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1 Original Article Journal of Apiculture 31(3) : 183~194 (16) DOI:.17519/apiculture Distribution and Ecological Characteristics of Cocoons of the Solitary Bees Osmia cornifrons and O. pedicornis (Hymenoptera: Megachilidae) Hyung Joo Yoon*, Kyeong Yong Lee, Sun Young Kim, Yeong Mi Kim 1 and Cheon Rak Kwon 2 Department of Agricultural Biology, The National Academy of Agricultural Science, RDA, Wanju 55365, Republic of Korea 1 Silkwork Business Center, Agricultural Resource Management Institute, Sangju 371, Republic of Korea 2 Yecheon Entomology Institute, Yecheon 3682, Republic of Korea (Received 4 September 16; Revised 26 September 16; Accepted 27 September 16) Abstract Osmia cornifrons and O. pedicornis are a cavity-nesting solitary species used mainly as an apple pollinator in Korea. To elucidate the characteristics of those bees for the efficient the pollination of agricultural crops, we investigated the distribution, the number of cocoons per straw, sex ratio, and characteristics of cocoon at different locations in. We collected four species of Osmia spp. consisting of O. cornifrons, O. pedicornis, O. taurus and O. satoi in Yeongwol, Bonghwa and Yechon locations. O. pedicornis was the dominant species, which accounts for 59.6% to 76.4% in the three locations. The mean number of Osmia spp. was varied in different 3 locations. In the number of cocoons per bamboo straw, there were from 7.6 to 8.5 in O. cornifrons and from 6.8 to 7.1 in O. pedicornis at one side straw, and from 16.3 to 16.5 in O. cornifrons and from 13.3 to 13.5 in O. pedicornis at both sides straw. The sex ratio of Osmia spp. were from 1:1 to 1: 2.8 in O. cornifrons and from 1:2.8 to 1:5.5 in O. pedicornis. The sex ratio of O. cornifrons and O. pedicornis was different in 3 locations. The female and male cocoons of O. cornifrons were the long ellipticalshaped and grey-brown color. The width, length and weight of the female cocoon were 5.6.6mm,.2.8mm and 89.5.mg, respectively. The width, length and weight of the male cocoon were 5..4mm, 9..6mm and mg, respectively. The female and male cocoons of O. pedicornis were the long elliptical-shaped and polished dark-brown color. The female cocoon had the width of 6.5.6mm, length of mm and weight of 89.5.mg. In case of the male cocoon, the width, length and weight of were 5..4mm, 9..6mm and mg, respectively. In both species of O. cornifrons and O. pedicornis, the cocoon size of the females was significantly larger than that of the male. Between those two species, the latter had a relatively bigger size than the former. *Corresponding author. yoonhj23@korea.kr 183

2 184 Key words: Solitary bees, Osmia spp., O. cornifrons, O. pedicornis, Distribution, Sex ratio, Cocoon 16,325 (Michener, 7).,,,, (Michener, ) % (Klein et al., 7),, 1 (James and Pitts-Singer, 8). (Free, 1993; Dag and Kammer, 1). Mason bee (Osmia spp.) (Hymenoptera) (Apoidee) (Megachilidae) (Osmia), 339 (Michener, 7). (Osmia cornifrons) (O. pedicornis) 1 1.,,,. 4 5.,., (Torchio, 1989; Bosch and Kemp, ; Lee et al., 16). 7 (Lee and Woo, 1994), 5,,, 3 (Kwon et al., 1997; Lee et al., 2).,, (Yamada et al., 1971; Maeta, 1978; Maeta and Kitamura, 1981; Torchio, 1981, 1985; Bosch, 1994; Bosch and Kemp, 1; Cane, 8). 196 (Maeta and Kitamura, 1964, 1965; Yamada et al., 1971), % (Sekita and Yamada, 1996)., % 5, 1 2, (Maeta and Kitamura, 1981) (Kim, 1999; Lee et al., 2), 199 (Lee et al., ). 2 ( ), % (Yoon et al., 13).,.,.

3 185.,,..,.,,,,., 3 4 (Maeta, 1978). ( 15cm, 7mm) (33.2cm, 7mm). 5,. ( )., ,.,,, Maeta(1978)..,, Lee and Woo(1994) Kwon and Huh(1995),.,, (Lee and Woo, 1994).,.. (Maeta, 1978).,.,, (Bosch, 1994).,,, (scopa) (Maeta, 1978; Torchio and Tepedino, 198).,,,.

4 186 PASW 18 (IBM Inc., 9) One-way ANOVA test, Tukey s HSD, T-test. One-way ANOVA test,,,. ( ). T-test.,,,,, 4, Table %, 13.7%, (O. satoi).4% 3, (O. taurus). 4.3%. 1.3%, 3.9%. Lee et al.(14) %, 21.9% %,.9%, 8.3%. 2, 59.6%, 23.%,.9%,.3% %, 2.%, 13.1%. 3, 59.6~76.4%., 3, 13.7~23.%. 1.6~ ~9.1%..9%.,,.3~.4%.. 7 (,,,,,, ),.,,,, (Lee and Woo, 1994). Lee et al.(2) 3, 5, 5 (45.7%), (41.8%), (11.6%), (.85%), (.3%).,,,,. 4, (52%), (27%), (15%), (5%) (Kim, 1999). 4 (58%), (24%), (17%) 3,,, (Kim et al., 1994),,. Lee et al.(14) ,

5 %, 13.%, 11.2%. 4 (3 ), (2 ) ~76.4%,.. 3.9~2.%, 3.9~13.1%. Table 2., , , ,.6~.9 (One-way ANOVA test: F=1.493, df=2, 249, p=.227). (Fig. 1), 3 14, 1, 2, 4. 9 (17.2%), (13.8%). 1 16, 2, 14, %. 1 (2.2%) 17 (1.1%), 2 13.,., , , Table 1. Rate of Osmia species collected by trap nests at different locations Collected locations n Osmia spp. (%) O. pedicornis O. cornifrons O. taurus O. satoi Mixed Osmia spp. Natural enemies & nesting competition insects (%) Yeongwol Bonghwa Yechon ) Trap nests were placed from late March to early April in. 2) Trap nests contained 5 bamboo straws (Yeongwol, Bonghwa: both sides trap; Yechon: one side trap). 3) n mean number of trap nests supplied. 4) Mixed Osmia spp. means O. pedicornis, O. cornifrons, O. taurus, and (or) O. satoi were located inside one bamboo straw. Empty nests (%) Table 2. The number of cocoons per bamboo straw in trap nests of O. cornifrons and O. pedicornis collected at different locations Collected locations Type of trap nests n O. cornifrons n O. pedicornis Yeongwol One side / both sides Both sides Bonghwa One side / both sides Both sides Yechon One side ) n mean number of trap nests. 2) There were no significant differences in number of cocoons per straw in trap of O. cornifrons and O. pedicornis collect at different locations at p<.5 using One-way ANOVA test. 3) There were significant differences in type of trap nests of O. cornifrons at p<.5 and O. pedicornis at p<.1 using One-way ANOVA test and Tukey s HSD.

6 188 (F=.385, df=2, 563, p=.68). (Fig. 2), 1 12, 7 (31.6%) 8 (29.8%) 61.4%., 12, 7 (33.3%), 8 (16.7%), 9 (14.8%) 64.8%. 7 (19.4), 8 (17.2%), 9 (22.2%) 58.8%., 3,.., Table 2 A 5 A (n=29) (n=171) Rate (%) Rate (%) No. of O. cornifrions cocoons per bamboo straw (one side/both sides, Yeongwol) No. of O. pedicornis cocoons per bamboo straw (one side/both sides, Yeongwol) B 5 B (n=51) (n=54) Rate (%) Rate (%) No. of O. cornifrions cocoons per bamboo straw (one side/both sides, Bonghwa) No. of O. pedicornis cocoons per bamboo straw (one side/both sides, Bonghwa) C 5 C (n=179) (n=36) Rate (%) Rate (%) No. of O. cornifrions cocoons per bamboo straw (one side, Yechon) No. of O. pedicornis cocoons per bamboo straw (one side, Yechon) Fig. 1. Propotional distribution of number of cocoons per bamboo straw of O. cornifrons at Yeongwol (A), Bonghwa (B) and Yechon locations (C). Fig. 2. Propotional distribution of number of cocoons per bamboo straw of O. pedicornis at Yeongwol (A), Bonghwa (B) and Yechon locations (C).

7 머리뿔가위벌(Osmia cornifrons)과 뿔가위벌(O. pedicornis)의 분포 및 고치의 생태적 특성 189 면소통인 경우, 머리뿔가위벌은 7.6~8.5개인데 반하 위벌의 고치수는 7.8±3.7개라고 하였다. 이런 지역 여 뿔가위벌은 6.8~7.1개로 머리뿔가위벌이 뿔가위 및 장소별로 나타나는 고치의 수의 차이는 채집 시기, 벌보다 소통 내 고치수가 많은 것을 알 수 있었으며, 지역의 기후, 식생 등에 따라 암벌의 산란양이 변할 통계적으로도 고도의 유의성이 인정되었다(T-test: 수 있기 때문으로 판단된다(Tepedino and Toricho, t=3.723, p=.1). 양면소통역시, 머리뿔가위벌이 1982; Bosch and Kemp, ). 또한 뿔가위벌류의 종류 16.3~16.5개로 뿔가위벌은 13.3~13.5개보다 소통 내 에 따라 소통 당 고치수의 차이는 뿔가위벌류의 크기, 고치수가 많았다(T-test: t=2.378, p=.21). Lee et 영소 및 산란습성 등에서 나오는 차이일 것으로 생각 al.(14)은 전국 32개의 사과원에서 채집한 머리뿔가 된다. 위벌과 뿔가위벌의 대나무소통 내 고치수는 각각 4.7 뿔가위벌류를 채집할 때 주로 대나무소통을 이용 ±.1개와 4.9±.1개로 뿔가위벌이 머리뿔가위벌보 하는데, 목적에 따라 길이에 의해서 구분되는 단면 다.2개 많은 것으로 보고하였다. 반면에 영월, 봉화, (15cm, 내경 7mm)과 양면(33.2cm, 내경 7mm) 소통을 예천 등 3개 지역에서 채집한 본 실험결과는 머리뿔 사용한다. 단면과 양면 소통의 소통 내 머리뿔가위벌 가위벌은 7.6~8.5개인데 반하여 뿔가위벌은 6.8~7.1개 로 머리뿔가위벌이 뿔가위벌보다 소통 내 고치수가 A 더 많았고, 사과원보다 고치수도 약 2배 이상 많은 것 으로 조사되었다. Lee et al.(16)은 소통 내 머리뿔가 25 Cocoon length (mm) 3 Cocoon length Mud thickness Mud thickness (mm) A B No. of O. pedicornis cocoons per bamboo straw (one side) 6 B 5 Cocoon length Mud thickness Mud thickness (mm) Cocoon length (mm) No. of O. pedicornis cocoons per bamboo straw (one side) Fig. 3. Relation between cocoon length and mud thickness at number of cocoons in bamboo straw of O. cornifrons (A) and O. pedicornis (B). Fig. 4. Cocoons in bamboo straws of O. cornifrons (A) and O. pedicornis (B).

8 19 Table 3. The sex ratio of O. cornifrons and O. pedicornis cocoons collected at different locations Collected locations Sex ration of Osmia spp. (Female: male) n O. cornifrons n O. pedicornis Yeongwol 2,386 1:1 98 1:5.5 Bonghwa 2,4 1: :3.3 Yechon 2,1 1: :2.8 1) Total number of trap nests is 498 in Yeongwol, 466 in Bonghwa, and 585 in Yechon. Table 4. The size of O. cornifrons cocoons collected at different locations Collected locations Yeongwol Bonghwa Yechon Type of trap nest One side / both sides Both sides One side / both sides One Sex n Female Male ab Female Male Female Male b Female Male a 1) n mean number of cocoons surveyed. 2) There was significant differences in sex and cocoon size of O. cornifrons collected at different locations at p<.1 using Turkey's pairwise comparisons test. Width (mm) Length (mm) Weight (mg) 1.9~2. (T-test: t=6.658, p=.1)., (T-test: t=19.239, p=.1). ( ) (Fig. 3)., (Fig. 3A, Fig. 4A),,., (Pearson correlation=-.872, p=.1).,,, (Pearson correlation=-.841, p=.1). Table 3., 1:1, 1:1.9, 1: ~2.8. 1:2.8, 1:3.3, 1: ~5.5. 1:1~1:2.8, 1:2.8~1:5.5. Jeon et al.(), 1:5.8,. Kwon et al.(1997) 1:2.4, 1:12.8, Kim(1999) 6 1:1.7, 1:3.6.

9 191 A B a b, Osmia cornuta (Toricho and Tepedino, 198; Kim, 1999; Bosch and Vicens, 5).,, (Maeta, 1978; Kim, 1999)., 1.~2.8, 2.8~5.5.. a Fig. 5. Female (a) and male cocoon (b) of O. cornifrons (A) and O. pedicornis (B). 1:1.9, 1:8.4, (Lee et al., ),. Bosch and Camp(),. Osmia lignaria (Tepedino and Toricho, 1982). b Table 4 5.8~6.mm,.3~.7mm, 83.8~2.2mg (One-way ANOVA test:, F=.88, df=2, 477, p=.916;, F=1.175, df=2, 477, p=.3;, F=2.216, df=2, 473, p=.1). 5.~5.1mm, 8.9~9.1mm, 44.8~63.5mg (, F=1.171, df=2, 497, p=.311;, F=2.373, df=2, 497, p=.94;, F=5.176, df=2, Table 5. The size of O. pedicornis cocoons collected at different locations Collected locations Yeongwol Bonghwa Yechon Type of trap nest One side / both sides Both sides One side / both sides Both sides One side Sex n Female a a Male A A Female Male Female b b Male 5.5.4B.9.5B Female Male Female a a Male 5.7.4A A ) n mean number of cocoons surveyed. 2) There was significant differences in cocoon size of O. pedicornis collected at different locations at p<.1 using One-way ANOVA test and Turkey HSD. Width (mm) Length (mm) Weight (mg)

10 , p=.6). (Table 4), (, t=-11.66, p=.1;, t= , p=.1;, t=-18.8, p=.1), (, t=.71, p=.484;, t=.634, p=.526;, t=-1.34, p=.1)., ( 5.6.6mm,.2.8mm, 89.5.mg) ( 5..4mm, 9..6mm, mg) (, t=.7, p=.1;, t=29.368, p=.1;, t=27.8, p=.1). Maeta(1978) 5.7~6.mm,.7~11.mm, 4.9~5.2mm, 9.2~9.5mm,..,. (Fig. 5Aa, 5Ab).,, (Fig. 4A). (Table 5), 6.4~6.6mm, 12.6~13.1mm, 139.4~158.4mg (, F=3.64, df=2, 416, p=.58;, F=8.71, df=2, 416, p=.1;, F=11.273, df=2, 416, p=.1). 5.5~5.8mm,.9~11.3mm, 9.9~3.3mg (, F=9.7, df=2, 4, p=.1;, F=16.814, df=2, 4, p=.1;, F=1.933, df=2, 4, p=.146). (Table 5),, (, t=-1.729, p=.84;, t=-23.17, p=.21;, t=-1.336, p=.182), (, t=- 2.65, p=.1;, t=-3.996, p=.1;, t=-2.336, p=.1). (Table 5), ( 6.5.6mm, mm, mg) ( 5.6.4mm, mm, mg) (, t=32.67, p=.1;, t=38.275, p=.1;, t=39.375, p=.1).. 6.~6.4mm, 12.~12.6mm, 5.2~5.3mm, 9.2~9.5mm (Maeta, 1978),.,.,.. (Fig. 5Ba, 5Bb).,, (Fig. 4B)....,,,,.,, 3,,,, 4, ~76.4%..

11 193, 7.6~8.5, 6.8~ ~16.5, 13.3~13.5., 1.~2.8, 2.8~5.5., 5.6.6mm,.2.8mm, 89.5.mg. 5..4mm, 9..6mm, mg mm, mm, mg, ( 5.6.4mm, mm, mg).. ( : PJ1). Bosch, J Improvement of field management of Osmia cornuta (Latreille) (Hymenoptera, Megachilidae). Apidologie 25: Bosch, J. and N. Vicens. 5. Sex allocation in a solitary bee: do females behave in agreement with Fisher's theory? Behav. Ecol. Sociobiol. 59: Bosch, J. and W.P. Kemp. 1. How to manage the blue orchard bee, Osmia lignaria, as an orchard pollinator. Sustainable Agriculture Network, Washington, DC. Bosch, J., F. Sgolastrac and W.P. Kemp.. Timing of eclosion affects diapause development, fat body consumption and longevity in Osmia lignaria, a univoltine, adultwintering solitary bee. J. Insect Physiol. 56: Cane, J.H., 8. Bees (Hymenoptera: Apoidea: Apiformes). Encyclo. Entomol. 2: Dag, A. and Y. Kammer. 1. Comparison between the effectiveness of honeybee (Apis millifera) and bumblebee (Bombus terrestris) as pollinators of greenhouse sweet pepper (Capsicum annuum). Am. Bee. J. 141: Free, J.B Insect pollination of crops. 2nd edn. Academic Press, London. IBM PASW Statistics PASW Core System User s Guide, IBM inc. USA. James, R.R. and T.L. Pitts-Singer. 8. Bee Pollination in Agricultural Ecosystems. 232 p, Oxford University Press, New York. Jeon, S.J., J.T. Kim and N.W. Kim.. A study on nesting behaviour of Osmia spp. (Hymenoptera, Insecta) from Korea. J. Basic Sci. Res. Kyungsan Univ. 4(1): Kim, I.S Studies on the ecological characteristics and utilization of Osmia spp. as pollinators pp. A thesis for the master degree of Kongju National University. Kim, I.S., J.C. Yun, G.B. Eum, G.M. Choi, S.W. Lee and Y.J. Kwon Study of usage of Osmia cornifrons in apple orchard. Annual reseach report. pp NAAS, Wanju, Korea. Klein, A.M., B. Vassiere, J.H. Cane, L. Steffan-Dewenter, S.A. Cunningham, C. Kremen. 7. Importance of crop pollinators in changing landscapes for world crops. Proc. Royal. Soc. London. B. 274: Kwon, Y.J. and E.Y. Huh A new species of Osmia mason bees from Korea, as a potential pollinator of fruit trees. Korean J. Apicult. : Kwon, Y.J., S.J. Suh, E.Y. Huh, Y.S. Yeo, S.J. Suh and J.H. Chung Utilization of pollinating insects in orchard. 116 pp. Institute of Agricultural Science in Gyungbook National University. Lee, H.S. and K.S. Woo Genus Osmia (Hymenoptera; Megachilidae) from Korea. Korean J. Apicult. 9: Lee, K.Y,. H.J. Yoon, K.S. Lee and B.R. Jin. 16. Development and mating behavior of Osmia cornifrons (Hymenotpera: Megachilidae) in the constant temperature. J. Asia Pac. Entomol. 19: Lee, K.Y,. H.J. Yoon, I.G. Park, C.R. Kwon and S.C. Lee.. Survey on the Current Status of Mason Bees in Apple Orchard of Korea. Korean J. Apicult. 25: Lee, K.Y,. S.B. Lee, I.G. Park, P.D. Kang and H.J. Yoon. 14. Distribution Status of Mason Bees, Osmia spp. (Hymenotpera: Megachilidae) in Apple Orchard of Korea. Korean J. Apicult. 29: Lee, S.W., K.H. Choi, D.H. Lee, D.A. Kim, H.K. Ryu and Y.I. Lee. 2. Distribution and collection of Osmia bees in the mountain areas of Korea. Korean J. Appl. Entomol. 41: Maeta, Y Comparative studies on the biology of the bees of the genus Osmia in Japan, with special reference to their management for pollination of crops (Hymenoptera, Megachilidae). Bull. Tohoku Natl. Agrci. Exp. Stn. 57: Maeta, Y. and T. Kitamura Studies on the apple pollination

12 194 of Osmia. I. Idea and present conditions in utilizing Osmia as pollinators of apples in Japan. Kontyu 32: Maeta, Y. and T. Kitamura Studies on the apple pollination of Osmia. II. Maeta, Y. and T. Kitamura Pollinating efficiency by Osmia cornifrons (Radoszkowski) in relation to required number of nesting bees for economic fruit production. Honeybee Sci. 2: Michener, C.D.. The bees of the world. Johns Hopkins University Press, Baltimore. Michener, C.D., 7. The bees of the world. Johns Hopkins University Press, Baltimore. Sekita, N., T. Watanabe and M. Yamade Population ecology of Osmia cornifrons (Hymenoptera: Magachilidae) in natural habitats. Bull. Aomori Apple Exp. Sta. 29: Tepedino, V.J. and P.F. Torchio Temporal variability in the sex ratio of a non-social bee, Osmia ligtiaria propinqua: extrinsic determination or the tracking of an optimum? Oikos. 38: Torchio, P.F Field experiments with Osmia lignaria propinqua Cresson as pollinator in almond orchards: II, 1976 studies (Hymenoptera: Magachilidae). J. Kansas Entomol. Soc. 54: Torchio, P.F Field experiments with the pollinator species, Osmia lignaria propinqua Cresson as in apple orchards: V, , methods of introducing bees, nesting success, seed counts, and fruit yields (Hymenoptera: Magachilidae). J. Kansas Entomol. Soc. 58: Torchio, P.F Biology, immature development, and adaptive behavior of Stelis montana, a cleptoparasite of Osmia spp. (Hymenoptera: Megachilidae). Ann. Entomol. Soc. Am. 82: Torchio, P.F. and V.J. Tepedino Sex Ratio, Body Size and Seasonality in a Solitary Bee, Osmia lignaria propinqua Cresson (Hymenoptera: Megachilidae). Evolution 34(5): Yamada, M., N. Oyama, N. Sekita, S. Shirasaki and C. Tsugawa The ecology of the megachilid bee Osmia cornifrons and its utilization for apple pollination. Bull. Aomori Apple Exp. Sta. 15: 1-8 (in Japanese). Yoon, H.J., K.Y. Lee, I.G. Park, A.E. Kim, Y.M. Kim and P.D. Kang. 13. Current status of insect pollinators use for horticultural crops in 11. Korean J. Apicult. 28: 9-18.

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