social behavior and nests of some nomiine bees in india (hymenoptera, halictidæ)

9
Insectes Sociaux, Paris. Volume XlIl, 1966, n ~ 3, pp. 145-154. SOCIAL BEHAVIOR AND NESTS OF SOME NOMIINE BEES IN INDIA (HYMENOPTERA, HALICTID.,~E) ~1) By Suzanne W. T. BATRA (The University of Kansas, Lawrence, Kansas, U. S. A.) Social or semisocial behavior among bees is known in most Apidae (Apis, Bombus, Meliponinae), in some Ceratininae (Allodape-Exoneura group) and in various Halictinae; however, its existence in Nomiinae is here reported for the first time. A few species of Nomia are colonial, each female making her own cell cluster although several share a common burrow. In Nornia (A ustronomia) australica Smith as many as three females may share a burrow but each is said to make and provision her own cell cluster (RAYMENT, 1956). Nomia (Hoplonomia) punctulata Dalla Torre is primarily solitary but when occasionally more than one female are found in a burrow, each sister makes her own celi cluster (HIRASHIMA, 1961). That Nomia (Pseudapis) ruficornis Spinola may be social or semisoci, a,1 is suggested by cel,1 ,clusters illustrated by GUTBIER (1915) which seem too large to have been constructed and provisioned by a single female. Other species of Nomia that have been ,studied are (1) Contribution No 1304 from the Department of Entomology, The University of Kansas, Lawrence. This paper is the second of a series on behavior of Halic- tidae in India; the others are BAwrut and Mm~ENE~ (1966) and BATnA (1966). The author wishes especially tO thank Dr. A. S. ATWALfor providing facilities at Punjab Agricultural University, Ludhiana. Dr. I. H. H. YAaaow arranged for my study of specimens at the British Museum (Natural History) and Dr. C. D. M~C~E- SEn kindly checked specimens against types there as well as reviewing the manuscript. Field work was supported by the American Philosophica.1 Society (Penrose Fund, Grant No. 3628) and a Sigma Xi-RESA Grand-in-Aid of Research. Some of the'field work was financed by a grant (No. FG-IN-215) made the Punjab Agricultural Uni- versity by the United States Department of Agriculture under Public Law 480 (Project A7-ENT-19). -Preparation of the manuscript was sup,ported by National Science Foundatioff Grant GB3151 (C. D. MIC~ESEn). Dr. K. A. STOCKHA.~MEnkindly transIated :the summary into German. INSECTES SOCI,IUX,TOME XIII, N ~ 3, 1966. 11

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Page 1: Social behavior and nests of some nomiine bees in India (Hymenoptera, Halictidæ)

Insectes Sociaux, Paris. Volume XlIl, 1966, n ~ 3, pp. 145-154.

SOCIAL BEHAVIOR AND NESTS

OF SOME NOMIINE BEES IN INDIA

(HYMENOPTERA, HALICTID.,~E) ~1)

By Suzanne W. T. BATRA (The University of Kansas, Lawrence, Kansas, U. S. A.)

Social or s emi soc i a l b e h a v i o r a m o n g bees is k n o w n i n m o s t A p i d a e (Apis, Bombus, M e l i p o n i n a e ) , i n some C e r a t i n i n a e (Allodape-Exoneura group) a n d i n v a r i o u s H a l i c t i n a e ; howeve r , i ts ex i s t ence i n N o m i i n a e is he re r e p o r t e d for the f i rs t t ime .

A few species of Nomia are co lon ia l , each f e m a l e m a k i n g he r o w n cell c l u s t e r a l t h o u g h seve ra l s h a r e a c o m m o n b u r r o w . I n Nornia (A ustronomia) australica S m i t h as m a n y as t h ree f e ma l e s m a y sha r e a b u r r o w b u t each is sa id to m a k e a n d p r o v i s i o n he r o w n cell c l u s t e r (RAYMENT, 1956). Nomia (Hoplonomia) punctulata Dal l a T o r r e is p r i m a r i l y s o l i t a r y b u t w h e n o c c a s i o n a l l y m o r e t h a n one f e m a l e are f o u n d in a b u r r o w , each s i s te r m a k e s he r o w n celi c lu s t e r (HIRASHIMA, 1961). T h a t Nomia (Pseudapis) ruficornis S p i n o l a m a y be social or semisoci, a,1 i s sugges t ed by cel,1 ,c lusters i l l u s t r a t e d by GUTBIER (1915) w h i c h seem too la rge to h a v e b e e n c o n s t r u c t e d a n d p r o v i s i o n e d by a s ing le f emale . O t h e r species of Nomia t h a t have b e e n ,s tudied are

(1) Contribution No 1304 from the Department of Entomology, The University of Kansas, Lawrence. This paper is the second of a series on behavior of Halic- tidae in India; the others are BAwrut and Mm~ENE~ (1966) and BATnA (1966).

The author wishes especially tO thank Dr. A. S. ATWAL for providing facilities at Punjab Agricultural University, Ludhiana. Dr. I. H. H. YAaaow arranged for my study of specimens at the British Museum (Natural History) and Dr. C. D. M~C~E- SEn kindly checked specimens against t ype s there as well as reviewing the manuscript.

Field work was supported by the American Philosophica.1 Society (Penrose Fund, Grant No. 3628) and a Sigma Xi-RESA Grand-in-Aid of Research. Some of the'field work was financed by a grant (No. FG-IN-215) made the Punjab Agricultural Uni- versity by the United States Department of Agriculture under Public Law 480 (Project A7-ENT-19). -Preparation of the manuscript was sup,ported by National Science Foundatioff

Grant GB3151 (C. D. MIC~ESEn). Dr. K. A. STOCKHA.~MEn kindly transIated :the summary into German.

INSECTES SOCI,IUX, TOME XIII, N ~ 3, 1966. 11

Page 2: Social behavior and nests of some nomiine bees in India (Hymenoptera, Halictidæ)

1 4 6 SUZANNE W. T. BATRA

2: i.

. " . :i~:;~i S:.: E

S .?

Fro. 1 a n d 2 a r e to t h e s a m e scale . Fig. 1 : n e s t 2 of Nomia capitata b r o k e n a t X) . Fig. 2 : n e s t 1 of N. oxybeloides.

FIG. 3 a n d 4 . - - U p p e r (fig. 3) a n d l o w e r (fig. 4) cell c l u s t e r s o f ne s t 2 of N. eapitata, v i e w e d f r o m above .

FIG. 5 a n d 6, - - L o w e r cell c l u s t e r o f n e s t 1 o f N. oxybeloides (fig. 5) a s v i e w e d f r o m b e n e a t h a n d (fig. 6) as v i e w e d f r o m above ,

FIG. 7 to 9. - - Cel ls o f N. oxybeloides. Fig. 7 : s e m i d i a g r a m m a t i e cell w i t h feces, s t r a n d s of m y c e l i u m a n d p e r i t h e c i a ; ( a ) i n n e r l a y e r of w a t e r p r o o f l i n i n g ; (b) p e r i t h e e i a ; (e) o u t e r l a y e r o f w a t e r p r o o f l i n ing . Fig. 8 : p lug , as v i ewed f r o m i n s i de cell. Fig. 9 : cel l w i t h egg a n d po l l en ba l l .

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S O C I A L B E H A V I O R AND N E S T S O F S O M E N O M I I N E B E E S IN I N D I A 147

solitary, each female m a k i n g her own nest, a l though dense aggrega- tion,s occur in some species and several adults may temporar i ly wait in the parenta l nests before leaving to make their own nests (see CRoss and BOHART, 1960; STEPHEN, 1959).

Nomia capitata was described f rom China but the Indian specimens agree wi th the type in the Bri t ish Museum (Natural History). Voucher specimens for any necessary fu ture checking of identifications are ~ in the Snow Entomologica l Museum, The Universi ty of Kansas.

Nests of Nomia capitata Smith and N. oxybeloides Smith were found on the campus of the P u n j a b Agr icul tura l University, Ludhiana . This par t of P u n j a b has a subtropical , semiar id climate. The nests were observed and excavated, then females f rom these nests were measured and dissected to determine caste using the methods of MICHENER and WILLE (1961).

Nomia capltata Smith.

Nests of this minute species were in a lawn composed of c lumps of short grass and the pros t ra te xerophytes , Indigofera linifolia and Helio- tropitim strigosttm. It was in sun all day. Four nests, in a level area of about 6 X 2 meters, were f o u n d and excavated between Septem- ber 21-and October 29, 1964, when nest ing activi ty ceased.

The bees apparen t ly ceased nest ing activities because their sole pollen source, Indigofera linifolia (Leguminosae) , stopped b l o o m i n g in November. However, N. capitata d i d not resume activi ty when the plant began blooming again in F e b r u a r y and no bees were seen dur ing the sp r ing and until I left L u d h i a n a in June. In Bangalore, Mysore State (South India) , however, this bee was collecting pollen on I. lini- folia on December 30 and m a y be active there all year.

The ent rances of nests (fig. 1) were 2 to 2.5 m m in d iameter and hidden by the vegetat ion (one was ac tual ly in the center of a grass clump). One new bu r row wi th one bee and no cells had a radial t umulus of loose sand, 1.7 cm in d iameter and 8.0 mm high; the other

3 nests had no tumuli . The near ly s t ra ight vertical bur rows were in firm, sl ightly moist sand reach ing m a x i m u m depths of 38, 47 and 48 em. Bur rows were wider t han the nest entrances, 2.5 to 3.3 m m in d iameter and apparen t ly wi thou t any secreted lining. Cells were vertical, radia l ly symmetr ica l , elongate oval and a r ranged in groups along hor izonta l ly r ami fy ing laterals of the same diameter as the main

In the above figures, the letters outside the cell clusters refer to corresponding vertical burrows, the letters inside the clusters refer to contents of cells as follows :

u : used empty cell from which adults have emerged; ~ : female pupa; p : preplipa ; 1 : large larva; m : medium larva; o : cell with pollen ball; n : nevcly-made, empty cell.

X marks position of the main burrow in the eluster.

Page 4: Social behavior and nests of some nomiine bees in India (Hymenoptera, Halictidæ)

148 SUZANNE W. T. BATRA

burrow (fig. 3 and 4). Vertical burrows surrounding each group (or cluster) of cells and sometimes extending horizontally" beneath t h e m part ial ly isolated the cells from the surrounding soil. Cells, having a smooth, shiny, water repellent lining, measured 8 to 10 mm long and 3.0 to 3.5 mm wide. The necks were 2.0 to 2.5 mm in diameter and cells after oviposition were plugged with 1.5 to 3.5 mm of loose soil. The three complete nests contained : -

NEST 1 . - -Four female bees. Three cell clusters (at depths of 27, 37 and 45 cm) with a total of 24 cells. The lowest cell cluster contained five empty (new .~) cells and a eell with an incomplete pollen ball. The middle cluster had six cells containing brood (1 small larva, 1 medium larva, 2 large larvae, 2 prepupae) and five empty cells. The upper cell cluster contained one female pupa and six empty (old) cells.

NEST 2 (fig. 1).--Six bees. Two cell clusters (at 35 and 44 cm) with 17 cells; the lower cell cluster eontained 13 cells, the upper cluster had four ceils (see.fig. 3 and 4 for contenbs).

NEST 3 . - -Four bees. One cell cluster (at 33 cm) with six cells; the cluster contained four large larvae and two prepupae.

The-yoffngest stages of the brood were found deepest in the nests and t()wards the outside of the individual clusters. Nomia capitata apparent ly enlarges the nest by adding new cells at the per iphery of. old" clusters and more important ly by beginning new clusters below, the upper clusters gradually being abandoned.

Pollen balls (3.0-3.5 mm long, 2.0-2.5 mm wide and 1.7 mm high), were oblong but hemispherical in cross-section, of firm, slightly moist, olive-green pollen. They were attached to one side of the cell near the bot tom, as in N. oxybeloides (fig. 9) and as in Halictinae with vertical cells, but in contrast to many Nomia species in which the pollen occupies the bottom of the cell. The egg was laid in a shallow groove. Probably provisions were made from pollen of Indigofera linifolia, the only plant f rom which foragers were seen taking pollen, although they visited Heliotropium strigosum for neetar.

Larval development is similar to that in other species of Nomia (Caoss and BOHART, 1960 ) . The greenish, apparent ly fluid fecal material was thinly smeared all around the lower third of the cell. Some cells containing prepupae also contained long, vertical stands of fungus mycelium without fruit ing structures and mites were in cells containing larvae. Althohgh Nomia capitata was about to cease activity for at least 8 months, the brood was in various stages of development.

Females from each of the three nests are classified into groups A to E according to the system used by MIEHENER and WILLE (1961) which is summarized as follows : Groups A, B, and D include insemi-

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S O C I A L B E H A V I O R AND N E S T S O F SOME N O M I I N E BEES IN INDIA 149

na ted females , C and E are not insemina ted . Group A females have the la rges t oocyte over 1.0 m m long and nea r ly r e a d y to lay; g roups B and C have the la rges t oocytes be tween 0.3 m m and 1.0 m m long; g roups D and E have la rges t oocyte less t h a n 0.3 m m long. Total wear ( sum of n u m b e r of n icks in bo th fo rewings and re la t ive mand ib le wear) p r o b a b l y ind ica tes age of the female. Mand ibu la r wea r was

d i v i d e d into classes ranging~ f r o m 1 (unworn) to 6 (much worn , sub- apical tooth lacking) .

Nest 1 con ta ined four b e e s as fol lows : (1) gua rd ing bee, g roup A, total wea r 14; (2) pol len collector, g roup A, w e a r 4; (3) forager , g roup C, wea r 2; (4) bee flying f r o m nest, g roup E, wea r l (unworn) .

Nest 2 con ta ined six b e e s : (1) guard, g roup D, wea r 1; (2) and (3) pol len col lectors , bo th g roup A, wear 2 and 4; (4) f emale leaving nest , g roup B, wea r ! ; (5) and (6) t aken f r o m nest, g roups A and E, wear 18 and 1, respect ively .

Nest 3 w i th four bees : (1) pol len collector, g roup B, wear 3; (2) seen to leave nest and go to a n e a r b y flower and take nectar , g roup A, wear 9; (3) and (4) f r o m nest, g roup A, wear 4 and group B (damaged and w e a r u n k n o w n ) .

F r o m the above da ta it m a y be seen tha t the pollen collectors (groups A and B) were also p r e s u m a b l y laying eggs. Th ree addi t iona l poller~ col lectors were t aken on flowers. T h e y were all g roup A wi th total w e a r o f 10, 8, and 3. The wea r of the pollen col lectors f rom nests var ied f r o m 2 to 4 . In each of the nests, however , there was a group A bee tha t was not col,lecting pollen at the t ime and was more w a r n (9, 14, 18) t h a n a n y of the o ther inhab i t an t s . These m a y have been ~ld fem,ales tha t had or ig ina l ly s t a r t ed each nest (i. e., m o t h e r s of the pol len collectors) or older sisteT.s of the pol len col, lec tors ; in any ca, se they a p p a r e n t l y were con t inu ing to ovipo.sit.

The two pol len-col lectors in n e s t 2 were coopera t ive ly p r o v i s i o n i n g a single cel l . (only one cell w i th an incomple te pollen ball was found w h e n the nes t was excava ted) . All pollen col lectors had nec ta r in thei r crops.

F o rag i ng ac t iv i ty began each day be tween 9 and 10 a. m., a f t e r the f lowers of I. l inifolia had been open for a b o u t 30 minu tes . Before the flowers opened, some bees m a d e br ie f r inspec t ion >> fl ights nea r the i r nests. The f lowers r e m a i n e d open and bees fo raged unt i l abou t 2 p . m . Pol len collectors vis i ted 11 to 16 f lowers per m inu t e , and a l ighted on vege ta t ion a t in te rva l s to pack the pollen. F ive pol len- collect ing t r ips las ted 6 , to 28 minu tes . F o r a g e r s were not seen on cloudy, re la t ive ly cool (21 ~ C) days.

The heads of gua rd bees could usua l ly be seen at the e n t r a n c e s of the th ree nes ts con ta in ing severa l f ema les bu t the u n c o m p l e t e d b u r r o w wi th only one bee was not guarded . Guards were r a t h e r t im id but could be induced to s t r ike at a s t r a w wi th their mand ib le s . T h e y were p resen t in the a f t e rnoon (when there was no fo rag ing ac t iv i ty) as well

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150 S U Z A N N E W . T. B A T R A

as in the m o r n i n g bu t were absen t dur ing cloudy, cool wea the r . F o r a g e r s of ten br ief ly g u a r d e d nes ts before leaving. Guards t ha t were dissected be longed to g roup A (wear 1 4 ) a n d D (wear 1). T h r e e of the four f emales in nes t 3 g u a r d e d it in tu rn .

No dif ferences in size (head wid ths ) could be seen be tween bees be longing to the va r ious ova r i an g roups or be tween bees wi th va r ious degrees of w e a r (p robab ly of di f ferent ages).

Nomla (Pseudapis) oxybeloides Smith.

Two nests of this species were excava ted on November 26 (fig. 2) and Decem ber 5, 1064, a th i rd nes t was excava ted May 25, 1965. The first two nests were 7 m apar t , in a level, l i t t le-used sandy p a t h a m o n g sparse vege ta t ion ; the th i rd nes t was at the base of a low sand r idge f o r m i n g an i r r iga t ion channe l in a cu l t iva ted field. All nes ts were in sun t h r o u g h o u t the day.

The entraffces of the nes ts were 3.0 to 3.5 m m in d iamete r , w i thou t tumul i . In the a u t u m n nests , the u n b r a n c h e d vert ical m a i n b u r r o w (diameter" 4.3-5.5 m m ) wa, s l ined wi th a wa t e r repel len t seoret ion s imi la r t o ' t h a t which l ines cells; these two nests r eached dep ths of 23.5 a n d 51.0 cm; the spr ing nest ( bu r row d i ame te r 4.5 m m , wi thou t l ining) was 21.0 cm deep. Ceils (fig. 7 and 0), s imi la r in shape and a r r a n g e m e n t to those of Nomia capitata, m e a s u r e d 9.5 to 15.5 m m long and 5.0 to 5.7 m m wide. T h e necks were 2.7 to 4.0 m m in d i ame te r and a f t e r ovipos i t ion p lugged by 1.5 to 3.0 m m of soil, the inside of the soil p lug (fig. 8) h a v i n g concent r ic r ings in the su r face and being s o m e w h a t depressed in the center . The shiny, w a t e r p r o o f cell l ining could be s epa ra t ed into two ce l lophane- l ike layers . The ver t ical b u r r o w s s u r r o u n d i n g one of the g roups of cells in te r laced below (fig. 5), l a rge ly i so la t ing the c lus te r f r o m the soil; o the r cell c lus ters r e sembled those of N. capitata.

The three nes ts eon ta ined cells as fol lows : - Nest 1 (with 12 females) h a d two cell c lusters . The uppe r cluster ,

11 em below the surface , con ta ined 12 cells (2 e m p t y cells, 4 old cells, 3 large ,larvae, 3 p r epupae ) . The second cluster , 21.0 c m f r o m the sur face (see fig. 6 for con ten ts ) , had 12 ceils.

Nest 2 (with one group B, u n w o r n female) had one cell c lus ter 14.7 em below the sur face . I t con ta ined six cells (3 old, used cells and 2 pre- pupae) . Below the c lus ter was a b l ind b u r r o w ex tend ing to a depth of 51.0 cm.

Nest 3 (new spr ing nest w i th one g roup A, wea r 3, female) had two cells (one egg, one incomple t e ly prov is ioned) wi th 3 ver t ical b u r r o w s descending f r o m a la tera l a t a dep th of 17.0 cm and showed the beg inn ing of c lus ter f o rma t ion . Below the cells was a shor t bl ind bu r row .

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S O C I A L B E H A V I O R AND N E S T S O F S O M E N O M I I N E B E E S IN I N D I A 151

Nomia oxgbeloides is polylectic. The sweet-smelling, yel low pollen ball (fig. 9) was' 5.0 m m long, 4.0 m m wide and 3.0 m m high. T h e brood in the two fall nests consis ted of four larvae tha t had not yet defecated and 15 prepupae , suggest ing that this species m a y over- win ter as p repupae . Nomia oxgbeloides m a y clean old cells and re-use them, for some of the brood in the uppe r cell e luster (nest 1) was younge r t han most of the brood in the lower cluster.

It was not possible to s tudy the soeial behavior of this species because the popula t ions in fall were no longer oviposit ing or foraging a l though t imid guards were present . However , the eomposi t ion O f the popula t ion in nest 1 suggests a degree of social behavior resembl ing tha t of N. capitata. All females in this nest were insemina ted and belonged e i ther to group B (ovaries modera te ly enlarged) or group D (ovaries s lender) . Some of these females m a y have belonged to group A (egg-layers wi th large oocytes) dur ing summer , thei r ovaries in fall having become smal ler t h rough resorp t ion of ooeytes. A high percentage of�9 them were worn, indicat ing tha t they p robab ly had foraged an f f had excavated in the nest. The total wear of group B females ranged f rom 3 to 35; of group D females f rom 4 to 36. The re was no dif ference in size (head widths) between females of var ious ages (degrees of wear) or be tween females wi th different degrees of enlar- gement of the ovaries.

Associates in(~luded m i t e s in m a n y cells conta in ing p repupae or larvae. Whi t e s t rands of myee l ium with black per i theeia and Asper- gillus-like eonidiophores , converging to fo rm a ma t at the bot tom of the cell, were found in several c e l l s conta in ing p repupae (fig. 7), growing on the feces and between the layers of the cell lining.

Nomia nasicana Cockerel l .

Th ree nests of this species were found on October 8, 1964, in an area 3 m square in a south-fac ing grassy e m b a n k m e n t of loose pow- dery clay soil pa r t ly shaded by Acacia trees, next to the Grand T r u n k road where it crosses the Beas River (Punjab) . T h e y had ne i ther tumul i nor guards . One nest, tha t had 3 females enter ing it wi thout pollen, was pa r t ly excavated.

The c i rcu la r ent rance , h idden by grass, was 4.0 tuna in diameter . The unbranched , vert ical b u r r o w w a s fol lowed for 30.0 cm. It was 5.0 m m in d iameter . One empty cell, wi th feces on the upper distal wall, was found at a dep th of 16.0 em. It was horizontal , bi la teral ly symmetr ica l , 11.0 m m long and 5.0 m m wide. The inhabi tan t s of the nest were no.t obta ined for dissect ion; however , this species m a y be sol i tary; the three females in the nest might well have been together only for h iberna t ion .

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152 S U Z A N N E W . T. BATRA

Discussion.

Nomia capitata appears to have reached a level of social organization that is considerably more advanced t h a n is known for any other nomiine bee, and N. oxybeloides may be social to a similar extent.

The social organization of N. capitata differs from that of other. primitively social bees (MIEHENER, 1958) pr imari ly because foragers, probably sisters, apparent ly lay eggs although a more worn individual, perhaps the mother (the nest-founding female) or an older sister, continues to oviposit in the same nest. There does not seem to be inhibition of oviposition of daughters or younger sisters as is common in social insects. If all individuals in the nest are of the same gene- ration (the most worn be being a sister ra ther than the mother), the behavior would be classified as semisocial (MICHENER, 1958), similar to that found in Augochloropsis sparsilis (Vachal) (MmHENEa and LANGE, 1959) except that in N. capitata foragers probably oviposit.

Social behavior in this species may have evolved from colonial behavior sucl{ as exists in Nomia punctulata and N. australica in which females share a burrow but each provisions and oviposits in her own cell cluster. In Nomia capitata and perhaps in N. oxy- beloides one may say that the type of reproduction characteristic of solitary bees probably persists but the females share cell clusters and even cooperatively provision the same cells.

Summary.

Social or semisocial behavior in nomiine bees is here reported for the first time. In Nomia capitata and perhaps in N. oxybeloides, presumed sister bees share cell clusters and cooperatively provision the same cells. Unlike other primitively social bees, the sisters simultaneously forage and oviposit. There is no apparent inhibition of the oviposition of daughters, al though their nest-founding mother or perhaps an older sister appears to stay in the nest and continue to oviposit.

Nesting behavior and nest s t ructure of Nomia capitata and N. oxy- beloides are similar; cells are vertical and clustered. In contrast Nomia nasicana makes horizontal cells scattered along the main burrow.

R~sum~.

Un comportement social est d~erit pour la premiere fois chez des abeilles nomiine. Chez Nomia capitata et probablement N. oxybeloides, il existe des abeilles sceurs qui partagent les cellules et les approvi- sionnent ensemble. Contrairement h d 'autres abeilles h comportement social 'primitif, les sceurs butinent et pondent des ceufs simultan~ment.

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SOCIAL BEHAVIOR AND NESTS OF SOME NOMIINE BEES IN INDIA 153

L e s f i l l e s n e m o n t r e n t p a s d ' i n h i b i t i o n a p p a r e n t e d e p o n t e m a l g r ~ q u e

l e u r m ~ r e a y a n t b~t t i t le n i d , o u p e u t - ~ t r e u n e v i e i l l e s ceu r , r e s t e d a n s

le n i d e t c o n t i n u e ~ p o n d r e d e s c e u f s .

L e e o m p o r t e m e n t e t l a s t r u c t u r e d e s n i d s d e s N o m i a capi tata, N. o x y - beloides e t N o m i a nas i cana s o n t d ~ e r i t s .

Z u s a m m e n [ a s s u n g .

Z u m e r s t e n M a l w i r d f i b e r s o z i a l e s o d e r h a l b s o z i a l e s V e r h a l t e n v o n

B i e n e n a u s d e r U n t e r f a m i l i e d e r N o m i i n e n b e r i c h t e t . B e i N o m i a capi- rata u n d v i o l t e i e h t a u e h b e i N. oxybe lo ides t e i l e n s i e h m e h r e r e B i e n e , n

- - w a h r s e h e i n l i e h S e h w e s t e r n - - e i n u n d d e n s e l b e n Z e l l k o m p l e x u n d

v e r p r o v i a n t i e r e n a u e h d i e Z e l l e n g e m e i n s a m . G a n z i m G e g e n s a t z z u

a n d e r e n s o z i a l p r i m i t i ~ c e n B i e n e n v e r p r o v i a n t i e r e n d i e s e N e s t g e n o s s e n

n i c h t n u r , s o n d e r n l e g e n a u e h E i e r . D i e F ~ i h i g k e i t E i e r z u p r o d u z i e r e n

i s t i n d i e s e n B i e n e n o f f e n b a r n i e h t n n t e r d r i i e k t , o b w o h l s i c h i n j e d e m

N e s t e i n e i e r l e g e n d e s I n d i v i d u u m b e f i n d e t , d i e e n t w e d e r d i e N e s t -

g r f i n d e r i n , o f f e r e i n e / i t t e r e S e h w e s t e r s e i n m u s s . I m N i s t v e r h a l t e n

u n d i n d e r S t r u k t u r d e r N e s t e r s i n d s i c h N. capi ta ta u n d N. oxybe lo ides s e h r ~ i h n l i e h . D i e Z e l l e n s t e h e n v e r t i k a l u n d s i n d i n G r u p p e n a n g e o r d -

n e t . I m G e g e n s a t z d a z u f e r t i g t N. nas icana h o r i z o n t a l l i e g e n d e Z e l i e n ,

d i e e n t l a n g e i n e s T u n n e l s v e r t e i l t s i n d .

LITERATURE CITED

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BATRA (S. W. T.) and MICHENEn (C. D.), 1966. - - Nest and descr ipt ion of a new bee, Systropha punjabensis f rom India. J. Kansas Entomol. Soc., 39, 4 (In press).

CRoss (E. A.) and BOHART (G. E.), 1960. - - The biology of Nomia (Epinomia) trianguli[era with compara t ive notes on other species of Nomia. Univ. Kansas Sci. Bull., 41, 6, p. 761-792.

GUTBXEn (A.), 1915. - - Essai sur la classif ication et sur le d6veloppement des nids des gu6pes et des abeilles. Horae Soc. Ent. Rossieae, 4 t , 7, p. 1-56, pl. I-II.

HmASmMA (Y.), 1961. - - Monographic s tudy of the subfami ly Nomi inae of J apan (Hymenoptera , Apoidea). Acta Hymen., 1, 3, p. 241-303.

MICHENER (C. D.), 1 9 5 8 . - The evolut ion of soeial behav io r in bees. Proc. Tenth Internat. Congr. Entomol. (Montreal) , 2, p. 441-447.

MICHEr~ER (C. D.) and LANC, E (H. B.), 1959. - - Observat ion on the behav ior of Braz i l ian ha l ic t id bees (Hymenoptera , Apoidea) IV. Augochloropsis, wi th notes on ex t ra l imi ta l forms. Am. Mus. Novitates No. 1924, p. 1-41.

MICHENWa (C. D.) and WILLE (A.), 1961. - - The b ionomics of a p r imi t ive ly social bee, Lasioglossum inconspicuum. Univ. Kansas Sci. Ball., 42, p. 1123-1202.

RAYME~T (T.), 1956. - - The lVomia austratica Sin. complex, i ts t axonomy, morpho- logy and biology, wi th the descr ipt ion of a new mut i l l id wasp. The Australian Zoologist, 12, 3, p. 176-200 and 4 plates.

STEPHEN (W. P.), 1959. - - Ma in ta in ing a lkal i bees for a l fa l fa seed product ion Oregon Agr. Exp. Sta. Ball., 568, p. 1-23.