly termed as 'melipona' bees are another important non-Apis group of true social bees which resembles to Apis species as

these form large colonies and yield sufficient honey for human consumption. The indigenous people of tropical America and Africa have for centuries managed various species of stingless bees for honey. Continuously changing climatic conditions and over exploitations of forests and batTen lands for agriculture have been the major hurdles in natural propagation of wild bees. In north India alone there has been about 73 per cent decline in their natural popu lation during 1976-78 (Jain, 1993). Looking upon their uti li ty and importance and their implications in preserving flora, there is a need to protect and conserve these bees for pollination.

Table 2. Association of non-Apis pollinations with different crops

Crop/plant Family Bee species Reference

Alfalfa Leguminosae Megachile bicolor, M. disjunta, Kapil eta/., 1974;Abrol,1986 (Medicago sativa) M. flaviceps, M.femorata, M. lanata,

Nomia oxybeloides, N.divisus, N.pusil/a, Pithitis smaragdula and Xylocopa fenestrata

Megachile nana, M. flaviceps, M.femorata Kapil eta/., 1975 and M. cephalotes

Braunsapis spp Kapil and Jain, 1980

M. lanata, M.cepha/otes, M.cephalotes Abrol, 1985 and Pithitis smaragdula

M. flaviceps and M. nana Abrol,1986

Crop/plant Family Bee species Reference

Berseem Leguminosae M. flaviceps and M. nana Abrol, 1986b (Trifolium alexandrium)

White clover (T.repens) Leguminosae Bombus asiaticus and B. albopleuralis Abrol, 1987

Red clover (T.repens) Leguminosae Bombus asiaticus and B. albopleuralis Abrol ,1987

Pigeon pea Leguminosae Megachile lanata, M.bicolor, M.flavipes, Chaudhary and Jain 1978 (Cajanus cajan) M.cephalotes and M.femorata

Megachile lanata, Xylocopa fenestrata, Abrol,1985 X. pubescens, M.bicolor and M.cephalotes

Sunhemp Leguminosae X. pubescens and X. fenestrata Kapil and Dhaliwal, 1968 ( Crotolaria juncea)

Megachile lanata, M. fasciculata and Grewal and Singh, 1978 X. fenestrate

Megachile lanata Abrol and Kapil, 1986

M. bicolor Abrol,1987a

Pea (Pisum sativum) Leguminosae X. fenestrata, X. pubescens and Kapil eta/., 1975 Megachile lanata

Braunsapis spp Kapil and Jain, 1980

X. fenestrata, X. pubescens, Abrol,1985 M.cephalotes and M.flavipes

B.albopleuralis, Bombus asiaticu Abrol,1987 and Lasioglossum spps

Sweet potato Convolvulaceae X. fenestrata, B. albopleurali Abrol, 1987 (Ipomoea batatas) and Bombus asiaticus

Egg plant Solanaceae B. asiaticus Abrol,1987 (Solanum melongena)

X. fenestrata, Ameigilla delicata, Batra,1967 A. subcosrulea, Nomia caliphora and Pithitis spp

Onion (Allium cepa) Liliaceae Nomioides spp Kapil et at., 1975

Lasioglossum spp, Nomioides spp Abrol,1987 and X. fenestrata

Field mustard Cruciferae Nomioides, Megachilids, Andrenids Kapil et at., 1971 (Brassica campestris) and Halictids

20

- --~ _ _., _....-

Crop/plant Family Bee species Reference

Andrena i/erda and A. leaena Abrol,1986

Rape (Brassica napus) Cruciferae Andrena i/erda Mohammad, 1938

Halictids Rahman, 1940

Raya (Brassica juncea Cruciferae Andrena i/erda Kapil et at., 1971

A. teaena and And rena ilerda Abrol 1985,1986

Taramira (Eruca sativa) Cruciferae A. leaena, A. ilerda, Colletes Kapil et at., 1971 and Halictus spp

Cabbage and cauliflower Cruciferae A. ilerda, Lassiogtossum spp and Batra,1967 (B. oteracea) Pithitis smaragdula

Raddish Cruciferae Anthophora spp, Nomia spp, Batra,1967 (Raphanus sativus) Lassiogtossum spp and Colletes spp

Pumpkin and squashes Cucurbitaceae X. fenestrata, X. pubescens Hatictus Atwal,1970 (Cucurbita spp) and spp Nomioides spp

Smooth loofah Cucurbitaceae X. fenestrata, X. pubescens Kapil et at., 1965-1970 (Luffa aegyptica) and P. smaragduta

Cucumbers Cucurbitaceae Nomia spp, P. smaragduta, Nomioides Kapil et at., 1965-1970 (Cucumis melo) variegata and Halictids

Lasiogtossum spp Abrol and Bhat 1987

Cotton (Gossypium spp) Malvaceae Lithurgens attratus Batra 1977

Corriander Umbelliferae Nomioides spp, Halictidae and Abrol1985 (Corraindrum sativum) X. fenestrate

Saunf Umbelliferae Halictis spp and X. fenestrata Abrol1985 (Foeniculum vulagre)

Carrot (Dacus carota) Umbelliferae Lasiogtossum spp, Sphecoides Hy/eaus, Batra 1967 Nomioides, Braunsapis and Pfthftis smaragdula

Jowain Umbelliferae Andrena spp, Nomioides. Halictus spp Batra 1967 ( Traechyspermum amm1) and Lasioglossum spp

Orange and lemon Rutaceae Lasioglossum spp and X. fenestrata Batra 1977 Citrus spp

Guava (Psidium guajava) Myrtaceae X. pubescens, X. fenestrata Batra 1977 and Megachile lanata

Mango (Mangifera indica) Anacardiaceae Xylocopa spp, Megachite spp, Nomia Batra, 1967 spp and Lasioglossum spp

'l1

Crop/plant Family Bee species Reference j Pomegranate Punicaceae Nomioides, Lasioglossum and Batra, 1967 (Punica granatum) Halictus spp

Apples (Pyrus malus) Rosaceae Co/fetes nursei, Lasioglossum spp, Abrol, 1987 CaHulum and Osmia cornifrons

Andrena spp, Bombus haemorrhoidalis, Anon.1986 Halictus vacchalli, Osmia spp, Pithitis spp, X.fenestrata and Nomia spp I

Almond (Pamygdalus) Rosaceae Lasioglossum spp and Xylocopa valga Abrol1987

Cherry (Pavium) Rosaceae Xylocopa valga and Nomia spp Abrol1987

Pear (Pcumminis) Rosaceae Xylocopa valga and Nomia spp Abrol1987

l -

22

2.0 Carpenter Bees

L arge carpenter bees (genus Xylocopa) are wood-nesting bees and their are difficulties in mass-rearing of Xy locopa and in the high

levels of nectar robbing exhibited by the bees. They are generalist pollinators of broad geographical distribution that exhibit varying levels of sociality. Their foraging is characterized by a wide range of food plants, long season of activity, tolerance of high temperatures, and activity under low illumination levels. These traits make them attractive candidates for agricultural pollination in hot climates, particularly in greenhouses, and of night-blooming

Xylocopafenestrata foraging on chaksu

crops. Carpenter bees have demonstrated efficient pollination service in passionflower, blueberries, greenhouse tomatoes and greenhouse melons. Current challenges to the commercialization of these attempts lie in the difficulties of mass-rearing Xylocopa, and in the high levels of nectar robbing exhibited by the bees.

The carpenter bees (Xylocopa spp.) have not been cultured in a hue sense although their nesting in ce1iain areas has been encouraged by placement of soft timbers in which they can construct nesting tunnels. Because of their large size (almost an inch in length and about half as wide), they resemble large bumble bees but do not have a n·ue pollen basket on the hind leg. They are usually metallic black. The bees are solitary but numerous ones may be attracted to soft timber in which they can tunnel. This tunnel may be 1 foot long or longer and about one-half inch wide. There may be numerous cells separated by partitions formed by chips of wood cemented together. About 30 to 31 days are required for development from egg to adult.Because of their lack of gregariousness, these bees are only of limited value where appropriate nesting timbers can be provided. They also have a strong tendency to cut holes in the bases of flowers that have long slender corolla tubes.

Gupta (2003) has reported 45 species and subspecies of carpenter bees under the genusXylocopa Latreille, 1802 in the Indian region. GenusXylocopa Latreille is the single genus included within tribe Xylocopini, in subfamily Xylocopinae of the recently redefmed family Apidae. Earlier it was included under family Anthophoridae, now merged into Apidae (Michener, 2000). Many species are found up to the Pacific islands in the east and towards Russia and Africa in the west.

2.1 GenusXylocopa Latreill

Xylocopa Latreille, 1802: Hist. Nat. Ins. xv1: p.432; suppressed by Commission Opinion74(1965).

23

Xylocopa Latreille, 1802: Hist. Nat. Ins ... xvi: p.379; Type species: Apis l'iolacea

Linnaeus, 1758, by designation ofWestwood, 1840: Vol. land 2: p.86.

Genus Xylocopa Latreille is the single genus included within tribe Xylocopini, in subfamily Xylocopinae of the recently redefined family Apidae. Earlier it was included under family Anthophoridae, now merged into Apidae (Michener, 2000). Carpenter bees are almost worldwide in distribution.

Some of the principal characters helpful in recognjzing species of Xylocopa are: their large size; loss of stigma, the very long prestigma and marginal cell (Danforth, 1989), and the strongly papillate distal parts of the wings. Other distinctive features are: quite long first flagellar segment, longer than the combined length of second and third; short but distinct proboscis with heavily sclerotized components, the postpalpal part of the galea expanded like a blade and presumably used to cut into the corollas of tubular flowers to rob the nectar. All carpenter bees have three submarginal cells in their forewings but the first and second are sometimes pattly or wholly fused owing to the disappearance of the posterior part or the whole of the first submarginal crossvein. An unusual feature of most male Xylocopini, not known in any other bees, is a large gland opening on the metanotal­propodeal line. Its product seem to play a role in comtship, and its presence results in unusual sexual differences in the form and structure of the posterior part of the thorax, which becomes elongated when the gland is large (Minckley, 1994 ). Unlike other tribes of

Xylocopinae, Allodapini and Ceratini, bees of the genus Xylocopa have no aralia, though a densely hairy plata often projects somewhat between the claws. Often one can recognize

aXylocopa by their typicallyriate flying pattern.

Michener (2000) synonymized Lestis Lepeletier and Serville, 1828 and Proxylocopa Hedicke, 1938 withXj1/ocopa following the cladistic analysis ofMinckley ( 1998). Both are presently reduced to the rank of subgenera of Xylocopa. Subgenus Lestis is known

from Australia with only two species. Subgenus Proxylocopa includes the only ground­nesting carpenter bees. Its 16 species are distributed in desert areas of some pa1ts of Europe, Israel towards east up to western China (including parts of Quetta and Kashmir). Eardley (1983) illustrated the presence of the mesosomal gland in males of both subgenera, denoting their affinities withinX:rlocopa. Identification keys, separately made for the subgenera found in the Western Hemisphere and for the Eastem Hemisphere, were presented in Michener (2000). The subgenera and their included species known from our specified region are detailed in this book. Additions in localities of species known from Sri Lanka have been made from Wijesekara (200 I).

2.1.1 GenusXylocoptl SubgenusBilww ~Ia, 1938

24

Xylocopa (Biluna) Ma, 1938: p. 276; Type species: Xylocopa nasalis Westwood, 1838, original designation.

This subgenus is a group of large, elongate bees that, as far as known, nest solely in bamboos. Associated with the very smooth inner surface of this nesting substrate, are morphological modifications of the legs and mandibles. Males can be distinguished from those of all other taxa by the combination oflacking a basi tibial plate and of spines on the outer apex of the tibia. Females can be distinguished by two features of the mid-tibia: a dense mat of short, stout setae and absence of spines on the outer apex. Hurd and Moure

( 1963) considered Biluna to be closely related to subgenus Xylocopa. Minckley (1998) placed this genus as clearly allied with a clade consisting of Xylomelissa, Rhysoxyfocopa

and Nodula. In most analyses, Bifuna is shown as the most basal taxon of this clade. The subgenus ranges from India and Sri Lanka to the Lesser Sunda Islands, the Philippines and

Taiwan north to Kashmir and central China and Japan. Michener (2000) indicated that there are about five species. Based upon Maa's ( 1946) revision, Hurd and Moure ( 1963) gave a key to the species of this subgenus.

Included species

1. (lllripennis Lepeletier

• XylocopaauripennisLepeletier, 1841

• Xylocopa hemichlora Cockerell1929, Syn.

• Xylocopa phenachroa Cockere11 1929, Syn.

• Xylocopa semipurpurea Cockerell1929, Syn.

• Distribution: Calcutta, Barrackpore, Sikkim, Da1jeeling, Myanmar, China, Malaysia, Sri Lanka: Polonnaruwa.

• Tlu·ee subspecies of auripennis namely, iridipennis, mcgregori and mimetica arc recognized, among them X auripennis iridipennis Lep. has been recorded from our reg1on.

2. (lllripemzis iridipennis Lepeletier

• Xylocopa auripennis iridipennis Lepeletier, 1841

• Xylocopa ch/oroptera Lepeletier sensu Horne, 1870, Misident.

• Xy/ocopa pictipennis Smith, 1874, Syn.

• Distribution: Northwest Provinces, Punjab, Northern Rajasthan to Bengal, Madras, Bangalore, Myanmar up to Sumatra, China.

3. nmwlis Westwood

• Xyfocopa violacea (Linnaeus) sensu Donovan, 1800, Misident.

• Xylocopa nasalis Westwood, 1838

• Xylocopa nasalis nasalis Westwood, 1838

25

•

•

•

•

•

•

•

Xylocopa dissimilis Lepeletier, 1841, Syn .

Xylocopa lunulata Lepeletier, 1841 , Syn .

Xylocopa dissimilis Lcpeletier sensu Smith, 1854 (in part), Syn .

Xylocopa auripennis Lepeletier sensu Smith, 1874 (in patt), Syn .

Xylocopa anzethystina (Fabricius) sensu Smith, 1878, Misident.

Xylocopa cyanoptera Taschenberg, 1879, Nomen Nudum

Xylocopa lunulata minensis Cockere ll, 1909, Syn .

• Distribution: Northwest Provinces, Punjab, Kumaon, Srinagar (Kashmir), Bareilley, Kotdwara, Dehradun, Nagpur, Nasik, Surat, Bombay, Gboorgballi Estate, Bangalore,

Goa, Trivendrum, Cochin, Coimbatore, Madras, Godavari Distt. , Kotagiri, Malabar, Raxaul, Pusa, Chapra, Pankhabari , Siliguri , Datjeeling, Cuttak, Sikkim, Assam, Shillong, Sibsagar, Darrang, Naga Hills; Tenasserim, Myitkyina in Upper Myanmar, Fort Stedman, Yawnghwe State, Inle Lake S. end of Taungdo, S. Shan States, Moulmein, Mandalay; Java, Sharp Peak Island at Fukien, Sri Lanka: Habarana; China: Mongwan, Yunnan;

Tenasserim, Malaysia, Borneo, Philippines, Palau and Madagascar.

4. trcmquebarorum concolomta (Ma)

• Xylocopa pictifrons con colora/a Ma, 1938

• Distribution: Sikkim (12,500'), Singla (Dcujeeling district), Kumaon, China, Java

5. trauquebarorum tranquebarorum (Swederus)

• Apis tranquebarorum Swederus, 1787

• Xylocopa orichalcea Lepeletier, 1841, Syn.

• Xylocopapictiji-ons Smith, 1852, Syn.

•

•

•

Xylocopa attenuata Perez, 190 I , Syn .

Xylocopa kelloggi Cockerell, 1931 , Syn .

Xylocopa onzeiana Ma, 1936, Syn .

• Distribution: Described from Ile de Fer? and Taiwan. Smith (1854) recorded from Bengal and China (as oriclzalecea); Cockerell (1931) noted from Chusan and Formosa (as

kelloggi), Java, Gnatong (Sikkim), Kumaon.

2.1.2 GenusXylocopa Subgenus Copoxyla Maa, 1954

• Xylocopa (Copoxyla) Maa, 1954: p. 211; Type species: Apis bomb. iris Christ, 1791, =

Xylocopa cyanescens Brulle 1832, by original designation. [Cluist's name has sometimes been regarded as an invalid trinominal, as below].

26

j

l

1

This is a subgenus that includes the smallest, bright-metallic species of the Xylocopini. They are more or less morphologically similar to Ctenoxylocopa. Females can be distinguished by the combination of a smoothly rounded thorax, the absence ofpreapical spines beside the pygidial spine, the presence of tridentate mandibles, basitibial plate extending nearly to base of tibia. Males differ from the males of Ctenoxylocopa by the presence of a propodeal triangle, small body size, face enti rely dark, a s ingle spine at the outer apex of the hind tibia, apex of basitibial plate bifid, gonostylus with a weakly

developed medially projecting lobe, and seventh tergum with pair of apical dentiform processes (Minckley, 1998).

This subgenus occurs from the MediteiTanean basin north to Slovakia, east to Russia. There are three to possibly six species (Michener, 2000). Popov and Ponomareva (1961) gave a key to Russian species. One species, as noted below, reaches our area in the extreme northern region.

Included species

1. cyanescens Brulle

• Apis bomb. iris Christ, 1791, Unav.

• Xylocopa cyanescens Brulle, 1832

• AJ'Iocopa min uta Lepeletier, 1841, Syn.

• A:);focopa tau rica Erich son, 1841, Syn.

• Xy/ocopa virescens Gistel, 1857, Homo. (nee Lepeletier, 1841)

• A:) locopa canula Rondani, 1874, Syn.

• Xylocopa virescentis Strand, 1917, Syn., replacement for virescens Gistel.

• Distribution: East ofMediterranean basin, northern region ofKashmir, and S. E. Russia.

2.1.3 GenusXylocopa Subgenus Ctenoxylocopa Michener, 1942

• Xylocopa (Ctenopoda) Ma, 1938: p. 285 (nee McAtee and Malloch, 1933); Type species:

Apisfenestrata Fabricius, 1798, by original designation.

• Xylocopa (Ctenoxylocopa) Michener, 1942: p . 282, replacement for Ctenopoda Ma, 1938; Type species: Apisfenestrata Fabricius, 1798, auto basic.

• Baana Sand house, 1943 : p .530, replacement for Ctenopoda Ma, 1938; Type species: A pis f enestra fa Fabricius, 1798, auto basic and by original designation.

Minckley ( 1998) in his cladistic analysis judged this subgenus most closely related to some members of the subgenus AJ1locopa s. st1: Distinctive features of males are: posterolateral lobes ofpronotum prolonged posteriorly so that they are nearly in line with the preepisternal groove on the mesepisternum, and the spiracles on tergum 3 bear an

27

elevated, posteriorly directed, scalelike process. Females can be distingu ished by the well-developed bas i tibial plate with a row of tubercles along each lateral edge, apical margin of mandibles with two teeth, and hind tibia with one spine on its outer apex.

Michener (2000), while compiling its distribution, noted that this subgenus occurs from Natal, South Africa, north to Gambia and Ethiopia, northeast to the Arabian Peninsula, Israel, Iraq, trans-Caspian Russia, and southeast to Madagascar, Mauritius, Sri Lanka, Pakistan, India and Myanmar. Around six species were listed by Hurd and Mome (1963) and were revised by Maa (1970). Two of the described species are found in our region;

among them, X f fenestrata is almost cosmopolitan.

Included species

1. basalis Smith

• Xylocopa basalis Smith, 1854

• Distribution: Pakistan: Wazirabad, Karachi; India: Alwar, Pali, Mount Abu, Bombay, Veeraval, Junagarh.

2. fenestrata fenestrata (Fabricius)

• Xylocopa fenestmta Fabricius, 1798

• Xylocopa indica Klug, 1807, Syn.

• Xylocopa lunata Klug, 1807, Syn.

• Xylocopa serripes Burmeister, 1876, Syn.

• ~J 'locopa gardineri Cameron, 1902, Syn.

• Xylocopa serripes Hedicke, 1938, Homo. (nee Burmeister, 1876)

• ~1'locopa hedickei Maa, 1940, Syn.; replacementforserripes Hedicke.

•

•

28

Xylocopa bombayensis Maa, 1954, Syn .

Distribution: Srinagar, Ramban, Udhampur, Kangra Valley, Choa, Gandhala Reserve Forest, Kallar Kahar, Delhi, Bijnor, Mailani, Kichha, Naini Tal, Debra dun, Roorkee,

Bareilly, Tret, Mussoorie, Agra, Alwar, Jodhpur, Asirgarh, Mandla, Nagpur, Bombay, Ratnagiri, Media, Ahmedabad, Salsette, Goa, Coimbatore, Malabar, Bangalore, Ramnad,

Nalla Malai Hills, Barkuda Islan, Chilka Lake, Hazaribagh, Parasnath, Calcutta, Dakhindari Salt Lake, Manbhum, Shantiniketan, Kharagpur, Raneeganj, Darjecling, Companyganj (Assam); Nepal: Pipal Hali, Nepal Terai; Myanmar: Minhla, Maymyo, Shah Plateau, Mandalay, Tatkon, Shwebo; Sri Lanka: Niroddumunai, Trincomalee, Colombo, Kandy, Puttalam, Anuradhapura, Hambantota, Galle, Matale, Ratnapura; Maldives; Persia: Bendar Abbas; Baluchistan: Pasni, Mehan Coast, Pakistan: Rawalpindi, Karachi and, Celebes. [A:))locopa serripes Hedicke, 1938 was described from

l

Bushire, Iran and another female was recorded from Nepal; Ma (1938) considered the second locality very improbable].

2.1.4 GenusXylocopa SubgenusKoptortosoma Gribodo, 1894

•

•

•

•

•

•

•

• •

Koptortosoma Gribodo, 1894: p. 271; Type species: Koptortosoma gabonica Gribodo, 1894, by designationofSandhouse, 1943, 1943: p. 561 [Michener, 1997].

Koptorthosoma Dalla Torre, 1896: p. 202; unjustified emendation of Koptortosoma Gribodo, 1894.

Cyaneoderes Ashmead, 1899: p. 70; Type species: Cyaneoderes fairchildi Ashmead, 1899 =Bomb us coeruleus Fabricius, 1804, by original designation.

Coptorthosoma Perez, 1901: p. 3; unjustified emendation of Koptortosoma Gribodo, 1894.

Xylocopa (Orbitella) Ma, 1938: p. 305 (not Douvialle, 1915); Type species: XyLocopa confusa Perez, 1901, auto basic.

Xylocopa (Maiella) Michener, 1942: p. 282, replacement for Orbitella Ma, 1938; Type species: X; locopa confusa Perez, 1901 , auto basic.

Ewyapis Sandhouse, 1943: p. 551, replacement for Orbitella Ma, 1938; Type species: Xylocopa confusa Perez, 1901, auto basic.

Xylocopa (Eoxylocopa) Sakagamiand Yoshikawa, 1961: p. 413 , nomen nudum .

XyLocopa (Cypho;._y/ocopa) Hurd and Moure, 1963: p. 283; Type species: Xylocopa

ocuLaris Perez, 1901 , by original designation.

• Xylocopa (Aji-oxylocopa) Hurd and Moure, 1963: p. 264; Type species: Apis nigrita Fabricius, 1775, by original designation.

• XyLocopa (Oxyxylocopa) Hurd and Moure, 1963: p. 275; Type species: Xylocopa varipes Smith, 1854, by original designation.

• Xylocopa (Lie.ftinckella) Hurd and Moure, 1963: p. 286; Type species: Xylocopa smithii Ritsema, 1876, by original designation.

This is the largest subgenus within Xylocopa, widely distributed in the Eastem Hemisphere. According to Minckley ( 1998) it is the sister group of Mesotrichia due to overall resemblances. The female scutellum has a sharp transverse tnmcation overhanging the metanotum (except in X sinensis), similarly in males the beginning of the posterior thoracic declivity is sharply angled (except in X sinensis). Males have unmodified tegulae, unlike the elongate tegulae ofMesotrichia.

Michener (2000) compiled its overall distribution as throughout sub-Saharan Africa as well as the Mediterranean countries of Africa (Morocco to Egypt), Dalmatia,

29

southwestern Asia and, in east up to Philippines, Taiwan and Japan, and south through Indonesia, New Guinea, and the BismarckA:rchipelago to southernmostA:ustralia.

Included species:

1. aestuans aestuans (Linnaeus)

• Apis aestuans Linnaeus, 1758

• Apis leucothorax DeGeer, 1773, Syn.?

• Xylocopa confusa Perez, 1901, Syn.?

• Distribution: Tauk (N W Province); Baluchistan: Pasni, Mek.ran Coast; Punjab: Khewra,

Salt Range, Karor Range, E. Rawalpindi, Choa, Shah pur, Lahore; Karachi; India: Delhi, Kangra Valley, Dehradun, Kichha, Naini Tal, Bijnor, Meemt, Basha Ghat (M.P.), Sawai

Madhopur, Kota, Hoshangabad, Bclgaum, Poona, Nasik, Andheri, Salsette (Goa), Bangalore, Cuddapah, Razampeta, Palnis, Coimbatore, N. Vellore, Madura, Kurnool (Tamilnadu); Raxaul, Bhaga1pur, Pusa, Taljhari, Santhal Parganas (Bihar); Chilka Lake (Orissa), Santin iketan, Birbhum (W. Bengal), Assam, Malaysia, Sri Lanka: Anmadhapura, Matale; Myanmar: Mergui; Jore, Singapore, Sumatra: A:nei Kloof, West coast of Sumatra, Fort de Kock; Java: Lambreth, Buitenzorg; Batavia, Borneo, Ambo ina, Saigon, Nepal: Chatri Gouri, Nepal Terai, and Africa.

2. abbotti (Cockerell)

• Mesotrichia abbotti Cockerell , 1909

• Xylocopa coerulea (Fabricius) sensu Bingham, 1897, Misidcnt.

• Distribution: Trang (Lower Siam), Tenasserim.

3. b1yorum (Fabricius)

• Apis bt}'orum Fabricius, 1775

• Xy/ocopa aestuans (Linnaeus) sensu Smith, 1864 and 1874, Misident.

• Distribution: Sikkim, Sri Lanka, Myanmar, Malabar up to Australia; Northern India (Smith, 1874).

4. coerulea (Fabricius)

• Bomb us coeruleus Fabricius, 1804

• Xy/ocopa semiarmenia Wiedemann, 182 1, Syn.,

• CyaneoderesfairchildiAshmead, 1899, Syn.

• Xylocopa caeruletformis Meade-Waldo, 1914, Syn.

• Xylocopa.fusca Meade-Waldo, 191 6, Syn.

30

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5.

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6.

• •

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7.

•

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8.

•

• 9.

•

•

10.

•

Xylocopa caerulea Auctt.

Distribution: Sikkim, Myanmar, Sri Lanka, Malaysia, Indo-China, Borneo, Java, New Caledonia(?).

flavicollis (DeGeer)

Apisflavi-collis DeGeer, 1778

Apis citronella DeGeer, 1 778, Syn .

Apis collaris Olivier, 1789, Syn .

Xylocopa divisa Klug, 1807, N. Syn. (by Hurd, Personalcomm.)

Distribution: Ma ( 1938: p. 328) indicated it (as Xylocopa divisa Klug) was distributed in Africa, but Dalla Torre (1896) added India to its distribution. In fact, it has never been

found in any part of Asia.

jlavonigrescens Smith

Xy locopaflavo-nigrescens Smith, 1854

Xy locopa malayana Cameron, 1901, Syn .

Mesotrichia confusa viridissima Cockerell, 1918, Syn .

Distribution: Sikkim, Dmjeeling, Sylhet (Assam), Andaman Islands, South Tenasserim,

Singapore, Siam; Penang (Malaysia); Tavoy (Myanmar); Upper Tenasserim.

hafiziiMa

Xylocopa hafiz ii Ma, 193 8

Distribution: Karwar, Jog, N. Kanara Dish·ict, Mahabaleshwar (Maharashh·a); Soccorro, Malabar (Goa); Sagar, Sbimoga, Coorg, Pollibetta (Karnataka); Nagarcoil, Trivendrum, Tenmalai (Kerala).

minor Maidl

Xy locopa (Koptorthosoma) 1ninor Maidl, 1912

Distribution: Sikkim .

provida Smith

Xy locopaprovida Smith, 1864

Distribution: Smith (1864) described from Mysol and Wigiou, Sunda Islands and was later recorded from North em India (Smith 187 4 ); Sri Lanka.

rujicornis Fabricius

Xylocopa rujicornis Fabricius, 1804

~1

• •

• •

Xylocopa verticalis Lepeletier, 1841, Syn .

Xylocopa bryorum (Fabricius) sensu B ingbam, 1896, Misident.

Xylocopa ceylonica Cameron 1901, Syn .

Xylocopa cia Pi crus Maidl, 1912, Syn .

• X (Koptorthosoma) separata Perez sensu Maidl, 1912, Misident.

• Xylocopa verlicillata Lieftinck, 1957, Syn.

• Distribution: Kashmir: Srinagar; South India: Bangalore, Barkuda Island, Madras; Orissa: Puri, Chilka Lake, Ganjam District; Bihar: Dalkhola, Pumeab; W. Bengal: Calcutta, Siliguri, Jalpaiguri, Camilla; Sikkim: Mangpu, Barnckpore, Murshidabad; Sri Lanka: Andaravela, Anuradhapura, Badulla, Mannar, Murunkan; Philippine: Luzon; Annam, Siam, Java, Malaysia: Kuala Kangsar, Perak; Sumah·a: Medan, Celebes up to

Australia(?).

11. separata Perez

• XylocopaseparataPerez, 1901.

• Distribution: India (Exact location not known)

2.1.5 GenusXylocopa SubgenusMaaiana Minckley, 1998

32

Xylocopa (Maaiana) Minckley, 1998: p. 32; Type species: Xylocopa bentoni Cockerell, 1919: p. 172, by original designation.

Ma ( 1938) noted that subgenus Nodu/a is represented by two "natural groups". Minckley ( 1998), while making cladistic analysis of the subgenera, recognized that the group typified by X bentoni should be separated as the subgenus Maaiana. He included five species in this subgenus namely, X angulosa, X ben toni, X bicristata, X punctigena and X punctilabris. A key to the species, as the punctigena group of Nodula was presented by Maa (1954) and in modified form by Hurd and Moure (1963). Michener (2000) indicated that six species belong to this subgenus, the sixth being X meyeri.

Males of this subgenus have an abrupt beginning of the posterior declivity on the

scutellum, and possess a mesosomal gland reservoir (different from Nodula), have a spine on the outer apex of the hind tibia (different from Xylomelissa), and have unmodified propodeal spiracles (different from Rhysmylocopa). Females of Maaiana can be distinguished from those of the subgenus Rhyso,\}"locopa by the presence of a carina dorsal to each lateral ocellus and a sharply rounded scutellum.

This subgenus is nan·owly dish·ibuted, from Turkestan and Afghanistan to India and Sri

Lanka. Michener (2000) included six species [wrote: species are the first six. in the key to the males ofNodula prepared by Hurd and Moure (1963)] in this genus and further wrote it to be confined to India and Sri Lanka. However, Xylocopa angulosa and Xylocopa

I

bicristata were described from Afgharustan ("Wama, Nuristan"), and Xylocopa punctilabris was found in Turkestan. In addi tion to these, X meyeri was included by him as sixth species (described from south I ndia). It was originally placed in Nodula by Ma

(1938) and according to Minck ley (personal communication), should remain in Nodula until the type is studied. The remaining two are as follows:

Included species

1. bentoni Cockerell

• Xylocopa benton i Cockerell, 1919

• Xylocopa kotzschi Hedicke, 1938, Syn.

• Distribution: Nmih Western Frontier Province: Abbottabad, Andarab, W. Hindu Kush (Afghanistan); Dhannsala, Kaisdhar, Kullu; Uttar Pradesh: Mossy Nullah, Mussoorie, Dehradun, Ranikhet [on Robinia sp., Acacia sp. and Calotropis sp.], Lol ba, Garhwal, Kuma on, Dehradun, Chakrata.

2. punctigena Ma

• Xylocopa (Nodula) punctigena Ma, 1938

• Distribution: Punjab: Hamirpw· Range, Hoshiarpur, Uttar Pradesh: Bhadraj, Mussoorie

GenusXylocopa Subgenus Mesotrichia Westwood, 1838

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•

Mesotrichia Westwood, 1938: p. 112; Type species: Mesotrichia ton·ida Westwood, 1938, monobasic.

Xylocopa (Platynopoda) Westwood, 1840: p. 271; Type species: Apis latipes Drury, 1773, by designation of Ashmead, 1899: p. 71.

Xylocopa (Audinetia) Lepeletier, 1841: p. 203; Type species: Apislatipes Drury, 1773, by designation ofSandhouse, 1943: p. 529.

Platinopoda Dalla TaiTe, 1896: p. 202, laps us for Platynopoda Westwood, 1840 .

Andineta Ashmead, 1899 p. 71, incorrect subsequent spelling for Audinetia Lepeletier, 1841.

Andineta Ashmead, 1899 p. 97, incorrect subsequent spelling for Audinetia Lepeletier, 184 1.

Xylocopa (Hoplitocopa) Lieftinck, 1955: p. 27; Type species: Xylocopa assimilis Ritsema, 1880, by original designation.

Xylocopa (Hoplmylocopa) Hurd and Moure, 1963: p. 260; Type species: Xylocopa acutipennis Smith, 1854, by original designation.

This subgenus, like Koptortosoma, can be distinguished from other subgenera by the

right-angular or acute scutellar beginning of the fema le thoracic declivity. Females have craterlike supraocellar pits and their hind trochanters roughly triangular in ventral view. Males can be distinguished by posteriorly elongate tegulae. Most males have greatly widened front or middle tarsi (except X assimilis Ritsema and X acutipemzis Smith). In a narrow sense this subgenus contains species with the anterior coxal spine of the male short. However, in the historically well-known group Platynopoda, males have a long anterior coxal spine (Michener, 2000).

Th is subgenus is widely distributed in sub-Saharan Africa and northeast as far as Iran. Many species are identified that belong to the oriental region, occuning in Sri Lanka notih to Kashmir, eastward through the Lesser Sunda Islands, Sumatra, Borneo and southeast Asia to the Philippines.

Included species

1. acutipennis Smith

• Xylocopa acutipennis Smith, 1854

• Xylocopa splendidipennis Ritsema, 1876, Syn.

• Distribution: Singla, Sevok, Kalimpong, Sitong, Pasbok, Tindharia, Mangpu (all tn

Datjeeling District), Sikkim, Sylhet, N. Khasi, Garo Hills; Myanmar: Dwana Hills, southern Myanmar, Tenasserim; Nepal: Kathmandu, Nayorkorte, Nepal Valley.

2. assimilis Ritsema

• Xylocopa assimilis Ritsema, 1880

• Xy/ocopa gastrica Maa, 1939, Syn.

• Distribution: Kashmir.

3. /atipes (Drury)

• Apis latipes Drmy, 1773

• Apis gigas DeGeer, 1773, Syn.

• Mesotrichia (Platynopoda) latipes basiloptera Cockerelll917, Syn.

• Distribution: Bengal: Pankhabari, Singla, Darjeeling, Kalimpong, Calcutta, Balasan Forest, E. Himalayas, Mangpu; Sikkim; Assam: Therria Ghat, Khasi Hills, Sibsagar; Meghalaya : Shillong, Naga Hills, Myanmar: Logae, Tavoy, Upper Tenasserim: Mergui, Tenasserim, Mekane, Hopin, Myitkyina Dist., Maymyo; Malaysia: Johore, Perek; Singapore; Sumatra, Borneo, Sarawak, Sundakan, Sintong; Siam. China, Amboina,

Philippines.

4. magnifica (CockereiJ)

• Mesotrichia latipes var. magnifica Cockerell, 1929

34

j

Distribution: Siam, Bengal: Balasan Forest, E. Himalayas, Maini Muk.h, Chittagong

Hills Tracts.

5. perforator Smith

• Xylocopa pe1jorator Smith, 1861

• Xylocopa pe1joratrix Schulz, 1906, Unjustified Emend.

• Distribution: Sri Lanka, Sumatra: Medan; Java: Buitenzorg; Lambok; Ternate (Sunda Islands).

6. tenuiscapa Westwood

• Xylocopa (P!atynopoda) tenuiscapa Westwood, 1840

• Xylocopa !atrei!lei Lepeletier, J 841, Syn.

• Xylocopa viridipennis Lepeletier, 1841, Syn.

• Xylocopa /ativentris Blanchard, 1844, Syn.

• Xy locopa albo-fasciata Sichel, 1867, Syn.

• Xylocopa esica Cameron, 1902, Syn.

• Xylocopa tenuicornis Ashmead 1904, Lapsus

• Distribution: Srinagar, Kashmir; Saharanpur, Dehradun, Sat Tal, Nagpur, Ratnagiri District, Mahabaleshwar, Satara, Sangli, Tavargatti, Belgatml, Karwar, Bassein, Marmagao, Soccorro, Shimoga, Bangalore, Trivendrum, Peermade, Kodaikanal , Palni Hills, Barkuda Island, Chilka Lake, Bellary, Palkonda Hills, Kangumaduga, Seshachalama, Cuddapah District, Vempali, Nadur, Javadi Hills, Denkanikota, Salem, Yercaud, Kuttur, Cumbun, Anamalai Hills, Tiruchirapalli, Palnis, Kodaikanal, Vellore, Orissa: Barkul, Gopkunda Island, Chill<a Lake, Puri, Bihar: Nalanda, Purneah, Ranchi, Champaran, Bettiah; Bengal: Balasan Forest, Calcutta, Panighata, Dakhindari Salt Lake,

Murshidabad, Sbantiniketan, Birbhum, Dmjeeling District, Pulta near Barrackpore, Saraghat, Kalimpong, Sikkim; Assam: Companyganj, Khasi Hills, Margherita; Nepal: Monda; Sri Lanka: Ampara, Colombo, Galle, Hambantota, Monaragala, atnapura, Bandaravela, Negombo, Perdcniya, Niroddumunai near Trincomalee; Maldives, Andamans, China (Yunnan), Siam (Chieng Mai), Java and Timor.

2.1.6 GenusXylocoptt Subgenus Nodula Ma, 1938

• Xylocopa (Nodula) Ma, 1938: p. 290; Type species: Apis amethystina Fabricius, 1793, by original designation.

Subgenus Nodu/a is allied to Maaiana, Xy/omelissa (s. lata) and Rhysoxylocopa. Both

sexes have their beginning of posterior thoracic declivity abruptly rounded. The males of Nodu/a can be distinguished from those of Xy/ome/issa by the presence of a spine on the

35

outer apex of the hind tibia of the male and from those of Rhysoxylocopa by the unmodified propodeal spiracles. Females of Nodula bear two spines at outer apex ofhind tibia and can be distinguished from those of the subgenus Rhysoxylocopa by carinae dorsal to the lateral ocelli and a sharply rounded scutellum.

Included species

1. amethystiua amethystina (Fabricius)

• Apis amethystina Fabticius, 1793

• Xylocopa ignita Smith 1874, Syn.

• Xylocopa amethystina sigiriana Cockerell, 1911, Syn.

• Distribution: Punjab, Northwest Provinces, Kumaon, Bombay, Malabar, Sri Lanka: Am para, Anuradhapura, Matale, Polonnaruwa.

2. amethystiua phallerocepha/a Cockerell

• Xylocopa amethystina phanerocephala Cockerell, 1920

• Distribution: Debra Dun, Bombay, SoccmTo (Goa), Quilon (Kerala), Vishakhapattanam, Golkunda, Nilgitis, Ootacamund, Bababuddin Hills, Bangalore (Kamataka), Kotagiri (Nilgiris), Salem, Cuddapah, Kuttur and Ooty (Tamilnadu); Sri Lanka: Niroddumunai, M. Iluppala, N. Sri Lanka.

3. madureusis Friese

• Xylocopa madurensis Friese, 1913, DEZ VII: p. 88.

• Distribution: Madura, Madras, Kodaikanal.

4. meyeri Dusmety Alonso

• Xylocopa (Xylocopa) meyeri Dusmety Alonso, 1924

• This species was placed, possibly inadvertantly, in the subgenus Maaiana by Michener (2000), when he stated that the fiTst six species in Hurd and Mome's (1963) key to males of Nodula were the six species those were placed in Maaiana. This differs, however, from Minckley's original placement of only five species into Maaiana, as indicated above. Neither Michener nor Minckley bas examined the type of meyeri, however (Minckley, pers. comm.), and at present we consider it to be a species of Nodula, following Ma's (1938) treatment of the species as a possible synonym of X madurensis, whose placement inNodula is not in doubt.

• Distribution: Sou them India.

5. uigrotarsata Ma

• _A);focopa (Nodula) nigrotarsata Ma, 1938

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• Distribution: Guduholli, Bombay.

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prashadi Ma

Xylocopa (Nodu/a) prashadiMa, 1938

Distribution: Madras, Coonoor, Nilgiris .

nmwkrislmai Ma

Xylocopa amethystina sensu Cockerell, J 9 J 1 (nee Fabricius)

Xylocopa ignita Cockerell, 1919 (nee Smith), Female .

Xylocopa amethystina Dover, 1922 (in part, nee Fabricius)

Xylocopa ramakrishnaiMa, 1938: p. 296 .

The specimens that constitute the syntype series of this species have apparently not been examined since its description, and presumably the taxon still lacks a designated lectotype. It was considered by Ma (1938) to be closely related to X prashadi.

Distribution: Dodabetta, Niligirs (Tamilnadu) .

remota Ma

Xylocopa (Nodula) remota Ma, 1938

Distribution: South India(?) .

GenusXylocopa SubgenusNyctomelitta Cockerell, 1929

• Xyfocopa (Nyetomelitta) Cockerell , 1929: p. 303; Type species: Bombus tranquebaricus Fabricius, 1804, by original designation.

Both sexes of the species of this subgenus can be distinguished from all other members of the genus Xylocopa by their large size, large eyes, convergent above, and greatly enlarged ocell i, greater in diameter than that of antenna! socket. Upper carina in mandibles with subapical tooth. In males, outer apcxofh ind tibia with one spine, hind femur w ithout basal spine or tubercle, apex ofbasitibial p late bifid and mesosomal reservoir present. Females

w ith graduli on Tl-T5 (Cockerell, 1929; Ma, 1938 and 1940; Minck ley, 1998). All species are nocturnal and are exclus ively found in the oriental region.

Included species

1. proximata Ma

• Xylocopa (Nyetomelitta) proxima fa Ma, 1938

• Distribution: Andaman Islands.

2. tranquebarica (Fabl"icius)

• Bombus tranquebaricus Fabricius, 1804

37

• Xylocopa rL!f'escens Smith, 187 4, Syn.

• Distribution: Pachgani (W. Ghats), Bombay, Karwar, Bangalore, Bababuddin Hills, Coorg, South Malabar (Tamilnadu), Coonoor, Sappel, Palghat, Darjeeling, Jalpaiguri, Sikkim, Rangoon (Myanmar), Petrsut Reserve, Katha Division, Maymyo, Shan Plateau, in Sri Lanka: Kanthaley, Niroddurnunai (Trincomalee), Vilankulam, Colleetek (N.P.), Siam, Tenasserim, Java, Sumatra and Borneo.

2.1.7 GenusXylocopa SubgenusProxylocopa Hedicke, 1938

• Xylocopa (Proxylocopa) Hedicke, 1938: p. 192; Type species: X olivieri Lepeletier, 1841, by original designation.

• Ancylocopa Maa, 1954: p. 190, 198; Type species: Xylocopa nitidiventris Smith, 1878, by original designation.

This subgenus include rather small, dull-colomed species, notable for their long faces. Minck ley ( 1998) noted that this feature probably is associated with loss of the strong

mandibular musculature for gnawing in wood. Females can be easily distinguished by the well-developed basitibial plate located at the base of the hind tibia and by the well­

developed elevated pygidial plate; metapostnotum absent. Males can be recognized by outer apex of hind tibia with two spines; hind femur without basal tubercle or spine;

mesosomal gland reservoir consisting of two small, widely separated pouches. A more distinct feature of males is the presence of short, ovate, parapsidal lines, no more than three times as long as wide. Some species have large ocelli, denoting nocturnal habits. They are known for their exclusively ground-nesting habits, and have accordingly long been considered a separate genus, presumably more primitive than other Xylocopines, though Minckley's ( 1998) phylogenetic analysis indicated otherwise.

The subgenus ranges from Albania, Greece, and Israel east to westem China (Michener, 2000). The lone species that reaches our area is X rl(/a.

Included species

1. rufa Friese

• Xylocopa o/il•ieri rL![a Friese, 190 1

• Xylocopa eril·anensis Perez, 1901, Syn.

• Distribution: Turkestan, Turkmenia, Quetta to extreme northern India.

2.1.8 GenusXylocopa SubgenusXylocopa Latreille, 1802 s.str.

• Xilocopa Latreille, 1802: HNGP .. xvi: p.432; suppressed by Commission Opinion 743 (1965).

• Xylocopa Latreille, 1802: HNGP .. xvi: p.379; Type species: Apis violacea Linnaeus, 1758, by designation ofWestwood, 1840: Vol. 1 and 2: p.86.

38

Minck ley ( 1998) studied the type species along with X valga to delineate the cladistic relationships for this much restricted subgenus. He concluded that X violacea shows clear

affinities with the New World subgenus Xylocopoides, whereas X valga is most closely related to the Old World subgenus Ctenoxylocopa. Females have two spines on the outer

apex of the hind tibia and the apical edge of their mandibles are tridentate. Males can be distinguished by combination of face without yellow maculations (i.e. completely black); the rounded profile of the posterior part of the thorax; the two spines on the outer apex of hind tibia; hind femur without basal tubercle or spine.

This subgenus contains 8 diverse species ranging from Europe (nmth as far as Germany) up to Russia and in the east up to Afghanistan, and western Himalayas (Michener, 2000). Besides the following two species, X rogenho.feri Friese might also be found in Kashmir orBaluchistan(Ma, 1938).

Included species

1. Vftlga Gerstaecker

• Xylocopa valga Gerstaecker, 1872

• Xylocopa ramulorum Rondani, 1874, Syn.

• Xylocopa convexa Smith, 1878, Syn.

• Xylocopa pyropyga Friese, 1913, Syn

• Distribution: Widely dish·ibuted in Palaearctic, Meditenanean, north-European and the Siberian regions besides following territories in Asia: Jhelum Valley (5200'), Gilgit, Srinagar (Kashmir); Haran Plateau, Sindh Valley, N. W. Frontier Province: Karakal,

Bumboret Valley, Chih·al; Ustui Gol, Rambhar Valley, Chit-raJ, Abbotabad, Chamba, Shah p ur (Punjab); Golpara (Assam); Turkey: Taurus Pampe, Asia Minor; Turkestan,

Nun-bulak; China: Kogyar, E. Turkestan, Yarkand, Hechyei, Kohtai, Kansu, Manchouli.

2. violacea Linnaeus

• Apis violacea Linnaeus, 1758

• Apis insubrica Myller, 1766, Syn.

• Xylocopa.femorata Fabricius, 1804, Syn

• Distribution: Thelum Valley, Gilgit, Srinagar in Kashmir; Sindh Valley, North West Frontier Province: Karakal, Bumboret Valley, Chitral, Abbotabad; Punjab: Chamba,

Shahpur, Ludhiana, Pathankot; Turkey: Taurus Pampe, Turkestan in Turkmenia; China: Kogyar, E. Turkestan, Hechyei, Kohtai, Kansu, Manchouli.

2.1.9 GenusXylocopa SubgenusZonohirsuta Ma, 1938

• Xylocopa (Zonohirsuta) Ma, 1938: p. 300; Type species: Xyfocopa collaris Lepeletier,

1841 (nee Apis collaris Oivier, 1789) = Xylocopa dejeanii Lepeletier, 1841, by original designation.

Minckley (1998) noted this subgenus is similar to Prosopoxy locopa. In Zonohirsuta the basi tibial plate in females is open basally, both of the lateral edges are strongly carinate and project perpendicularly from the tibial surface. Female hind tibial outer apex with two spines and beginning of posterior thoracic declivity of female abruptly rounded. In males hind tibial outer apex with only one spine; metapostnotum present and mesosomal gland reservoir present.

This subgenus has been recorded from Tibet, India and Sri Lanka thence eastward tlu·ough Myanmar to southeast Asia and the Philippines, Indonesia up to Sulawesi (Michener, 2000). Xylocopa dejeanii and its subspecies have been listed below, the other two, Xylocopa xanti and Xylocopa mazarredoi are known from Palawan and Borneo, respectively.

Included species

1. dejeanii dejeanii Lepeletier

• Xylocopa collaris Lepeletier, 1841 , Homo. (nee Olivier, 1789)

• Xylocopa dejeanii Lepeletier, 1841

• Xj;locopa b1yanti Cockerell, 1919, Syn.

• Distribution: Kumaun, Allahabad, Sikkim, Myanmar, Malaysia up to Siam, Sri Lanka: Anuradhapura, Colombo.

2. dejeanii bhowara Ma

• Xylocopa collaris bhowa ra Ma, 193 8

• Distribution: Kalutara and Yakambi in Maharashtra; Trivendrum, Temnalai, Cochin in Kerala; Sri Lanka: Kandy, Niroddumunai, Colombo, Ampara, Galle, Kalutara, Anuradhapura, Matale, Badurelia; Palypitiya (N W Province); Andamans: Hope Town (Pani Ghat).

3.

•

•

•

4.

•

40

dejeanii bingluuni Cockerell

Xylocopa colla ris Lepeletier, 1841 (in part)

Xylocopa dejeanii binghami Cockerell, 1904

Distribution: Nalaonda, Dehradun, Datjeeling, Teesta, Siliguri, Samsing, Kalimpong, Riijang, Singla Bazar, Sikkim; Sibsagar, Naga Hills, Marghertia, Sadiya, Khasi Hills (Assam); Khamba Jong (Tibet).

dejeanii nigrocaerula Smith

Xylocopa nigro-caerula Smith, 1874

• Distribution: Sri Lanka, Celebes.

5. dejeanii penaugensis Cockerell

• Xylocopa collaris penangensis Cockerell, 1918

• Distribution: Malaysia: Penang, Kwala Kangsar, Perak, Johore; Myanmar: Mergui; Upper Tenasserim; Moulmein, Tenasserim, Thaungyin Valley, H lebwe, Katba division,

Bodaung, S. Tougoo.

Other subspecies of X dejeanii known from other east Asian counn·ies (including China)

are:

• dejeanii alboxantha Maa

• dejeanii chenghuoi Maa

• dejeanii sauteri Friese

• dejeanii yangweiella Maa

• dejeanii indecisa Cockerell and LeVeque

• dejeaniifuliginata Perez\

• dejecmii kanoiYasumatsu

Carpenter bees are almost worldwide in distribution . Some of the principal characters helpful in recognizing species of Xylocopa are: their large size; loss of stigma, the very long prestigma and marginal cell (Danfmth, 1989), and the strongly papillate distal pat1s of the wings. Other distinctive features are: quite long first flagellar segment, longer than

the combined length of second and third; short but distinct proboscis with heavily sclerotized components, the postpalpal part of the galea expanded like a blade and presumably used to cut into the corollas of tubular flowers to rob the nectar. All carpenter bees have three submarginal cells in their forewings but the first and second are sometimes partly or wholly fused owing to the disappearance of the posterior part or the whole of the first submarginal crossvein. An unusual feature of most male Xylocopini, not known in any other bees, is a large gland opening on the metanotal-propodealline. Its product seem to play a role in courtship, and its presence results in unusual sexual differences in the form and structure of the posterior part of the thorax, which becomes elongated when the gland is large (Minckley, 1994). Unlike other n·ibes of Xylocopinae, Allodapini and Ceratini,

bees of the genus Xylocopa have no arolia, though a densely hairy plata often projects somewhat between the claws. Often one can recognize a Xylocopa by their typical lyriate flying pattern.

M ichener (2000) synonymized Lestis Lepeletier and Serville, 1828 and Proxy locopa

Hedicke, 1938 with Xylocopa following the cladistic analysis ofMinckley (1998). Both are presently reduced to the rank of subgenera of Xylocopa. Subgenus Lestis is known

41

from Australia with only two species. Subgenus Proxy locopa includes the only ground­nesting carpenter bees. Its 16 species are distributed in desert areas of some parts of Europe, Israel towards east up to western China (including parts ofQuetta and Kashmir). Eardley (1983) illustrated the presence of the mesosomal gland in males of both subgenera, denoting their affinities within.X~}'/ocopa. Identification keys, separately made for the subgenera found in the Westem Hemisphere and for the Eastem Hemisphere, were presented in Michener (2000).

2.2 The Biology and Life History of Carpenter Bees

42

Large carpenter bees belong to the h·ibe Xylocopini within the subfamily Xylocopinae (Hymenoptera: Apidae). They are currently grouped into a single genus, Xylocopa (Minckley. 1998). The genus comprises at least three clades (Leys et al., 2002) and ca. 470 species (Michener, 1963). Carpenter bees occur in tropical and subtropical habitats around the world, and occasionally in temperate areas (Hurd and Moure , 1963).

Biogeographical analyses suggest that the genus probably has an Orientai-Palaearctic origin, and that its present world distribution results mainly from independent dispersal events (Leys eta/., 2002).As implied by their name, carpenter bees dig the ir nests in dead or decaying wood, except for the subgenus Proxylocopa that nests in the soil (Gottlieb eL

al. , 2005). The wood-nesting carpenter bees construct two main types of nests: (i) unbranched (also called linear), with tum1els extending in either one of both directions from the nest entrance. Linear nests are usual ly consh11cted in hollow or soft-centered

plant material, such as reeds; (i i) branched nests (> tunnels), usually constructed in h·ee trunks or timber (Gerling et al., 1989).The type of nest conshucted usually varies with

species, but some species show plasticity in nest architecture, depending on the nesting substrate available to them (Steen and Schwarz 2000). The nesting female lays one or a

few eggs along a tunnel during a brood cycle, provisions them, and constructs partitions of masticated wood to separate the offspring from one another. Maternal care in carpenter

bees also involves guarding of the immature offspring and feeding of the newly matured ones by trophallaxis (Gerling eta/. , 1981, 1983., Steen, 1978). In some species, helper females participate in offspring care rather than nesting independently, thus nesting can be social (see below). Some species are univoltine, whereas o thers produce more than one brood per year (Steen and Schwarz 2000). The activity season of carpenter bees spans 8-12 months, depending on species (Gerling et al 1983, Ben Mordechai eta!.. 1978,

Camillo and Garofa lo 1982, Camillo et a/., 1986)). Carpenter bees in temperate areas hibernate during the cold season (Steen and Schwarz 2000, Sugiura 1995), but emerge to forage on wam1 winter days (Gerling et al., 1983, Ben Mordechai eta!. , 1978). The mating behavior of carpenter bees been described for 38 species belonging to 16 subgenera (Leys and Hogendoorn 2008).

Variation in mating sh·ategies among subgenera has been recorded. In some subgenera,

males search for females at nesting sites, flowers, or landmarks (non-tenitoriality). ln others, they monopolize resources used by females, such as flowers or nesting sites (resource-based territoriali ty). Males may also monopolize areas lacking resources for

females (non-resource-based territories, or leks) Gerling eta!., 1989, Leys 2000).

A phylogenetic analysis suggests that resource defense is the ancestral state, and that this

mating system is conelated with low color dimorphism between males and females and a small size of the mesosomal pheromonal gland (Leys and Hogendoom 2008). Territorial

males chase away intruding males (Leys 2000, Sugiura 2008), which they identify by s ight and by the odor emitted from the intruders' mandibular g lands (Hefetz 1983). They also use a pheromone secreted from their mandibular gland to mark their territory (Hefetz 1983). When females enter the territories, males follow and try to mount them (Leys 2000, Rosenboim 1994). Observations of copulations in carpenter bees are extremely rare (Leys 2000) and were recorded only for a handful of species. In X varipuncta, matings take place in the non-resource territories {Alcock 1993), while in X sulcatipes and X jlavon(fa, they occur at high elevation during flight (Gerling eta/., 1983, Rosenboim, 1994, Watmouth, 1974).

2.3 Social Organization

Sociality, involving non egg-laying guard bees and a dominant egg-laying forager, has been described for ten species of Xylocopa. In nests of the African species X com bus/a,

fi rst eclos ing daughters remain in their natal nests and perform guarding duties while their mothers produce a second brood (Bonelli 1976.). Similarly. in nests of X pubescens sociality generally occurs after the emergence of the young, where either the mother is the reproductive and a daughter guards or vice versa (Hogendoorn and Leys 1993). Matrifilial nests of X virginica (comprised of a mother and her daughters) also show reproductive skew, and guard ing individuals become reproductive in the following year. In these nests, the mother perfonn s all nest maintenance, foraging, cell preparation and oviposition, whereas the younger inactive females only perforn1 guarding duties (Gerling and

Hennann 1978). Nests of X sulcatipes can be matrifilia1, composed of sisters, or involve the jo ining of unrelated females (Stark 1992). Some X sulcatipes nests are initially quasi social (no reproductive division of labor), but after a brief period of reproductive competition involving oophagy, a division of labor is usually established. Eventually most nests contain one reproductive and a guard (Stark eta/., 1990). The helping role of female offspring bas been suggested promote greater maternal investment in daughters

than in sons, leading to the female-biased sex ratio recorded in X sulcatipes (Stark 1992). In both X pubescens and X sulcatipes, the reproductive females produce 100% of the offspring while the guards produce none (Hogendoorn and Velthuis 1999). Nests of X sonorina also exhibit high reproductive skew, where the forager (mother) reproduces and feeds nestmates via trophallaxis, and additional females (daughters and/or joiners) shar

43

44

guarding duties (Gerling 1983.). For X ji-ontalis, X grisescens, and X suspecta matrifilial, semisocial, and communal nests have been recorded (Camillo and Garofalo 1989). Genetic analysis of X aeratus and X bombylans. which form multi-female nests during part of the breeding season, indicated the presence of multiple matrilines in

approximately 50% of nests. Socially nesting females were frequently sisters in one of the populations studied, and were often umelated in a second population. The results also indicated that temporary high reproductive skew occmTed in multi-female nests, that is, that different females were reproductive during different parts of the season (Steen 1978). Several ecological and life-histmy variables were suggested to promote social nesting in carpenter bees. Social living was found to con·elate with late season (Hogendoorn and. Veltbuis 1999) and older age (Hogendoorn and Leys 1993) in X pubescens, possibly because matriftlial nesting only occurs when mothers produce their second brood. Nest structure was proposed as an additional factor that affects social organization: in some species, females in branched nests build and provision separate tunnels at the same time, which can result in a communal social organization. In other species, females constmct one tunnel for the fiTst brood generation and only construct a new tunnel after the first brood has reached maturity. This can then result in eusocial nesting, where the daughters of the first generation assist their mother in building and provisioning subsequent tunnels (Steen and. Schwarz 2000). Finally, a period of reproductive inactivity of mature offspring was proposed as a n·ansition step toward social living. Such a period occurs in some solitary species (such as X frontalis and X grisescens), where newly emerged adult females remain in their natal nest for 20-30 days. During this time, they are provisioned by their mother or by their oldest sister, if the mother is absent. In some species, this association becomes permanent in a fraction of the nests (e.g., in X suspecta (Camillo et al., 1986), which then become social. Improved defense against parasites and predators has been suggested to favor the evolution of social nesting in bees (Smith et al., 2003)).

Carpenter bee nests are attacked by several types of natural enemies, including parasitoid wasps and flies, predatory wasps, ants, tem1ites, and insectivorous birds (Balduf 1962). However, in X pubescens, the frequency of parasitism did not differ between social and solitary nests (Hogendoom and Velthuis 1993). Thus the role of guards in reducing nest parasitism is not supported so far. The most extensive work on the consequences of sociality has been carried out for X pubescens.

In this species, the frequency of social nesting increases as the reproductive season progresses. It has been suggested that this increase has evolutionarily been imposed on females by shmtage in nesting s ites (Gerling eta f. , 1981 ). Social nesters spend more time foraging outside their nests as compared with sol itary individuals, perhaps because the presence of the guard in the nest reduces the risk of prolonged foraging (Hogendoom and

Velthuis 1995). Social nesters also suffer fewer nest takeovers by intruders than solitary nesters, providing a possible benefit for social nesting w hen competition for nests is high. The guards, in turn, may benefit from increased indirect fitness (if related to the reproductive), and increase their chances of eventually taking over the nest (Hogendoorn and Velthuis 1995). Thus, social organization can affect the fitness of X pubescens

fema les. Social and solitary nesters that foraged within a greenhouse differed in their food-plant preferences. Social females directed more of their foraging to a pollen source (Portulaca oleracea) than solitary nesters, possibly because of their higher brood production rates (Keasar eta/. , 2007).

2.4 Foraging Ecology

2.4.1. Abiotic Requirements for Foraging

Carpenter bees tolerate high ambient temperatures during foraging, and most species are inactive at low temperatures. For example, the lower activity temperature thresholds are

23 oC for X. capitata (Luow and Nicolson 1993), 21 oC for X. sulcatipes, and 18oC for X

pubescens (Gerling et a/., 1981 ). Flower visit rates in X olivieri are highest at a

combination ofhigh (25-35oC) temperatures and low (1- 100 Lux) illumination levels

(Gottlieb eta/., 2005). X ari::onensis individuals that foraged on Agave schottii together with honey bees and bumble bees were active mainly during the late moming hours, while honey bees and bumble bees were more crepuscular. These patterns were suggested to reflect low competitive ability, together w ith high thermal tolerance, in the carpenter

bees (Schaffer eta/., 1979). X varipuncta maintains fli ght activity within an ambient

temperature range of 12-40oC (Heinrich and Buchmann 1986). This heat tolerance

suggests good heat regulation ability in carpenter bees, possibly controlled by a thermoregulatory center in the prothorax (Volynchik et al 2006.). The activity period of some species, for example, X su/catipes, X cearensis, and X ordinaria, spans most of the

dayl ight hours (Viana eta!., 2002, Bernardino and Gaglianone 2008.). In other species (such as X pubescens, X taban(formis, and X olivieri), activity is crepuscular (Abrol

1987, Janzen 1964). A few species are nocturnal: X. tenuiscapa forages on its pollen host on moonless nights (Somanathan and Borges 2001 ), and X tranquebarica (Burgett et al

2005) has been observed foraging on moonlit nights.

2.4.2 Water Balance

Carpenter bees often ingest excess water during nectar foraging. Analysis of nectar consumed by X capitata showed that it is very concentrated. Nevertheless, their hemolymph is only moderately concentrated, and their urine is very dilute. This suggests

that ions, rather than water, may be limiting for carpenter bees (Nicolson and Luow 1982). This hypothesis is supported by the observation that bees often excrete water

45

before and during flight, and that they often engage in water evaporation from ingested

nectar (Willmer 1988). A similar excess of water ingestion, which leads to copious

excretion and evaporation of water, was described for X pubescens foraging on the nectar

of Callotropis. On the other hand, physiological water requirements are finely balanced

with the water contents of Callotropis nectar in the sympatric species X su/catipes, possibly due to extended coevolution with this plant (Willmer 1988).

2.4.3. Nectar Robbing

Nectar-foraging carpenter bees often perforate the corollas of long-tubed flowers, and

thereby reach the nectaries withou t contact with the anthers. Such "illegitimate pollination" or "nectar theft" has been reported for X virginica and X micans foraging on

blueberries. Nectarrobbing in blueberries may reach 100% of the vis its (Delaplane 1995)

and significantly reduces fruit set and seed number as compared with plants visited by

honey bees ((Dedej 2004 ), but see (Sampson et a l 2004)). Nectar robbing by carpenter

bees has also been observed in the wild plants Petrocoptis grandiflora (Guitian et al

1994), Fouquieria splendens (Scott et a/., 1993), G/echoma longituba (Zhang et a/., 2007), and Duranta erecta (Navano and Medel 2009). Corolla tube perforation

contributed to the reproductive success of the plants in P. grandiflora and F sp/endens, indicating that the nectar robbers were dusted with pollen during foraging, and functioned

as poll inators. In G. longituba and D. erecta, on the other hand, nectar robbing by

carpenter bees reduced seed set, as compared w ith plants visited by legitimate pollinators

(Guitian et al., 1994, Zhang eta!., 2007, Navarro and Medel 2009).

2.4.4. Food Sources

Carpenter bees in natural habitats are generalist nectar and pollen foragers. For example,

foraging X cearensis were recorded from 43 p lant species in Bahia, Brazil (Viana eta/., 2002), while X /atipes and X pubescens foraged on 30 species in India (Raju and Rao

2006.); In lsrael,X pubescens and X su/catipes used 61 species as forage plants (Gerling

eta/., 1983.); X danvini in the Pacific is known to visit the flowers of79 plant species

(Sugiura 2008.); 28 plant species provide nectar and pollen for X ordinaria in Brazil

(Bemardino and Gaglianone 2008). Carpenter bees can also be trained to collect sucrose

solution from feeders in experimental settings. In laboratory experiments, X micans were

able to discriminate between sucrose solutions that d iffered in mean volume (I versus 3

microl iter) and concentration (10% versus 30%). They were indifferent to variability in

both nectar volume and nectar sugar concentrations. This risk indifference was recorded if the bees were fed or starved (Perez and Waddington 1996).

2.5. Crop Plants Pollinated by Carpenter Bees

46

Carpenter bees pollinate passionflower (Passiflora spp.) in their native habitats (Mcguire

1999) and in commercial agricultural settings (Corbett and Willmer 1980, Roubik 1995,

Freitas and Oliveira 2003, de Siqueira et a/., 2009). They provide better pollination

service than honey bees for this crop (Roubik 1995). Xylocopa subgenus Lestis has been successfully reared in greenhouses for tomato pollination in Australia. Their foraging act ivity led to an increase in tomato weight by I 0% relative to a combination of w ind and insect po IIi nation. The efficiency of carpenter bees in pollinating tomatoes is increased by their abili ty to buzz the anthers (Hogendoorn eta/., 2000). In a pilot study in Israel, the fruit set of greenhouse-grown honeydew melons was three times higher when poll inated by X. pubescens compared to honey bee pollination (Sadeh eta/., 2007).

Social and solitary nesters bad similar efficiency in pollinating this crop: they did not differ in the daily activity patterns and flower visitation rates. Pollination by both types of nesters led to similar fruit sets, fruit mass, and fruit seed number (Keasar eta/., 2007). Carpenter bees are important pollinators of cotton in Pakistan, India, and Egypt

(Watmouth 1974). X varipuncta is compared favorably with honey bees (Apis mel/(j'era)

as pollinators of male-sterile cotton in field cages in the USA (Sadeh et a/., 2007). However, X pubescens in Israel did not provide satisfactory pollination of cotton for hybrid seed production (D. Wei/, personal communication). Finally, the night-flowering cactus Cereus repandus (syn. C. peruvian us) is pollinated by X pubescens in Israel (Weiss etall 994).

2.6. Domestication and Mass Rearing of Carpenter Bees for Pollination

A major obstacle to the commercial use of native polli nators in agriculture is the need to mass-rear them, rather than collect them from nature. Devising efficient and cost­effective mass-rearing protocols for X pubescens is a necessruy step in this direction. Attempts to mass-rear carpenter bees have focused on the construction of nest boxes that are placed in natural habitats to enhance nesting success.

Skaife ( 1952) constructed observation nests of bamboo tubes and transferred hibernating X ca.ffra into them. Most of the females remained in these nests after they exited hibemation. Oliviera and Freitas (2003) designed and tested nest boxes for X fi·ontalis,

based on the general design ofLangstroth honey bee hives. Each of nine wooden frames in these boxes was modified to serve as an independenL~y/ocopa nest.

Colonization rates of these boxes ranged from 19 to 52%, and the proportion of males in the emerging brood was 0.38. Efforts to develop protocols for captive mating and rearing of carpenter bees have so far met w ith limited success (unpublished results). The endocrine and molecular pathways that underl ie reproduction in carpenter bees are yet unknown. Elucidation of these pathways wi ll help identify the bottlenecks in the bees' reproduction, which may include overwintering of adults, mating, sperm storage and choice, nest construction and/or brood care. lnfom1ation on the potential reproductive

pitfalls, and their physiological mechanisms, is expected to facilitate the development of

.11"7

effective captive breeding methods forXylocopa.

2. 7 Future Prospects of Carpenter bees

48

Carpenter bees possess several advantages as potential crop pollinators compared to other non-Apis bees. Many solitary bees have a short activity season and/or are specialist foragers, and therefore do not provide a broad alternative to honey bee pollination. Carpenter bees, on the other hand, have long activity seasons and feed on a wide range of plant species. In addition, they are capable of buzz-pollination. This makes them potentially more versatile as agricultural pollinators. Hibernation occurs in the adult stage, and females start foraging whenever temperatures reach high enough values. This means that it is relatively easy to manipulate the onset of foraging in greenJwuses. Another important advantage is that the genus has a worldwide distribution. This implies that local species ofXylocopa can potentially be used over wide areas, reducing the need to import exotic pollinators. The possibility to lure these bees into suitable artificial nesting material allows provisioning of nesting material that can be easily used in agricultural settings and moved to places where pollination services are needed. In spite of higher per-capita pollination efficiency in some crops, carpenter bees are clearly inferior to honey bees in terms of pollinator work force, as they do not form large nests. Therefore they are expected to conhihute most to crop pollination when honey bees are ineffective. For example, the high termoregulatory ability of carpenter bees enables them to forage at higher ambient temperatures than honey bees. This makes them attractive candidates as pollinators in hot areas and in hot microclimates, such as in glass houses. The crepuscular and noctumal activity of some species may also allow them to pollinate night-flowering crops, which are not visited by honey bees.

Several problems remain in the management of carpenter bees for crop pollination, which call for further research. Most important is the need to develop an efficient captive breeding program for carpenter bees, which would include controlled selection of genotypes, mating, and nest founding. Such protocols have already been developed for other non-Apis pollinators, such as Osmia lignaria and Osmia cornuta. They include guidelines for nest construction and placement, overwintering and transportation of the bees. A complementary challenge is to enhance reproduction of wild Xylocopa populations, through provisioning of nesting material to their natural habitat. The availability of nesting resources was shown to correlate with the community sh·ucture of wild bees. Moreover, experimental enhancement of nest site availability has led to dramatic increases in wild populations of Osmia rufa. These fmdings suggest that Xylocopa populations, and the pollination services they provide, may also benefit from nest site enhancement in agro-ecosystems. Additional information about the pathogens and parasites of the genus is needed as well. A combination of ecological, physiological, and molecular genetic studies is likely to provide these essential data.

3.0 Bumble Bees

B umble bees are important pollinators of various crops like tomato, pepper, cucumber,

watermelon, cotton, kiwi and strawberries and various other crops. The use of hand

pollination in ceria in greenhouse crops is very expensive, so it is necessary to exploit the

bumb le bee species for pollination purposes, which can be managed easily in the greenhouse

than honey bees. ln several countries beekeepers and scientists are looking for other pol linating

insects like bumble bees as an alternate pollinator. Economic aspects of the use ofbumble bees

as pollinators for many agricultural crops are nowadays receiving more attention. Bumble bees

are commercially used in Netherland and Belgium from 1987. Commercial bumble bee colonies

have been available in US since mid-1992 and arc now utilized by most tomato greenhouse

growers (Kuencman, 1995). Jany ( 1950) reported that the piercing of scarlet runner beans flower

by bumble bees helped other chief insect visitors like honey bee and cabbage butterflies to access

nectar of the flowers. If the bumble bee population found in large number then the maximum

ferii lization took place in beans and seed development was not affected by the formation ofholes.

Bumble bees are effective pollinators for tomato in greeenhouses, which were earlier pollinated

by vibrat ing the tomato trusses (van Koot and van Ravestijin, 1962 and Picken, 1984). The

mechanical method was an expensive and time intensive practice compared with pollination by

bumble bees (Banda and Paxton, 1991; Kevan eta/., 1991 ). Ravestijn ( 1987) placed artificially

reared bumble bee colonies in tomato greenhouse for pollination in Belgium. Banda ( 1990)

reported that bumble bees were more effective than honey bees or hand pollination as determined

by fruit set, fruit s ize, weight and number of seeds per frui t in tomato in Europe. Asada ( 1997)

utilized the Japanese native bumble bees for tomato pollination. Fruit set was 84-90% more by

using bumble bees and very few fruits (7-10 %) were found inferior. In New Zealand, trapped

nested colonies of long tongued bumble bees were used for pollinating tetraploid red clover,

Trifolium pratense L. during early bloom (Macfarlane et a/., 1983). They estimated that 24

colon ies of B. hortorum each with a queen and about 35 workers placed along one side of red

clover maximized seed production. Short tongued bumble bees were found effective pollinators

of crane beiTies, Vaccinium sp. (Macfarlane and Patten, 1997).

3.1 The Distribution and Diversity

The distribution of bumble bee fauna is still poorly understood and much needs to be

known. The documented infonnation is rather fragmentaty and far from complete. The

Indian species of bumbus has generally been restricted to higher elevations especially Himalayan ranges. Bingham (1897) listed 24 species of bumblebees from higher

elevations of Kashmir, Himachal throligh Sikkim and Assam. Mani (1962) listed fou r

species of bumble bees at e levations over 4000m at Himalayas. William ( 1991) recorded

28 species of bumble bees from Kashmir I Iimalayas. Abrol (1998) has recorded Bombus

haemorrhoidalis from intermediate areas of Jammu range. More than 300 species of

49

bumble bees have been identified from north temperate zones extending through Europe, Asia and North America. The bumble bee fauna has extensively studied in several counh·ies of the world which include France, Japan, Germany, Korea, Italy, U.K, Canada, Sweden, U.S.A, Newzealand, China and Finland. More than 25 species have been recorded from U.K (Prys-Jones and Corbet, 1987). Chang-Whan and Ito (1987)

identified 212 species ofbumble bees and 5 species of parasitic bumble bees (Psithyrus) from Korean pen insula. Bumble bee fauna of central Italy has been reported to consist of9

species of bumble bees and three species of Psithyrus (Intoppa and Pace, 1983). In a similar study, Ito (1985) recorded 15 species ofbumble bees and one species of Psithyrus

from North Korea. Teras ( 1985) reported the occunence of 12 species of bumble bees in southern Finland. In these countries much work has been done on their management and utilization for crop pollination (Hobbs et. a!. , 1962; Free, 1963). Roseler ( 1977) devised methods for rearing bumble bees in green houses and developed many suitable domiciles. Griffm eta!., ( 1990) developed techn iques for commercial rearing of Bomb us ter6stris

(L.); B. ruderatus(Fabr.) and B. subterraneus to match their emergence with flowering phenology of the crop. Several investigators have designed nest boxes of wood or polystyrene forrearing of bumble bees (Haemert eta/., 1990; Eijude, eta/. , 1991; Ptacek, 1991 ). However, the success rate varied from species Lo species.

3.2 Effectiveness as Pollinator

50

Honey bees were found to visit on blackcurrant (Ribes nigrum ), raspberry (Rubus idea us)

and strawberry (Fragria ananassa), but bumble bee queens were found only on black currant. Bumble bees spent less time on each flower and visited more flowers per bush than honey bees. Honey bees were numerous on raspberry than bumble bees, but bumble bee collected propmtionally more pollen. Only few bumble bee found to visit strawberry (Free, 1968). However Poulsen ( 1973) observed more number of bwnble bees visiting

field beans, working more rapidly as compared to honey bees. Honey bee workers and bumble bee queens were found to remove and deposit different amount of pollen in apple, because of different foraging behavior. Apple stamens restricted access to the ncctaries by shmt tongued Apis pa11icularly on Delicious group, because bees collected nectar side ways without contact with stigma. In contrast, Bombus queen approached the flowers from above, landing directly on the anthers and stigmas (Me Gregor, 1976). Bumble bees have long been suggested as useful alternative to honey bees in pollination under

greenhouse conditions where they do not appear to become easily disoriented as honey bees (Free, 1970). Richard and Pomeroy (1989) studied the poll ination of greenhouse

muskmelon, Cucumis melo by bumble bees in New Zealand. They observed lhat bumble bees foraged on melon flowers from dawn until dusk. Flowers pollinated by bumble bees attained 90% fruit with exportable weight. Ravestijn and Kraemer (1991) compared the

pollinating efficiency of honey bees and bumble bees on melon and found that fruit

l

numbers and weight/m1 area were slightly greater with bumble bees pollination than with

honey bees. Most pears are self unfruitful and need insect vector for cross pollination. Mayer et a/. ( 1987) found that in the Pacific Northwest, Bartlett and all other pear varieties require cross pollination. Pear blooms before over wintered queens of most bumble bee species have begun to rear their workers. Therefore, few bumble bee foragers were observed on pear flowers. The number of honey bees foraging pear bloom at the

same time was higher than the bumble bees. Free ( 1993) suggested that more bumble bees will forage at low temperature at which honey bees activity is limited for pear pollination. In Meditenanean coastal regions malformed fruits production of strawberries was prevented by bee-pollination method (Ahnet a/., 1988), but unfortunately honey bees do

not work on cold days when temperature was lower than l"2°C. Pinzauti ( 1993) suggested that grower should introduce bumble bees to their greenhouses especia lly in winter months to get early, high yield, large and regular shaped strawberries. Paydas eta/. (2000) reported both honey bees and bumble bees to be effective in early production of strawberries. In this respect bumble bees were generally more efficient in pollination in cold weather conditions. The effectiveness of honey bee and bumble bee pollination on fru it set and abortion of cucumber and' watermelon was studied by Stanghellini eta/.

( 1997). They revealed that cucumber flowers visited by Bombus impatiens had lower per cent fruit abmtion thanApis mellifera, when compared at equal bee numbers. Stanghellini

eta/. (2002) used the commercial bumble bee colonies as back up pollinators for honey bees to produce cucumber and watennelon in large quantities. Several poll inators have been tested on Capsicum annuum under greenhouse conditions. B. impatiens was foundn to be well established for sweet pepper pollination (Shipp et a/. 1994; Miesels and Chiasson, 1997). Arnon and Kammer (2001) compared the effectiveness of bumble bees and honey bees on sweet pepper pollination under greenhouse conditions in Europe and Israel. They found that the average yield in honey bee plot (22.6 kg) was similar to the bumble bee plot (23.4 kg). In Mediten·anean countries, greenhouses are not regularly heated and pollen production and its quality are declined by low temperature and it brings the fruit set problems in egg plant. Abak eta/. ( L 997) studied the effect of bumble bee and

vibrations on the yield and frui t characteristics of egg plants in unheated greenhouses. Yield increased by 25% using bumble bee pollination during three years of experimentation. Bumble bee pollinated fru its also increased by 14% in weight and 7% in length as compared to vibration pollination. Bumble bees were fou nd to be efficient pollinators of flowers in the unheated greenhouses dur ing the winter and spring months in

the Mediterranean coastal region. Dasgan eta/. (1999) investigated the effectiveness of bumble bees as an alternative pollinator for melon plants in Turkey. Fruit weight, fruit height, fruit diameter and number of seeds per fruit of bumble bee pollinated plants were significantly higher than honey bee pollinated. The yields obtained were found sin1ilar

(6kg/m2) by both these pollinators, while the average fruit weight (1 166 and 991 g), height

51

(15.42 and 12.63 em) and number of seeds (628 and 579 seeds/fruit) were higher in

bumble bee pollinated plants than honey bees. Zaitoun eta/. (2006) compared the effect

of honey bees and bumble bees on the length and sugar content in strawberry. The plants

pollinated by bumble bees had fruits with higher fruit length (4.2 em), diameter (4.1 em)

and volume (269.3 cm3) and per cent sugar content (7.8 %), than the honey bees pollinated

(fruit length 3.9 em, diameter3.7 em, volume 227.9 cm' and percent sugar content 7.6 %).

3.3 Bumble Bee ForagingActivity

52

The foraging activity of B. dahlbomii and B. terrestris was studied in native and non­

native vegetation in Chile by Ruz and Herrera, ( 1980 ). They observed B. dah/bomii in January, with very active colonies whereas the B. terrestris was found first time in wild

during April-May. B. terrestris was observed to forage faster than B. dahlbomii and other

native bees. Therefore, in long tenn it may cause impact over slower forager species while

exploiting the same resource. Free ( 1955) studied the foraging behaviour of four species

of bumble bees living in artifi cial nest boxes in E ngland. He found that the foraging

population showed slight peak at I 0-11 a.m. and a tendency to collect pollen load

increased during the day. About 15% of the foragers spent the night away from the nest.

He also worked on the division of labour in artificial boxes and found that half of the

workers had no specific duties, but 2/3nJ were either foragers or house bees. The amount of

pollen collected was related to the amount of brood in the nest. Paarmann (1977)

compared the activity ofhoney bees and bumble bees on morel Ia cheny Honey bees were

found to be more active around noon but bumble bees in the evening hours, however,

during cold days honey bees did not fly out whereas bumble bees were more active in the

aftemoons. Willmer et a!., (1994) studied the behaviour and activity pattems of Apis

mellifera and of 5 species of Bombus on raspberry in Scotland. They found that Bombus spp. favoured young flowers strongly, early in morning when pollen was more abundant

while A. mell(fera visited unselectively. Bombus spp. carried more pollen on their bodies

than A. meLI?fera and deposited more pollen on raspberry stigmas. Bumble bees also

foraged substantially for longer periods of the day in poorer weather. Asada and Ono

(1996) classified foraging behavior of bumble bee in four categmies viz. buzzing on

flower and hanging on anther, grooming pollen grains adhering to leg hairs, while

hanging on anther, flight from flower to flower and flight towards the cage and resting.

Abak and Guier ( 1994) reported that pollen amount and fertility of greenhouse egg plant

were generally lower in winter, but if the effective pollinators like bumble bees were used,

cultivars are able to set the fruit. Bumble bees were found to forage from 7:00 to 14:00

hours in January, 6:00 to 18:00 hours in February and March on sunny days. Bumble bee

pollinated flowers gave higher fruit yield (25%) as compared to vibration pollination.

Fruit size also increased 14% in weight and 7% in length and number of seeds per fruit

was higher in bumble bee poll ina ted fruits than from vibration pollination.

j

A B

c D

Bumble bee working on A, B ) citrus flowers C) cucurbit and D) brinjal flowers E) clovers, F) thistle

3.4 Foraging Activity on Different Crops Grown in Polyhouse

Fisher and Pomeroy (1989) recorded the bumble bee activity in greenhouse condition on melon plant in England. They observed that bees were entering and leaving the each hive during 5 minute period at 2 hour intervals from 0600 to 1800 hours iJl December. The numbers of male and hennaphrodite flowers visited by bees were also recorded. Ravestijin (1991) used bumble bee, Bombus terrestris for pollination of glasshouse

tomato inNaaldwizk. Flowering started on an average on 1 •' February± 7 days and bumble

bee colonies were placed in the glasshouse on 31 '' Janua1y. He found that the one active

btunble bee worker may pollinate at least 500 plants/day i.e. 250 m2 area of glasshouse. Therefore, use of 10-15 colonies/hac. were seemed more than sufficient to ensure effective pollination. Hive traffic and greenhouse foraging of Bombus impatiens was recorded on sweet peppers (Capsicum annuum L.) in Canada (Meisels and Chiasson, 1997). B. impatiens flight activity was measured every third day from 29 June to 13 July at 2 hour intervals from 0700 to 1500 hours. Hive traffic was found higher during the first 3 days from 29 June to 13 July and greenhouse foraging was greatest during the first 4 days from 29 June to 13 July. Abak eta I. ( 1997) observed the bumble bee, (Bombus terrestris) activity on egg plants grown in unheated greenhouse in Turkey. They found that bumble bees start foraging from 0700 to 1400 hours in January, 0600 to 1800 hours in Febmary and March. However, the peak activity was observed between 0900 to 1100 hours and then decreased gradually. The flight activity stopped between 1300 to 1400 hours and started again in aftemoon between 1500 to 1800 hour

3.5 Status of Pollination and Domestication Research in India

54

Studies conducted on the domiciliation of bumble bee (Bombus sp.) and their resource partitioning with honey bees are summarized here under. The main flora sustaining the queen population was identified under Nauni conditions. The queen can be directly trapped from Salvia moorcroftiana, Rosmarilts officina/is, Lavendula sp. and wild plants like Caryopteris bicolor, Scutelleria linearis. Studies on the rearing of bumble bee revealed that colonies can be initiated after trapping queens from these plants during moming and evening hours. Domestication of bumble bee was successfully done under

controlled conditions maintaining the temperature of 25-30°C and 60-65% relative humidity and by feeding them sucrose solution (50%) and corbicular pollen. Fifteen colonies were successfully reared with the range of population 80-100 individuals during 2006-07 continuously for 7-9 months. The best nesting cage was wooden box and for feeding bottle lid was easily accepted. Nest architecture of the artificial reared colonies were studied in two chambered wooden box and hoarding cages. All the nests contaiJ1ed number of cell layer and brood cells in the centre, where as number of honey pots were mostly arranged in cluster near the nest periphery. Foraging activity of three natural nests were recorded and found peaked in the morning hours. Bumble bee start foraging activity

from dawn to dusk (0630-1730 hours) period. Therefore bumble bee has long working hours. Time spent outside by the marked bees ranged from 10.83-21.33 min./bumble bee. The study revealed that the forager dose not exhibit different foraging trip after leaving the nest during different day hours.Natural nests were digged out after ceasing the nest activity. All these nests were built underground inside the abandoned site of rodents burrows and found attached to the root of adjoining tree or shrub. Nests were found to be interwoven with grass, moss, wooden pieces and leaves beneath which brood cells lies. The study revealed that bumble bee prefers darker and corner side of the ground and make involucrum to protect the nest from outside env ironment. Comparative foraging activity

of bumble bee and honey bee on the field grown cucumber, capsicum was recorded similar, but in bitter gourd bumble bee was found higher in number than honey bees. On pumpkin only bumble bee was observed to visiting the bloom shows its complete dominance. In sweet cherry bumble bees were playing an important role to pollinate the crop, where no honey bee was found visiting the bloom in Kurnarsain, Shim Ia district. A

new species B. assam en sis Bingham was reported for the first time visiting apple bloom at an altitude of2725 m above mean sea level in Narkanda, Shimla district. One attificially reared colony was kept inside the polyhouse at vegetable field on cucumber crop. Colony was kept without providing any additional food inside the rearing box. Bumble bee start visiting the flowers from 0600-1900 hours and found better acclimatized with controlled environment. Therefore it can act as an alternate source of pollinator inside polyhouse where the honey bee activity is limited. Solan area of Himachal Pradesh has thus exhibited diverse wild bee fauna like bumble bee. Because of the importance ofthese bees as crop pollination and the recent outbreak of Varroa to honey bees industry effort should be made to profitably manage their population through manipulations so that these bees may adjunctively work with the hive bees for enhancing the crop production.

3.6 Domestication

In present study queens trapped from the flowers in spring were kept singly in two

chambered boxes and hoarding cages maintained at temperature 25-30°C and 65-70% relative humidity. The queens were fed w ith sucrose solution (50%) and corbicular pollen.

Macfarlane eta!. (1990), have also reared the spring collected queens kept in two screen cages and fed with 50% sugar solution and ground corbicular pollen by maintaining at 18-

250C temperature. Most of the workers have reported the successful domestication of

queens collected during spring. Plowright and Jay (1966) have reared the hibernated

queens in plastic cages at 30°C temperature and 60% relative humidity. Frison (1927) found a greater incidence of colony formation in boxes containing a single queen than housing pair of queens, close to the present observations where the queens were kept singly inside the boxes. Similarly, Ptacek ( 1985) has reared the bumble bee in wooden boxes and fed with 60% honey solution and pollen dough made of corbicular pollen. Ono

56

eta!. , (1994) reared Bombus hypocrita and B. ignitus in laboratory from post hibernating queens. Queens were kept in wooden boxes consisting of brood chamber and feeding chamber and were supplied with sucrose solution (50%) and fresh pollen pellets.

Fifteen colonies were successfully domesticated for 7-9 months during 2006-07. In the present study, the average developmental time in B. haemorrhoidalis from egg (wax secretion) to adult was found in the range of 23.7 to 30 days. These fmdings are in confonnity with Mah and Bilinski, (2001) who found the average developmental time in B. ardens ardens, B. hypocrita sapporoensis and B. ignitus from egg to adult being 28, 27 and 28 days, respectively. Similar study was conducted by Gonzalez eta!. (2004), who reported 29.6 days to be the average developmental time in B. atratus. Yeninar and Kaftanoglu (1997) repmted three successive phases in colony development that is colony initiation, switch point and competition. They found that dming colony initiation queen makes egg cups, lays diploid eggs and start producing worker bee. With the first worker emergence, worker assists the queen in feeding. Switch point occuned on an average 18.9±1.3 days after the first worker bee emerged and competition point occurred at an average of 31. 8± 10 days after the emergence of first worker bee. Similarly, in the present study queen started to make egg cups, laid diploid eggs and start producing workers after 23±2 days of wax secretion.

Several workers have given various nest box designs for rearing the colonies in captivity. Macfarlane et al. ( 1983) found that box can be made from any type of material (plywood, concrete, polysterene, plastic), but should be constructed to allow for the escape of water vapours as excessive humidity makes colonies susceptible to mould. Rosier and Jay ( 1966) used different methods of rearing Bombus sp. to induce the colony in captivity. In

series I queens were kept in container consisting wax paper carton, but none of the eggs laid survived for more than few days. In series II queens were installed in a wooden container consisting of two boxes with glass roof and conugated floor. Queens built their egg cells directly on the cardboard floor of the outer box. During the development of first brood, some queens neglected larvae because of wax envelope surrounding them. The design of series III was similar to series II, but divided into two unequal parts. The larger chamber contained feeding tube and was lined with corrugated cardboard and had a glass roof and smaller chamber was lined with upholsters cotton. This design was more suited for rearing the queens in captivity. On the other hand, queens which had conshucted their first egg cells upon the pollen lump rarely built more cells upon the floor. In the present study two chambered wooden box used for rearing were found similar to Rosier and Jay (1966), but with out lined conugated floor and more number of queens were found to construct their egg cells upon the pollen lump than the floor of the box.

3.7 Advantages

Bumblebees are good pollinators of the crops for the following reasons:

The bumblebee is capable of vibrating the flower using the unique "buzz pollination" mechanism. The tomato flower needs vibration for proper pollination and fruit set under greenhouse conditions. The bumblebee does this in an optimal way far superior to other methods. The bumblebee is less affected by extreme weather conditions than the honeybee.

Bumble bees are cool weather operators. Unlike honey bees, bumblebees are active at low temperatures (5°C), in windy conditions and under cloudy skies. Their bodies have an interesting adaptation that assists them in being flight functional in cool temperatures when other insects cannot fly. Their thorax is almost always totally or pmiially black and often "bald". The black color absorbs heat quickly and warms up the flight muscles allowing them to fly after only being exposed to sunlight for a short time.

The bumblebee is better adapted to perfonn under confmed greenhouse conditions. Bw11blebees are not only excellent pollinators in open air, but are especially valuable in greenhouses and plastic tunnels .The bumblebee is less inclined to look for alternative sources of pollen and nectar outside the greenhouse. Therefore, it will stay in the greenhouse even if the latter is opened for ventilation purposes. Many species have longer tongues than honeybees, so they can pollinate flowers with long, nanow corollas. They are very hairy and their hairs are branched and so are perfect for picking up and transfening pollen. Bumblebees can completely replace manual pollination and use of hmmones, resulting in less labour costs. Higher fruit production and quality: In crops, such as tomatoes, peppers and blueberries, bumblebee pollination results in higher yield as well as larger and higher quality fruits.

51

· 4.0 Leaf Cutting Bees

H oney bees continue to provide generally satisfactory and frequently excellent pollination of most of our introduced flowering crops. A few crops exhibiting obviously specialized pollinator needs, such as Lucerne and perhaps red clover, have

been catered for by the introduction of specialist bees. Where numbers of these bees have been adequate, crop yields have increased. Alfalfa, Medicago sativa (L), is a high quality forage and green manure crop that originated in the Middle East. Alfalfa is largely sclf-fe1tile, but for mechanical reasons, flowers require bee visitation for pollination. Solitary bees and bumblebees are the most efficient pollinators of alfalfa. Honey bee efficiency, on the other hand, is low after opening alfalfa flowers several times. Iloney bee "learns" to collect nectar without tripping flowers, due to the specific structure of the alfalfa flower. For that reason, despite the abundance of honey bee in alfalfa fields, seed yield per hectare may be very poor when solitary bees and bumble bees are not present. The problem was successfully overcome for the first time in USA and Canada with the domestication and utilization of the solitary bee Megachile rotundata (Fabricius, 1793). Alfalfa flowers require visiting bees to trip the sexual column, thereby providing pollination and subsequent pod and seed set. However, tripping is done by a specialized group of bees, which enter the flowers and press their keel by their own weight by releasing male and female organs to distribute pollen and effect cross-pollination. Much reliance was thus placed on the solitary bees which were thought to have greater adaptation to the floral structure of the flower. Constant effmts led to the establishment of two solitary bees- Nomia melcmderi and Megachile rotundata. Between the two, the latter was proven to be more efficient pollinator besides having gregarious nesting habit, a point highly useful towards harnessing it in artificial domiciles. Stephen ( 1961) and Bohart ( 1962) have been able to develop its harnessing methodology using corrugated paper paper straws and dri lied wooden blocks.

4.1 Nesting biology

In India, around Hissar (29".1 "N (latitude) and 75°.46 E (longitude), a group of four megachilid bees Megachile hmJ'anaensis Rahman (ex. M. nana Bingh)., Chalicodoma rubripes (Morawitz.) (ex. Megachile jlavipes Spinola)., Chalicodoma lanata F. (ex. Megachile /a nata Lepel) and Chalicodoma cephalotes Smith (ex. Megachile cephalotes Smith) were found. They construct nests in pre-tunneled channels. These characteristics make them also suitable for hamessing and hence for organized pollination of alfalfa.

Castor leaf stalks or stems having tmmel diameters, ranging from 2-11 mm and 10, 15 and 20 em in length with one end closed were tied into bundles of 10 each and were hung at a location close to the crop area in October by Kapil and his co-workers. The bees started accepting them within a week. In March-April the operation was again repeated by hanging the bundles inside a hut specifically constructed for the purpose in the field facing north east. In addition to the bundles of castor stems new domiciles drilled wooden

blocks and grooved particle boards having 6 mm diameter tunnels were added to the a1tificial nesting site in February before the bees started post-winter activity. Some tunnels had paper straws and others were without. The experiments to detem1ine the efficiency of bees was carried out in field cages made of nylon netting (24 mesh) supported on angle-iron frames.

The data given in table 3 explain the acceptance of artificial nesting materials by the megachilid bees. In October-November, the acceptance although was poor, probably for reasons of short period of bee activity subsequent increases in acceptance were observed

from March-April. MhaJ)'Cmaensis had shown by comparison more preference for the castor stem tunnels over the mason bees was much higher in the field their visitation on the nesting site was surprisingly low. The addition of soda straws to the artificial nesting complex in March, attracted a large number of bees to nest in them. Among them, M.hmyanaensis had an overall lead in accepting the devices and showed preference for straws over the castor stem tunnels the acceptability being 77.0 per cent in soda straws and 32.4 per cent in castor stem tunnel. Analysis of acceptance of the castor stem tunnel length by the bees indicate that the bees generally prefen·ed shorter tunnels. M. hmyanaensis liked to nest more in tunnels having intemal diameter between 3.5 to 6.0 mm with major concentration between 4.0-5.5 mm as indicated by the frequency of acceptance in this range of the tunnel diameter. Whereas the mason bees preferred

between 4.5 to 8.0 1mn. The acceptance of the tunnel d iameter seems to be determined by the size ofthe bees' thorax.

During brisk activity period the leaf cutter bee constructed between 7.5 and 14.0 cells per nest during march-May, in different lengths of tunnels provided, which appears affecting

the length and diameter of the cells. The number of cells constructed during other seasons dropped by one-third. The seasonal effects were not, however, seen on the length of cells,

although some variations in diameter were recorded. Compared to castor stem tunnels more straw tunnels were accepted (table 4) during the same period. This suggests the preference of the bee for tunnels with homogenous tunnel rather than of castors having irregular tunnel d iameter. The space used for closing the tunnel was also seasonally influenced.

Because few mason bees visited the nesting site, not much information could be collected.

M . .flavipes preferred relatively wider tunnels with diameter close to 6.8 mrn, M. cephalotes constructed nests in 5.6 mm tunnels and M femora fa in still nan·ow tunnels

havi ng intemal diameter about 5.1 mm. The rate of cell construction remained unaffected: for example M.jlavipes made 3.8 cells/nest, M.femorata made 3.3 and M. cephalotes 3.7 cells. The cells of M. cephalotes measured 13.8 mm and those of Mflm'ipes 13.4 mm,

but they also used much space for closing the tUlmels thanM..flavipes and M.femorata.

4.2 Impact on seed setting

M. hcnyanaensis and M . .femorata were shown to be the most effective pollinators of

alfalfa. Kapil eta!. 1975 also concluded similarly from the rate of tripping. These bees

tripped 87.4 and 89.7 per cent respectively of the flowers visi ted by them while M . .flavipes tripped 82.4 per cent flowers. M. lzcnyanaensis as explained in the Table 4 also

competes well with the most popular exotic bee pollinator M. rotundata. An experiment

conducted on the effect of !vf. haryanaensis and M. flavipes on seed setting in which a

known number of bees were released in cages demonstrated the superiority of M hw)'alwensis over M.Jlavipes. But out of all bees the population of M . .flavipes available in the field at the time of blooming was the highest . Table 6 explains the impact of the

management of these bees ncar the alfalfa field at Hisar on seed setting. With constant

addition of artificial devices and an obvious increase in the population of bees, an increase

in seed production was observed through open pollination. In first year of study, the seed

production was 273 kg/ha and in next year it increased further to 370 kg/ha with

maximum yield being 480 kg/ha during the same year. ln the subsequent years, the yield

was still better as assessed from setting and naked eye assessment but the seeds did not

nature due to untimely rains.

4.3 Conclusion

en

Artificial hives are recommended to fmmers for many reasons: easy to handle, storage

and reuse. In addition, using hives could help for keeping bees in nature and saving them

from disappearance. Using and conserving wild solitary bees will increase alfalfa seed

production and also the productivity of several other crops.

Table 3: Acceptance of artificial nesting materials by megachilid bees

Year of Period of Materials No. No. of Tunnels accepted by bees

study nesting Provided Provided M. M.flavipes C. Total haryanaensis andM. cephalotes

femorata

1st year October- Castors 110 8 0 08

November [I.D.4-8mm]

March-April -do- 200 15 40 2479

April-May -do- 800 76 40 16132

July- August -do- - 87 - 18105

2oc~ year April -do- 968 314 56 53423

April Straws 700 539 57 15611

[I.D. 6mm]

*At natural nesting site

Table 4:Percentage acceptability of different kinds of nesting materials and the rate of cell construction by M. haryanaensis

Material No. No. Rate of acceptance* Av. Cell Nest provided nests nest April May May June Cells/ length/nest closing

provided accepted 20 5 20 5 nests [mm]

Castors 768 252 4.9 18.5 28.5 32.8 5.5 9.9 6.1 [1.0. 5-7 mm]

Straws 700 539 14.1 37.5 58.7 77.0 6.1 9.0 6.2

[I.D.6mm]

*Figures given are the percent acceptance of the total number provided.

61

5.0 Alkali Bees I o

I t is a highly gregarious solitary bee that nests in large numbers in saline soils with a silt loam or fine sandy loam texture. A solita1y species is one in which the female prepares and provisions the cell, deposits the egg, and then seals the cell completely unassisted. More than

one cell may be constructed, but only one at a time. After the cell is sealed, no further attention is given it, and the adult may die within a few days. observed that wild bee populations actually increased at least in the eastem half of the Un ited States because of i) opening up of forested areas, which created more favourable conditions for bees, ii) paving highways, which concentrated moisture along roadsides, iii) introduction ofweedsupon which the bees forage, iv) growing numerous crops upon which the bees forage and v) bringing desert areas into bloom (with irrigation). The world's only intensively managed ground-nesting bee, the alkali bee

(Nomia me/anderi Cockerell), has been used for >50 years as an effective pollinator of alfalfa (Medicagosativa L.) grownforsecd in the western USA(Cane, 2007).

5.1 Important Species of Nomia

The alkali bee, Nonzia spp. (Hymenoptera: Halictidae) has been known for many years to be a highly efficient and effective pollinator of alfalfa, particularly in western countries of the world where these bees are used in large-scale pollination oflegume crops.

• melanderi

• capitata

• oxybeloides

• angustitibialis

• fedorensis

• foxii

• howardi

• manee1

• nortoni

• robinsoni

• tetra:,onata

• universitatis

• nasi can a

5.2 Life Cycle and Habits

Alkali bees are nearly as large as honey bees. They are black, with iridescent copper-green stripes across the abdomen. The male bee has much larger antennae than the female. Gregarious bees are solitary individuals that endeavour to nest in close proximity to each other and alkali bee (Nomiamelanderi) belongs to this category. The adult bees emerge from late Jtme to late July, depending upon the location and season. The males appear a few days aheadofthe females. Before emergence, each bee is confined to its natal cell for 3 days as an egg, 8 days as a growing larva, 10 months as a full grown dormant larva, 2 weeks as a pupa, and several days as a hardening, maturing adult. During the approximate 1 month ofher active adult life, the female constm cts, provisions, and lays an

egg in each of 15 to 20 cells(Johansen & Mayer, 1982).

Mating occurs during the 3 days the entrance tunnel is under construction, usually during

the first day. The males patrol back and forth over the nesting site, and they will mate with any number of females; however, they rarely bother a mated female after she becomes

actively engaged in constructing the nest. About the third day after construction starts, the first cell is completed. Pollen is then collected and formed into a pellet in the cell, an egg is

laid on the pollen, and the cell is immediately scaled by a spiral ceiling and a soi l plug. After that work is started on the next cell, and no further attention is paid to the last one. Thereafter, the daily routine consists of fashioning another cell off the main tunnel, providing it with a pollen ball, depositing the egg and sealing the cell. About one cell is completed each day (Bohart and Cross, 1955). Usually only one nest is prepared and provisioned by a female. There is usually only one generation a year but two and sometimes three generations appear.

It bui Ids individual nests in the ground as many as 1 00 nests per square foot of soil. Being gregarious, alkali bees may constmct 100,000 or more nests in an area 40 x 50 feet. Nesting sites w ith an estimated 200,000 nests have been reported by Bohatt, 1952. The nest, a 10 nun (0.4 inch) vertical tunnel, may extend 10 inches below the surface but is

usually only 3 to 5 inches deep. There may be 15 to 20 cells usually arranged in a single comb-shaped cluster. Each cell is an oval cavity, slightly larger than the main tunnel, about one-half inch long, lined frrst with soil and then with a waterproof transparent liquid applied with the bee's glossa. Each cell is provisioned with a 1.5to 2 mm oval pollen ball, made up of8 to 10 bee loads of pollen mixed with nectar. The soil removed from the tunnel is dumped at the tunnel entrance to form a conical mound 2 to 3 inches across.

5.3 Bee Forages and Feeding Characteristics

Alfa lfa nectar and pollen constitute the primary source of food for most female alkali bees. They visit a few other plant species, for example, clovers, mint, onions, Russian thistle,

salt cedar, and sweet clovers. In alfalfa seed producing areas, however, most of the nests

63

are provisioned with nectar-moistened pollen balls derived ffom alfalfa(Johansen& Mayer, 1982). While foraging, alkali bees do not trip the alfalfa blossoms as rapidly as do the Jeafcutter bees, but almost every blossom they visit is tripped. Because females visit large number of flowers, they become highly effective. The males visit flowers for nectar only and only occasionally trip the flowers.The time of day that wild bees forage differs with the species involved. Those that feed only at dawn are referred to as matinal bees. Crepuscular bees feed both at dawn and near dusk. A few species are nocturnal in their foraging, but the great majority feed when the sun is shining, because that is when the majority of the flowers are open.

An alkali bee female trips at least 95 per cent of the luceme flowers as she gathers pollen for her nest. Under good conditions, each female wi ll trip about25,000 lucerne flowers

during her lifetime which translates to I /5- Y2 lb (91 -227 g)of clean seed. Females tend to forage within 1 mile (1.6 km) of the nest, altboughthey have been found up to 7 miles (11 km) from the nest( Johansen& Mayer, 1982). Kulkarni and Dhanorkar ( 1998) rep01ted that honey bees as main pollinating agents inniger fo llowed by sol itary, alkali and carpenter

bees.

5.4 Nesting Sites or Beds

nA

Although alkali bees are solitary, individuals make nests near each other (W1ight, 1997).

There are certain basic requirements of an acceptable bed. It must have a moisture supply capable of rising to the surface. This usually requires a hardpan layer a foot or more below a porous soil that tends to hold the moisture and permits its movement from the source of supply to the surface. Conditions should pennitrapid drainage of surface water. The under layer should range in texture from a silt loam to a sandy loam with no more than 7 percent clay-size particles. The surface should be finn but not have a hard crust. If some salt does not appear on the surface, about 1 pound of salt per square foot of surface should be raked into the flrst 2 inches. This seals the surface layer and thus slows down evaporation.

The bed should be kept relatively free of weeds. It should not be nooded during the active bee season or excessively disturbed by livestock or vehicles. When bee beds are constructed by alfalfa seed growers. about 3 feet of soil is removed from the selected si te.

The flat-bottomed excavation is then lined with 0.006-inch plastic film. The excavation is backfilled with an inch of soil, a 1 0-inch layer of gravel. and 2 feet of appropriate soi I. Salt is usually added to the surface as mentioned above. Water can be supplied through a piece ofti le that extends ffom the gravel bed to several inches above the smface.

Social or semisocialbehavior in nomiine bees is here reported for the fust time from India (Batra, 1964). InNomiacapitata and perhaps inN oxybeloides, presumed sister bees share

cell clusters and cooperatively provision the same cel ls. Unlike other primitively social bees, the sisters simultaneously forage and oviposit. There is no apparent inhibition of the

oviposition of daughters, a lthough their nest-founding mother or perhaps an older sister

appears to stay in the nest and continue to oviposit. Nesting behavior and nest structure of

Nomiacapilala and N oxybeloides are similar; cells arc vertical and clustered. In contrast

Nomianasicana makes horizontal cells scattered along the main burrow.

5.5 Qualities of Good Nesting Sites

There are three important factors determining the quality of alkali bee nesting sites,

whether natural or managed i) soi l moisture; (ii) soil composition and texture; and (iii)

vegetation (Johansen & Mayer, 1982).

5.5.1 Soil Moisture

Soil must be moist down to at least 12 inches (31 em). Soil moisture in good sites

varies from 8 to 32 per cent depending on soil type. Soil moisture can be measured

using a tensiometer. A reading of 15-25 centibars indicates adequate moisture regardless of soil texture. Dry nesting sites have been a limiting factor in alkali bee

production. Good natura l moisture conditions are associated with sha llow layers of calcareous hardpans lying a few inches to several feet below the surface.

Seepage water may be sub-in·igated nearby nesting sites when a shelf of this

impervious material lies along a river, canal, or pond. Where this occurs,

populations of alkali bees may build up naturally with little management. Most

nesting sites are man-made and require an artificial water supply prov ided with

shallow ditches m ade across or armmd beds, or some kind of subsurface

distribution system.

5.5.2 Soil Composition and Texture

Maintaining proper soil texture in bee beds is almost as important as providing

adequate water. These two factors are inteiTelated. A soil of poor texture can limit

the movement of wate r through the upper horizon even when water is abundant.

Conversely, soil of excellent texhrral qualities is of little value where water is in

short supply.Proper soil texture encourages excavation by bees and a llows

continuous capillary flow of subsurface water towards the surface, replacing water

in the upper layers at a rate equal to or slightly greater than the rate of evaporation.

The SLllface should not be crusty or fluffy. The goal of soil texture management is a

mo ist and moderately compact upper soil horizon which persists throughout the active bee season.

Uniform soil moisture and good nest digging condi tions depend largely on the

percentage composition of clay, sand. and si lt, w ith clay preventing capillari ty at

levels g reater than 25 per cent. Soils high in sand ( 45-80 %) are difficult to seal, and

excessive water movement and evaporation may occur. The likely result is a wet

bed with a dry, powdery surface. Sandy beds become quickly populated with

---·--~-~---~

maximum number of nesting bees but do not maintain good populations for more than 3-4 years. The best alkali bee beds have soil classed as silt loams with 2-6%

fine silt and 42-68% coarse silt. They contain 13-24% clay and 10-40% sand.

5.5.3 Vegetation

The surface should be essentially bare with sparse vegetation. Plants use up soil moisture and bees prefer to nest in bare ground. Nevertheless, a little vegetation can help protect bees from summerrains and reduce wind erosion.

5.6 Establishing the Bees

The final and critical step in developing a new bed is establishing a population of bees. This may be easy if the new site is close to existing beds which support large and growing bee populations. Surplus bees from these established sites ·will move to the freshly prepared surfaces which are relatively clear of vegetation .If a new bed is isolated from existing populations, bees must be introduced to the area either as adults or as immature. Several methods have been developed for transplanting/ transfening bees from one location to another, sometimes over great distances.

The most successful and widely-used method is to transplant cores or blocks of soil containing prepupae. This must be done in spring before the immature transform into pupae or adults. Cores are placed on pallets and loaded on to a truck for transport. They should be covered with moist canvas or burlap if they are being transported long distance. In this manner, several thousand cores can be moved by one semi-huck. When the cores arrive at a new site, they should be buried in 12 inches (30 em) rr·enches and puddle in with sparing amounts of water. Soil at the new site should be properly prepared before the transplants are installed. It should have about the same moisture content as the core.

Cores should be installed in straight lines at least 4-6 inches ( 10.2-15.2 em) apart to ensure good soil moisture. They should never be stored for any length of time before burying them; this increases the risk of desiccation and decreases the viability ofprepupae.

5. 7 Measurement of Bee Density

The number of foragi ng female bees required in field is not fully known, but it should probably exceed 3000 per acre (741 0 /ha) in lucerne. It is easier to measure bee densities by assessing nest concentration. A good natural nesting site wi11 average about 1 million nests per acre (2.5 million ha). One acre (0.4 ha) of bed with this number of nests should

provide for 1000 lb of clean seed per acre (1120 kg/ ha) on 200 acres (8 1 ha) of lucerne. The maximum population in artificial beds is about 5 Yz million nest per acre ( 13.6 million !ha)(Jobansen&Mayer, 1982).

5.8 Limitations

>- Their services are confined to areas where rainfall, particularly during the active season, is

unlikely.

The beds cannot be transported; therefore, the crop to be pollinated must be planted near the bed. The bed must be planned and constructed many months before ils pollination service is expected.

Bee bed may be lost--quickly and easily--to flooding, predators, parasites, diseases, or pesticides and other agricultural practices.

Alkali bee-blue band Alkali bee-blue band

Alkali Bee nest

Alkali bee nest sites -the mounds of excavated soil called tumuli

67

6.0 Stingless Bees and Meliponiculture

Members of the Apidae subfamily Meliponinae or "stingless bees" are social insects. Some species have clusters of as many as 80,000 individuals; other species, less than 100. The two important genera are Melipona and Trigona. They do not occur in the

United States but are present and of economic significance in Mexico as well as Central and South America. Trigona spp. also occurs in Africa, Southern Asia, and Australia. They are mentioned here because of their widespread distribution over the tropical and subtropical areas of the world, their value in the pollination of many crops, and their long-time culture for the production of honey and "wax".

These bees have been studied taxonomically by Schwarz (1948) and behaviorally by several men, especially byNogueira-Neto (1948a, b, 1950, 1951 ), Nogueira-Neto and Sakagami (1966), Ken (1946, 1 948, 1951 ), Sakagami (1966), Sakagami and Oniki (1963), Sakagami and Zucchi (1967), and Zucchi et al. (1967). Meliponicultme was reviewed and discussed from the practical standpoint by Ordetx and Perez (1966). The following discussion is drawn largely from the above references. The females possess weak or vestigial stingers but are unable to inllict pain with them, hence the term "stingless bees." Some species have mandibles sufficiently strong to inflict a rnild bite or to pull hairs, or they may crawl into the ears or nostrils of the intruders. Others emit a caustic liquid from the mouth that, in contact with the skin, causes intense initation. Most species, however, are not bothersome to man, and he may safely manipulate them with ease, even to having his face within inches of a Trigona nest containing many thousands of individuals.

Stingless bees were kept by man centuries before the ani val of Columbus or the common honey bee (Bennett 1964). Some species produce an acceptably delectable honey, as much as half a gallon per colony per year. Others produce less desirable, thin (35 percent moisture versus half that amount in our domestic honeys), strongly acid honeys. One species (Trigona (Lestrimellita) limao Smith) produces a honey used to induce vomiting (Bennett 1965). The most common species used in miliponiculture is Melipona beechii Bennett. When the wax is secreted from the glands on the abdomen of stingless bees it is simi lar in appearance to that of Apis mell(fera, but it is then mixed with propolis and the product, called cerumen or Campecbe wax, is more or less black. Cerumen is used for waterproofing on farms and in villages, in ink and lithography, and in other restricted ways.

Originally, the colonies were kept in gourds, tree trunks, or similar cavities, but an improved hive has been developed that permits easy manipulation and transpo1iation of these bees. This hive is about a cubic foot in volume--sufficient for the 3,000 to 5,000 bees in an A1. beechii cluster. If necessa1y, additional space can be added for larger clusters. A nest of Trigona clavipes (F.) in a hollow tree, sketched to scale by Sakagami and Zucchi ( 1967), was 8 by 8 by 50 inches and had a worker bee population that "apparently exceeded several tens of thousands." It contained "at

68

least 20" horizontal brood combs separated from the collection of pollen and honeypots. The size

of hive acceptable to a colony of this size was not given. Life histories and habits-The size of stingless bees varies from 2 to 14.5 mm. Trigona duckei Friese is the smallest species of stingless bee known; Melipona interrupta Latrielle is the largest. M beechii is slightly smaller than Apis mel/ifera. The colors of the different species va1y from black to brown, red, orange, yellow, and white. The nest entrance is frequently reduced to permit only a single bee to enter at a time. The nest may be covered by a membranous wax and propolis network, which envelops and protects the nest and brood. There may be a single or multiple layer ofbrood--the individual cells vertical in some species, horizontal in others --or the cells may be in a cluster like grapes. Some species use the brood cells only once, then they are destroyed and reconstructed. The honey and pollen are not stored in the brood comb but in irregular cells outside of the brood nest.

The queens of Trigona are reared in queen cells, similar to those of Apis mellifera. Melipona queens develop in cells that externally seem to be no different from those that produce drones and workers, usually one queen to three to six workers. The workers of Melipona fill the cell with food before the egg is deposited. Each colony has a single sovereign queen but tolerates numerous virgins. A 4,000 worker bee population of M beechii may have 50 virgin queens living ham1oniously with the mother queen. Mating occurs in the air.

6.1 l~01·agc Sources

Stingless bees havenumberofforage sources listed in table 5.

Table 5. Forage sources for stingless bees

Family Plant species Habit Flowering season

Acanthaceae Asystasia gangetica (L.) T. Anderson Herb August-December

Dicliptera cuneata Nees in Wall. Herb November -March

Anacardiaceae Mangifera indica L. Tree January-March

Annonaceae Annona squamosa L. Tree March-July

Polyalthia longifolia (Sonner) Thw. Tree March-August

Apocynaceae Tabernamontana divaricata (L.) R.Br. Ex. Roem. and Schult. Shrub Throughout the Year

Arecaceae Phoenix sy/vestris (L.) Roxb. Tree January-May

Asteraceae Cosmos bipinnatus Cav. Herb Throughout the Year

Tridax procumbens L. Herb Throughout the Year

Tagetes erecta L. Herb Throughout the Year

Bignoniaceae Spathodea campanulata Beauv. Tree February-April

Tecoma stans (L.) Kunth Tree September-March

Bombacaceae Ceiba pentandra (L.) Gaertn. Tree January-March

Burseraceae *Boswellia ovalifoliolata Bal. and Henry Tree March-April

Caesalpiniaceae Caesalpinia pu/cherrima L. Tree Throughout the Year

69

Family Plant species Habit Flowering season

Bauhinia purpurea L. Tree October-December

Peltophorum pterocarpum Backer ex. K. Heyne Tree March-June

Tamarindus indica L. Tree April-August

Capparaceae Crataeva magna (Lour) DC. Tree April-May

Combretaceae Terminalia pal/ida Brandis Tree May

Dipterocarpaceae *Shorea roxburghii Don. Tree March-April

*S. tumbuggaia Roxb. Tree April-May

Euphorbiaceae Jatropha gossypiifolia L. Herb June-October

J. pandurifolia Andr. Shrub Throughout the Year

Fabaceae Dalbergia sissoo Roxb. Tree April-August

Gliricidia sepium (Jacq.) Walp. Tree January-March

Pongamia pinnata (L.) Pierre Tree March-June

Pterocarpus santalinus L.f. Tree March-May

Rhynchosia beddomei Baker Shrub November-January

Lamiaceae Hyptis suaveolens (L.) Poit. Shrub September -November

Leonotis nepetaefolia R.Br. Herb October-January

Lecythidaceae Couroupita guianensis A ubi. Tree Throughout the Year

Liliaceae Lilium candidum L. Herb April-June

Malvaceae Malvastrum coromandelianum (L.) Garcke. Shrub Throughout the Year

Meliaceae Azadirachta indica A. Juss. Tree March-May

Mimosaceae Mimosa pudica L. Herb Throughout the Year

Moraceae Artocarpus heterophyllus Lam. Tree November -August

Ficus religiosa L. Tree March-August

Moringaceae Moringa oleifera Lamk. Tree Throughout the Year

Muntingiaceae Muntingia calabura L. Tree Throughout the Year

Musaceae Musa paradisiaca L. Herb Throughout the Year

Myrtaceae Psidium guajava L. Tree January-May

Syzygium cuminii (L.) Skeels. Tree May-June

Nyctaginaceae Boerhavia diffusa L. Herb Throughout the Year

Polygonaceae Antigonon leptopus L. Creeper Throughout the Year

Rhamnaceae ZizyphusmauritianaLamk. Tree Sept. -January March-June

Rubiaceae Morinda tomentosa Roth. Tree April-August

*Wendlandia tinctoria (Roxb.) DC. Tree May

Sapindaceae Sapindus emarginatus Vahl. Tree November-January

Sapotaceae Achras zapata L. Tree February-April

Solanaceae Capsicum frutescens L. Herb Throughout the Year

Solanum melongena L. Herb Throughout the Year

70

Family Plant species Habit Flowering season

Sterculiaceae Sterculia foetida L. Tree January-March

Verbenaceae Gmelina asiatica L. Shrub February-October

Tectona grandis L.f. Tree June-November

Zygophyllaceae Tribulus terrestris L. Herb June-October

Stingless bees are important pollinators of several crops (table 6) as given below:

Table 6. Crops pollinated by stingless bees

S.No Crop group Crop

1. Fruits Peach, Plum, Pear, Guava, Citrus, Litchi, Strawberry, Jack fruit, Bread fruit

2 Vegetables Cucumber,Water melon,Squash,Bittergourd,Sweet pepper, Egg plant, Onion

3 Pulses Pigeon Pea

4. Oil seeds Sun flower, Castor, Niger

5 Spices Cardamom, Coriander

6 Trees Indian jujube, Subabul, Soap nut, Kapok, TamarindSago palm, Rubber, Eucalyptus Some of the crops, like papaya, passion fruit, banana, custard apple, oil palm, cocoa, cashew, black pepper which stingless bees visit, are pollinated by quite different means.

Different nesting sites for stingless bees (Courtesy Dr AJS Raju).

71

--~~·--------~---

Different nesting sites for stingless bees (Courtesy Dr AJS Raju).

Stingless bee brood cells

Nest entrance types of Trigona iridipennis at Visakhapatnam: Arrow mark indicates nest entrance; guard bees are present at the entrance of some nests( Courtesy Dr AJS Raju).

6.2 Pollination of on-Crop ~pecie~

Stingless bees are efficient pollinators of non-crop species in natural habitats. They play a vital role in sustaining the forest flora, epiphytic orchids and rare aquatic plants. In some cases stingless bees may do harm for some crops by removing nectar or pollen without pollinating the flowers. This makes the flowers less attractive to the other insects or birds which are the effective pollinators of that crop. In some extreme cases stingless bees may cause more serious ha1m. Form example, stingless bees have been reported to damage the flowers of the canola crop and drive away the effective pollinators of passion fruit.

6.3 Advantages ofStingle$S Bees over Honey Bees

1. Stingless bees are generally harmless to humans and domesticated animals. So they can be safely kept close to a house and be handled by people who are allergic to honey bee stings.

2. Small size of stingless bees allows them to have access to many kind offlowers, whose opening are too narrow to permit penetration by other bees. Hence, they co­exist peacefully with commercial bees. They collect and utilize considerable nectar and pollen throughout most of the year, therefore, numerous flowers must

3.

4.

be visited and pollinated.

They can be manipulated in hives like honey bees. The hives are small, easily handled, and relatively inexpensive.

The foraging range of stingless bees is shorter than commercial bees. Hence they can be very wel l utilized for pollination.

5. Stingless bees are able to forage effectively in glasshouses. Honey bees usually do not forage well in confined spaces. However, if condensation develops inside the glasshouse panels they get trapped.

6. Stingless bee colonies are not able to swann away as honey bees because the mature stingless bee queen is tmable to fly. Stingless bees can, however, set up a new nest when the old one is full, but part of the colony always remains in the original location.

7. Stingless bees are resistant to the diseases and parasites of honey bees. They are not affected by the vims diseases of honey bees or mites. However, stingless bees have their own natural enemies but these are not shared with honey bees and are not very senous.

8. The byproducts ofhoney and cerumen are usable.

6.4 Disadvantages of Stingless Bees

9. Stingless bees cannot tolerate cold weather; therefore, they are limited to the tropical and subtropical regions.

I 0. The byproducts are produced only in small quantities, and they are less desirable than those of the honey bee.

ln general stingless bees possess many characteristics that enhance their importance as crop pollinators both as wild populations and managed pollinators. Their character such as perenniality, polylecty, floral constancy, recruitment and harmlessness make them better pollinators. Challenges to their wide spread use include the lack of availability of large number of colonies and the dearth of knowledge about meliponiculture. Improved domestication practices would increase colony avai lability for planned pollination there by reducing reliance on natural populations. Stingless bees provide substantial economic benefits by their crop pollination service which are to be quantified in many crops. Stingless bee pollination has not been properly studied for many crops there is still much to be learnt and there are many management problems with stingless bees still to be solved. Stingless bees display greater diet breadth and range of foraging behaviour than honey bees which make them the potential pollinators of future best suited to the needs of particular crops and habitat.

j

' j

7.0 Management of Wild Bees

T he expensive need for insect pollination in modern agriculture has made bees a vital factor in crop production. To most people, the word bee suggests only the common honeybee, perhaps also bumblebees; but the bees are in fact a super family of the order

Hymenoptera, containing an estimated twenty thousand species. They are in fact a group of flower-visiting wasps termed as bees that has abandoned the wasp habit of provisioning nests with insect or spider prey and instead feeds its larvae with pollen and nectar, collected from flowers or with glandular secretions u ltimately derived from the same sources. They are a large and diversified group, considered to be consisted of nine families (Michener, 1974). Except in certain Colletidae which carry pollen with nectar in the crop, the structures used for canying pollen consist of scopal hairs having various locations and arrangements. One such group is the

Masaridae, a family of wasps closely related to the Vespidae and in the superfamily Vespoidea. The other group of wasps which abandoned predation as a source of larval food was from an entirely different source than the masarids, namely the wasps of the superfamily Sphecoidea. From this group arose the bees. Bees, entirely dependent on flowers for food, could not have arisen before the appearance ofthe angiosperm plants.

How early bees arose from the sphecoid wasps is unknown; it might have been as late as the middle Cretaceous since angiosperms also became the dominant vegetation in middle

Cretaceous times. Primitive angiosperms had relatively shallow flowers, such as can be used as pol len and nectar sources by short-tongued insects, including many beetles, wasps, and the short­tongued bees. The first five families, listed as Colletidae. Halictidae. Oxaedae, Andrenidae and Mellitidae are characterized by usually sho11 mouthparts and are often grouped as the short­tongued bees. The other four farnilies-Fidelidae, Megachilidae, Anthophoridae and Apidae are classified as long tounged families, al l are equipped with elongated glossae, maxillary galeae and basal segments of the labial palpi forming the sucking apparatus for taking advantage of nectar sources with deep tubular flowers. Flowers with deep corolla tubes probably arose in co­evolution with their plincipal pollinators. The legume crops and their pollinators such as alfalfa­

megachilid constitute an imp011ant example. There is close relationship between bumble bee species and their major plant hosts accordingly to their length. The worldwide distribution of these bees and their remarkable ability for proliferation are the results of their higher degree of adaptability (Stephen et al., 1969, Michener, 1974).

In many colonies there are interrelations among individuals, such that behavior of one in fluences the behaviour or development of another. All these interrelations are termed social interactions.

Feeding of a larva by a bee is an example of a social interaction. As indicated previously, colonies of bees range from those that seem almost insignificant-two or three bees in a burrow in the ground or in a hollow stem- to the large colonies of the honeybees. The kinds and amount of division of labour and communication among bees in colonies very greatly. Species are often

25

called solitary, communal, social, and so forth. Such terms are generally applied to the most complex type of organization attained during life cycle of the species. In the majority of species of bees each female makes her own nest, or sometimes several of them, without regard to the locations of other nests of the same species. Such bees mass-provision the cells by placing enough pollen and nectar in each to provide for the entire growth of a larva. After oviposition, the mother seals the cell and goes on to construct and provision another. Ordinarily she dies before her progeny mature and emerge from their cells; therefore there is no contact between generations. Probably the majority of species of Solitary bees have only a single generation per year, the adults emerging and flying about during a relatively brief season, sometimes only two or three weeks. Such species pass the rest of the year in the nest. The feeding stage of the larva is ordinarily brief, often only a few days, and most of the year is passed in the pupal stage or as young adults either still in their natal cells or in special hibernating or aestivating places. Some solitary bees, however, regularly have two generations per year, for example one in spring and another in the autumn, while others go through a succession of overlapping generations so that, except in the spring, al l stages can be found at anytime during the watmer months of the year.

Nest aggregations occur most commonly among bees that bunow in the soil. Aggregations of such burrows may vaty from a few to 10 nests scattered that one wonders if they constitute an aggregation at all, to small, dense clusters of nests like those of Lasioglossum versatum in Kansas. Some bees that make burrows in stumps or logs instead of soil also fonn aggregations. For example. one may find numerous nests of carpenter bees (Xylocopa) in a single post or building. Bees mostly megachilids that construct exposed cells of mud and other materials brought to the site sometimes also form aggregations of nests: for example Chalicodoma muraria sometimes covers large portions of walls in southern Europe and North Africa with masses of its cells. Communal quasisocial and semi-social groups are so similar superficially that the convenient collective term parasocial has been proposed for them. Parasocial colonies are simply any colony in which the adult bees consist of a single generatio~ unlike the eusocial forms in which two generations of adults are ordinarily present. A communal colony consists of a group of females of the same generation using a single nest each making, provisioning, and ovipositing in her own cells. In the enormous genus Alldre/la, most species are solitary, some nesting in aggregations. However, Alfdre/la bucephala (Perkins, 1917) and Aferox live in colonies that are probably communal. These are small colonies with two to several females, usually of about the same age and of the same generation, cooperatively construct and provision cells. More than one bee working on a given cell. As in communal groups, each female has enlarged ovaries and is mated, indicating that each is an egg layer. Some species of Nomia are possibly quasi social, although knowledge of their colonies is inadequate (Batra, 1966a ). The best known of such species is N. capitata from India. These are small groups which show cooperative activity and division of labor among adult females as in eusocial groups. Polygynous young colonjes of other halictines (Vleugel, 1961) are often temporarily semisocial in that division of labor develops among gynes, one becoming the egg layer or queen, the others

7R

auxiliaries or, in effect, workers. These are family groups each consisting of one adult female and a number of immature offsprings which are protected and fed by the adults. The mother leaves or dies before or at about the time that the young reach maturity. There is no division of labour among adults, as is found in semisocial and eusocial groups. Young colonies of Bomb us, before workers are produced, are subsocial; the queen progressively feeds the growing larvae in a more

or less subdivided common cell. However, the true social Hymenoptera, for which the word eusocial was coined by Batra (1966b), live in colonies which are family groups consisting of individuals of two generations, mothers and daughters. Usually in bees a eusocial colony contains only one queen and the bulk of the females are workers (daughters). Division oflabor, with some individuals functioning as egg layers or queens and other as workers. that is, with more or less recognizable castes occurs in both the semi social and eusocial colonies but not in

the other types of colon ies.

Colonies are long-lived and sustained tluough periods of adversity by food for adult as well as larval consumption, stored in the nests but in brood cells. Integration within colonies is complex and involves a variety of behaviour patterns, pheromones and physiological adaptations that would have no obvious function in solitary forms. Aggressive behaviour among individual of the same colony is rarely evident, nor is the egg eating that is often associated with such behaviour. Communication concerning food sources and at swarming time, concerning nest sites is well developed in many of these bees. Larvae are fed at least in large part on glandular secretions of workers. Populations of colonies are commonly in the thousands (upto over 60000 for Apis. 180,000 for some species of Trigona), although some species often have colonies of only one or two hundred. Only Bombus and the highly eusocial bees store food in quantity outside of brood cells for use of adults and for transfer to larvae or brood cells as needed. In most groups of social insects interactions between adults and young (i.e., brood-eggs, larvae, pupae) are universal and important parts of the social organization. Exchange of food between larvae and adults is well known in ants and vespid wasps, and it bas often been supposed that larval activity or secretions are of great importance in maintaining the social group. In most kinds ofbees, however, there are

no contacts between adults and young ones because the cells in which the young ones are reared closed before the eggs hatch, each cell being mass provisioned with enough food to provide for the entire growth of the larva. Progressive feeding, which of course involves adult- larva contact from time to time during the growth of the larvae occurs among bees only in Apis, Bombus and most allodapines. Even the highly eusocial meliponines have mass-provisioned cells which,

together with the cocoons spun by the mature larvae, completely enclose the immature stages for the whole developmental period.

The number of species of solitary bees is greatest in the warmer, more arid sections of the world, particularly in the semideseti areas as typified by those of western North America, North America, South Africa, Australia, northwestern Argentina, and South-central Eurasia. An abundant and diverse solitary bee fauna is common adjacent to mountainous areas where

77

moderate rainfall conditions exist. The rich bee fauna found in mountains adjacent to arid or semiarid areas is only partially explained by the stratification into altidudinal zones. The varied soil type and exposures, rock niches, beetle holes in wood, and pithy-stemmed plants offer many diverse nesting niches. The world-wide distribution of bees and the remarkable proliferation of species attests to their high degree of adaptability. Numerous definitions have been proposed to

distinguish between social and solitary bees, but recent infmmation has shown that hard and fast distinctions cannot always be nude.

The biology and behaviour of the solitary bees has attracted the attention of an increasing number of research workers dUiing the past 15 years: those exploring the value of solitary bees for pollination pUI-poses (Bohart. 1972: Stephen, 1969) and those attempting to evaluate the significance ofbiological patterns as a supplementary tool for the determination of phylogenetic

relationships among bees (Michener. 1974). In the Northwest about half of the Megachile rotunda/a larvae of the first generation pupate and emerge as adults in the late summer. I11 some seasons a small percentage of the progeny of the second generation emerge as a third generation, although they usually have little time for nesting before being killed by cold temperatures. Some Megachile and H_rlaeus and some Anthophora have more than one complete generation and overwinter as prepupae. Some single-generation species overwinter as adults in their netal cells. For example, Osmia ligna ria usually emerges as an adult in April and dies for about three weeks, The type of life cycle described above is apparently an adaptation for early spring emergence,

although some species exhibiting it (for example, Osmia tera) do not until late spring. It is interesting to note that another species with relatively late emergence. Osmia cal(fomica, usually has some individuals overwintering as prepupae. Other bees overwintering as unemerged adults include most Osmia and Andreno and some Antbophora, Megachile, and Emphoropsis. Bumblebees undergo a life history similar to that of halictines. The overwintered female is sole egg layer for several generations, which overlap broadly because egg laying is continuous. The overwintermg female (queens) which are distinctly larger than the earlier generations of

workers, are usually produced after the worker: brood ratio is favourable for intensive feeding of queen brood. The Xylocopinae overwinter as emerged adults, as do halictines and many apids. However, both sexes of Xylocopines overwinter in a dormant condition and mate in the spring (some mating is reported to take place also in the fall). The females usually overwinter in the

natal nest with males from other nests often joining them.

In many megachilids, males appear more numerous but the exact ratio varies. Exact causes leading to the va1iance are not well understood. In Megachile rotumdata, the ratio of males to fema les in large samples taken from different nesting population bas been seen to vary from I: I to 10: 1. The difference in ratios is directly correlated with the diameter of the tunnel in which they

are nesting. Emergence of males in advance of females is evident in the alkali bee and in Megachile rotundata, as in most other species ofbees that have been studied. This phenomenon, referred to as protandry, is a general rule among solitary bees and is interpreted a evolutionaty

70

adaptation, assuring the presence of males fo r mating with emerging females.

The avai lability of suitable substrates for nesting is one of the most common factors limiting the population and distribution of bee species (tab le 3).The principal type nesting

microenvironments include soil, wood, small and large cavities and even fully exposed surfaces. Species nesting in soil may select horizontal to gently sloping surfaces or vertical banks. The vertical surface may be bare or overhung with vegetation, or rarely with a grassy cover, and its exposure may provide maximum minimum shade. Vertical banks are usually dry, but they may be moist in shaded gullies or dram ditches. The soil surface may be wet or chy, but appreciable moist is usually avai lable where and when the cells are constructed. At least one observation has been made of Anthophora species tunneling in moderately hard sand stone. Ceratina, some Megachile, Xylocopa and several genera of small sphecid wasps burrow in the soft p ithy plant stems of p lants such s raspberry, black-berry and sunflower in constructing nesting tunnels. A wide variety of nesting matetial is utilized by megachilids. Their nests may be found in snail

shells (Old World Osmia), pockets or cracks in rocks (many osmiines), attached to twigs or rock surfaces (some Dianthidirim), or in almost any kind of hole and crevices of vatying shape and size for nesting. Tendency of bees to renest in close proximity to their parent's nest is one of the main causes of gregariousness, and that selection of a peculiar soil condition has a minor effect. Most burrowing bees construct laterals from the main burrow with one or more cells arising from each lateral. Exceptions occur among species that construct a main burrow with one or more cells

attached to the end, along the sides, or atTanged within the main burrow in linear series. After the cells arc provisioned and capped, the nest, or a portion of it, is plugged. Some species only plug the area immediately exterior of each cell, others completely backfill the laterals, and sti II other plug the entrance when the nest is completed. Most species that arrange their cells in line series construct complete cells, i.e. provide walls as well as top and bottom for each cel l (most Megachi/e. Anthidium). In Megachile, the rounded base of each cell inserted into the concave apex of the cell below, resulting in a nearly intact, weakly differentiated column of cells.

Table 7. Nesting behavior, nest acceptance and temperature tolerance of subtropical Megachild and Xylocopinae bees

Bee species Crops pollinated Nesting behavior Nest tunnel Temperature accepted for activity

Diameter (mm) Length (em) oc

Megachile haryanensis Alfalfa Leaf cutter bee 4.5-5.5 10-12 26-43 (M.nana)

Chalicodoma rubripes Alfalfa Masson bee 5.5-6.5 10-12 28-43 (M.flavipes)

C.lanata(M.Ianata) Pigeon pea Masson bee 6.5-7.5 10-1 2 26-38

Chalicodoma cephalotes Pigeon pea Masson bee Utilizes resin/ 5.5-6.5 10-12 27-41 (M. cephalotes) animal faecal material for

partitioning and closing the cells /tunnels

79

Bee species Crops pollinated Nesting behavior Nest tunnel Temperature accepted for activity

Diameter (mm) Length (em) DC

Xylocopa fenestrata Cucurbits Carpenter bee 10-12 23-30 25-48

Xylocopa pubescens Cucurbits Carpenter bee 10-12 23-30 25-48

X.valga Almonds,apples Carpenter bee 10-12 23-30 5-30

X.aestuans Cucurbits Carpenter bee 10-12 23-30 10-47

Modified from Sihag(1994)

The number of cells per nest ranges from one to several thousand. Most solitary soil butTowing

species make only one nest, which contains as many cel ls as foraging conditions and the

reproductive potential of the bees allow. Nest of Nomia me/anderi, may contain from 5 to 24

cells, depending upon the availability of forage and th quality of the substrate in which they are

nesting. Some species ofOsmia normally select small pockets in rocks that accommodate only

one cell, but the same specie may place several cells in somewhat larger holes. !Yfegachile

rotundata will accept tunnels which accommodate only a sing le cell but more commonly uses

long tubular cavities in which it places as many as 20 cells. The reproductive potential of this

species is even higher (upto 40 eggs), but there seems to be an upperlimit to thenumbcrofcells it

places in a single tube, independent of its length.

The world wide distribution of these bees and their remarkable ability for proliferation are the

result of their higher degree of adaptabi lity. Floral fidelity is the major attribute of these species

making them sure of maintaining species characteristic within plant species. Based upon degree

of association in between bees and the plant species i.e. the number of plant types visited for

pollen collection (Linslay eta/., 1958), the bees are tenned monolactic (visiting one species)

oligolactic (visiting few related genera) or poly lactic (visiting many types of plants). The solitary

bee species arc mostly oligo lactic. D ifferences in seasonal adaptability, the pattern of origin vis­

a-vis individual bee specialization basically detetmine bees abundance for pollen col lection on a

particu lar· plant. The principle factor which determi nes the effectiveness of such poll inators for a

particular crop or plant species depends upon the bee abundance, bee flight period, bee flight

hours per day and the number of flowers visited per day. The factors which contribute to bee

survival in nature and their propagation depends upon the availabi lity of natural or man made

nesting devices of preferred dimensions, abundance of natural parasitic, or predators, incidence

of d isease or pesticide poisoning, and the natural brood mortality during active or the dormant

season. Most important is the synchronization ofthe bee flight period with the major b looming

period of the crop. This is achieved through appropriate provisions of nesting devices and

regulating development of adults so that there is synchrony in adults formation with crop

blooming. Following are the characteristic features of such bee management programmes for

c rop pollination.

Rn

..

i)

(ii)

iii)

iv)

7.1

Provision of appropriate nesting devices of brood cell formation.

Col lection and safe storage of brood nest of cells at low temperature.

Checking/controlled emergence of parasites or removal diseased cells.

Incubation of cells at appropriate temperature to regulate fom1ation

Protocols for Study of Non-Apis Solitary Bees

7.1.1 Pollinator Diversity

a. Collect the pollinating insects from target crops at weekly interval starting from commencement of the pollinator act ivity in the morning till the cessation of activity in the evening.

b. Collections should be made in time and space i.e. with the commencement of flowering of each crop till its cessation throughout the season

c. The collection should be made by five insect collection nets sweeps at all the random five spots equally distributed in the crop area.

d. Preserve the collected pollinators and get these identified from authentic sources for detennining the diversity of pollinator insects.

7.1.2 Habitat Diversity

I . Collect the non-apis pollinators from different agro-climatic zones as per the procedure described above from different crops/plants

2. Collections should be made in time and space i.e. with the commencement of fl owering of each crop till its cessation throughout the season

3. The collection should be made by five insect collection nets sweeps at all the random five spots equally distributed in the crop area.

4. Preserve the collected insects and get these identified from authentic sources for determining the diversity of pollinator insects.

Table 8. Nesting habits of solitary bees

Family Important genera Nesting habit

Andrenideae Andren a, Panurginus, Perdita, Pseudopanurginus Soil

Colletidae Colletes, Hylaeus Soil

Halictidae Agapostemon, Dufournea, Halictus. Nomia Soil

Melittidae Hesperapis, Melitta Soil

Anthophoridae Anthophora, Melissodes, Nomada, Soda water straws, bamboo stems, castor stems, drilled boards, pithy materials

Apidae Xylocopa -do-

Megachilidae Anthidium, Lithurgus, Megachi/e, Osmia Soda water straws, bamboo stems, castor stems, drilled boards

A1

Artificial domiciliation materials

3.2 Conservation Strategies

a. The best sites are bare or slightly vegetated tender silty loamy soils. These soils should be left undistw-bed raised in bunds for their natural nesting

Artificial domiciliation of non-A pis pollinators

Different nesting material viz., Soda water straws, bamboo stems, castor stems, drilled boards, pithy stems of varying size should be standardized for determining the nesting preference of non A pis pollinators. (table 8)

Step I. You need to prepare a shed in the field to place shelters for domiciling non- A pis bees

Step 2. Prepare cardboard/wooden boxes of size 1.5 xl.S feet. At least 3 such boxes with different nesting diameter should be evaluated for each treatment as given below:

Tl: Bamboo stem: 1 feet length with3 different diameter holes

1. 1.0-1.5 em,

2. 1.5-2.0 em

3. 2.0-2.5 em

T2 Castor stem: 1 feet with 3 different diameter holes

1. 4.0-5.0 llliU

2. 5.0-6.0 mm

3. 6.0-7.0 nun

T3 Pithitus stem : I feet with 3 different thicknesses

1. l.0-1.5cm

2. 1.5-2.0em

3. 2.0-2.5em

T4 Drilled wooden board with soda water straws inserted in holes

l. 4.5-5.5mm

2. 5.5-6.5mm

3. 6.5-7.5mm

T5 Soda water straw: Assorted size

T6 Sarkanda(Arundo spp) Canna indica

1. 1.0-1.5 em,

2. 1.5-2.0 em

R?

3. 2.0-2.5 em

Step 3. Observations to be recorded

Type of nesting material-----------------------

S.No Cell size Number of Size ofthe Bee Species/Family cells accepted cell accepted

Sarkanda

Drilled board with soda water straws

Percentage acceptance

Pithy stems

Shed for domiciles

Remarks

Step 4. Once some bees have accepted the nesting materials, study the following:

Specific Role of on- A pi .... · Species

1. Seasonal survey need to be made to identify non-Apis bee pol linators

2. Study their nesting sites

3. Factors affecting their population build up.

4. Study their behaviour under enclosures

83

5. Detem1ining the pollinator effectiveness under enclosures.

6. To determine the optimum number ofbees required per hectare of the crop

7. Specific foraging range ofnon-Apis pollinators

8. Providing nesting sites near target crop.

For the pollination studies of various crops, the conduct of the experiments and the observations should be recorded as per the following protocols

7.3 Non-APISBees and Future Prospects

84

Honey bees (Genus A pis Linnaeus) have often been credited with pollination services that are actually performed by other bee species. Hundreds of entomophilous crops are now known that are very poorly pollinated by honey bees. The act in major is shared by non­Apis or the so called wild bees (Parker eta/., 1987). There are few estimates available for the value ofnon-Apis pollination. Estimations declared the value of wild bee industry was well over US S l million per year in terms of expenditures and benefits in USA alone (Bohart, 1970, 1972). The benefits increased up to a range of US S 18-40 million in 1981 and, the total involvement of money over crossed US$ 81 billion (Levin, 1983).

Recent technological advances in agronomic practices have focused primarily on improving yield, increasing the number of crops grown and increasing the area of barvestable crops. These advancements have been applied indiscriminately to the majority

of crops and, in a very short duration, they have transformed farms into intensive monoculture systems. The positive results of these practices are impressive. The quality

and quantity of food bas increased, food costs have decreased, numerous fresh fruits and vegetables of high quality are available for much longer period, the quality and types of

prepared food products have greatly improved and, the large labour force once required has been reduced, at the same time crop area have increased. On the other hand, the technical advances and intensive farming practices have evolved numerous negative impacts on crop pollination and non-Apis populations (Richards, 1993).

A number of conservation studies have concluded that clearing land of trees and increased cultivation have inadvct1ently eliminated many of the nesting sites previously used by non-Apis pollinators (Renner, 1996; Cane, 2001 ). Frequent applications of broad­spectrum weedicides and pesticides have been responsible for the rapid decline of pollinator numbers within agricultural areas (Batra, 1995c). Changing inigation practices

have had long-term negative effects on soil nesting pollinators. Overgrazing of rangeland and the use of herbicides have indirectly reduced the presence of pollinators by decreasing diversity of pollen-nectar resources and, by eliminating required plant resources that are uti! ized by various wi ld-non-Apis bees in nest construction (Batra, 1979b and 1979c ). One of the consequences of an increased food supply for the world has been inadvertently

depopulating both numbers and species of native pollinators within agricultural environment (Roubik, 2001 ). This situation must be addressed if our agricultural ecosystem has to be sustained.

Second interesting aspect of pollination with honey bee for several crops lies in the fact that after one or two morning forages honey bees are least interested in pollen collection. It prefers collection of nectar on I y and pollens are attached to the scopal bristles accidentally (Sharma and Gupta, 1993). In such events, honey bee enters and after drinking nectar moves out of the corolla tube so swiftly that a flower hardly receives any help in tripping. In the field, less than one percent of the self-tripped flowers produce seed and most of the non-tripped flowers fail to do so. Studies available with regard to many leguminous crops

have rep011ed several non-Apis bees as intentional trippers and they continue to do so for many f-Ld l days (Zaleski, 1956). They deposit pollens in their nest chamber adjacent to their egg that hatch later as larva and feeds upon it. The process usually requires at least 3-15 days as per nest size of the species. Apparently, several wild-solitary bees are known for their effective pollination values abroad. Otherwise also, majority ofnon-Apis females are

supported with dense brush of pollen collecting scopa and in larger area of the body so that maximum amount of pollen load may be collected in each trip to the field (Torchio, 1987).

The benefits we derive from native pollinators are bebeved to be increasing as the honey bee industry experiences continued difficulties from mites and diseases. Furthennore the crops that are better pollinated by bees other than honey bees, are being grown more intensively. To protect the native bee pollinators, two alternatives have been suggested, one is the preservation and management ofhabitats and another is artificial domestication and management of bee species (Stephen and Every, 1970; Williams eta/., 1991). Bohart (1970, 1972) quoted that honey bees are not entirely satisfactory in thei r use fo r the maximum output, in enclosures. On the contrary, non-Apis species when invited to artificial nesting devices, have been found much more effective pollinators. Apparently, a

newer aspect has evolved which deals with "artificial domestication and management of wi ld bees for crop pollination (ADMP)".

In Notih America 3,500 species of wild-solitary bees have been recorded, commonly referred as "Pollen Bees" (Batra, 1994a). Alfalfa leafcutter bee (Megachile rotundata), blue orchard bee (Osmia lignaria propinqua), fuzzy foot bees (Anthophora pilipes) and, mustached bees (Anthophora abrupta) are some of the wi ld-bee cross pollinators which are successfully used in the arti ft cial domestication and management programme, to enhance the crop pollination (Batra, 1982, 1991, 1993, 1994b). Robinson et al, (1989b) detailed several parameters applicable for the pollinators. In brief, such progranm1es intends to sustain a good population of pollinators close to the crops by providing them

artificial nesting devices and, the normal principle usually understood was 'higher the number of pollinators higher wi ll be seed yield'. Kapil, Grewal, Kumar and Atwal (1970)

R!i

86

recorded some of the insect pollinators and noted their comparative abundance. However, after certain threshold limit population increase of the pollinator does not put any further positive impact upon the seed yield (Strickler, 1997). Instead, a consistency in increase in floral resource has to be maintained to sustain higher population of pollinator on the crop (Strickler, 1999; Strickler and Freitas, 1999), otherwise fear concerning escape of bee species would persist.

A few other successful programmes exist that has enhanced the number of native pollinators for fruit crops such as, using horn faced bees Osmia cornifrons in Japan and use of Osmia corn uta in Europe (Maeta, 1978; Torchia and Asineo, 1985; Maccagnani et al, 2003) and its introduction and establishment in USA (Batra, 1978a). Identically, many species of Megachile, Heriades and Osmia are found in sufficient numbers and need

intensive investigations in this direction because they are quite efficient visitors of several cultivated crops innotthem India (p.o. author).

Certainly, ADMP has emerged as a new entomological industry (Bohart, 1970). In India, the studies relevant to pollination of some more crops and the efficiency of several bee species were recently commenced by Kumar et al, (1994), Shatma and Gupta, (2001) and, Kapila et al, (2002). Some of these workers have made significant contribution by using the criteria such as amount of carried pollen loads, number of visits made on flowers etc. for different bee species. Megachiline bees have the maximum area for the collection of

pollen grains (beneath their abdomens) hence usually top the list with regard to carried

pollen loads.

1t is suggest that several correlated aspects should be undettaken in future such as, bee foraging studies coinciding with rotation of crops; population studies correlating different bee species on different crops; pollinator efficiency studies; effects of insecticides on various pollinator bee species; study ofimmatures, their developmental periodicities and, its variation under conh·oJied environment, so that supply of broods as seeds may be implemented in practice, alongwith blooming of different crops; brood transfer

techniques; studies relating parasites-predators and pests of adults and broods and, their mortality rate; impact of ecological factors on adults and broods; crop y ield studies; development of nesting devices for different bee species so as to retain maximum population of bees on crops etc. These will ultimately help initiate artificial domestication

and management progranune in the country for many useful bee species to obtain better seed yields. Domiciliation technology for domestication of wild solitary bees is available using different type of nesting materials such as soda water straws, bambbo stems, castor stems, drilled boards etc (for details see Kapil and Jain 1980;Abrol, 1991 ).

Aview of the experimental hut for domivcilation ofwild bees

Field hut for nesting solitary bees Pithy stems for Pithitis species

Management of solitary bees.

87

8.0 Conclusion and Future Strategies

I mproved agricultural practices has increased food supply over the past 50 years but a depopulation in both number and species of bee pollinators within agricultural environment bas resulted from land clearing, cultivation, irrigation and pesticidal use. The population of

honey bees presently is not sufficient to meet the huge demand of pollination above this honey bees can not pollinate al l the crops. Honey bee population may also fluctuate naturally in reponse to climatic calamity, the attack of paras ites and diseases. Reliance on a single agricultural pollinator is always prone to such crises. We conclude there is immediate need of diversification and conservation of poll inators.

The efficiency of insects as crop poll inators would depend on their biological characteti stics in relation to the crop and the environment in which they are needed. Each insect species which has been used as a pollinator so far would have its specific characteristics, which might be favorable or unfavorable from the standpoint of the user. The value of stingless pollinators is obvious from the farmer's point of view. Due to their compact colonies and safety for farmers and visitors, they can be used in areas where stinging insects are not desirable, as in greenhouses. However, very few surveys concerning pollination by stingless bees have been conducted in the temperate countries. The experiments to assess crop pollination efficiency by stingless bees and to improve colony management techniques are needed before they can be confidently used for the pollination of crops in greenhouses.

We stand at a crossroads, as the domesticated honey bees have suffered immense losses during thee recent years and there is a pollination crisis all over the world. There is a need to conserve altemative pollinators to improve crop and food security. Pollinators are essential to our environment. Seventy percent of the world's flowering plants, including more than two-thirds of the world's crop species, rely on pollinators to reproduce. One third of human diet depends upon the services provided by pollinators. Besides, poll inators are also important fo r production of forage crops required fo r cattle to produce daity products. As the beekeeping industry is in crisis and dur ing the past 50 years, an almost 50 percent decl ine in the number of managed honey bee colonies have been reported. The honeybee colonies are plagued by new pests and diseases and most recently, and most alarmingly, the beekeeping community is facing Colony Collapse D isorder, w here for still unknown reasons, worker bees simply abandon the hive. Evidently, it can no longer be safely assumed that honey bees wi ll provide all of farmers' future pollination needs. Efforts are therefore needed to explore manage conserve and multiply the alternative pollinators. The continuing trend toward larger monocul tures, insecticide use, and the concomitant lack of habitat-particularly a decline in the number and diversity of flowering plants available when crops are not in bloom--creates a landscape where few crop pollinators can survive.

To diversify our pollinators, we must better understand how to manage a variety of bee species as well as the habitat that supports them and their wild counterparts. Perhaps the silver lining of

88

Colony Collapse Disorder is its wake-up call to invest time, research, and energy into new managed pollinators and new ways of looking at farm management for the betterment of all pollinators. For agricultural as a whole the diversification ofpollinaton assemblages for crops is clearly important. Wild and domesticated non-Apis bees effectively complement honey bee pollination in many crops. Examples of management of non-Apis species for agricultural pollination include the use ofbumble bees, primarily for the pollination of greenhouse tomatoes, the sol itary bees Nomia and Osmia for the pollination of orchard crops, Megachi/e for alfalfa pollination, and social stingless bees to pollinate coffee and other crops. The value of the alfalafa leaf cutting bee M. rotunda (F.) as a better pollinator than honey bees for alfalfa has been clearly demonstrated by Richard, 1987. he concluded that the real impact of introduction of Megachi/e bees stating that alfalfa seed yield increased from 50 kg/ha to 350 kg/ha and with more careful handling it can be raised upto lOOOkg/ha.

Because information on the role wild pollinators in agriculture and the effects of agricultural methods on pollinators is largely speculative, research is critical to provide an understanding of this interaction. Little effort has been made to ensure the diversity of wild populations resulting in poor pollination and reduced y ields.

Research be encourageJ to determine which crops and wild flowers are bee pollinated, which bee species pollinate them best and at what densities and the economic benefits ofbee pollination per crop and per region. Relevant research and development and agricultural policies be adopted to ensure adequate populations of appropriate pollinators for different crops in different regions. Particular attention should be given to develop and improve teclmiques for the rearing of solitary bees and bumblebees, suppmt bee taxonomic research, promote a thriving beekeeping industry and poll ination services and advice to growers, encourage the use of native rather than exotic bees, monitor movement of commercially reared bees and the impact of diseases, and investigate the impact of honey bee introductions on native bee populations. Land use policies must be promoted to encourage appropriate management of agricultural, forested, semi-natural, conservation and amenity areas to improve habi tats for wild and managed bees. Particular attention should be given to conserve and restore natural vegetation, minimise soil surface disturbance which destroys nest sites, promote perennial herbaceous vegetation, bee forage seed mixtures and legumes and develop pesticides safe for bees. Agriculture has become more pollinator dependent because of a disproportionate increase in the area cultivated with poll inator dependent crops. If the trend toward favouring cultivation of pollinator-dependent crops continues, the need for the service provided by declining pollinators will greatly increase in the near future. The protection of native pollinators is critical. It would thus be prudent to set as ide areas for native pollinators in agro-ecosystems and to encourage their populations by providing forage and nesting sites for their conservation where native pollinators can thrive. The genera l public, pol icy makers and planners, and politicians must give due importance of pollination and po IIi nators, the seriousness of their dem ise, and the urgency for their conservation.

89

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Acknowledgements

The authors are grateful to the ICAR, New Delhi for supporting this publication

which will be helpful in sbating the information on emerging topic 'Non-Apis Bee

Pollinator' with the scientists and other stakeholders. Authors desire to express special

thanks to all expert sources for consultation, preparation and writing of this bulletin in

the present form.

92

D PAbrol

R KThakur

HD Kaushik

Sunita Yadav

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