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Contribution of Insect Pollination to Macadamia Integrifolia Production in Hawaii

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Contribution of Insect Pollination to Macadamia Integrifolia Production in Hawaii IPnsectroceedings PollI natof theIon ofhawaiian Macada emintomologicala In Hawaii society (2015) 47:35–49 35 Contribution of Insect Pollination to Macadamia integrifolia Production in Hawaii Jane M. Tavares, Ethel M. Villalobos, and Mark G. Wright1 Department of Plant and Environmental Protection Sciences, University of Hawaii at Manoa, 3050 Maile Way, Honolulu, HI 96822. 1correspondence: [email protected] Abstract. The honeybee, Apis mellifera, is commonly accepted to be an important pollinator in many agricultural crop systems in Hawaii. However, specific details on the importance of A. mellifera, along with other insect visitors have not been determined for macadamia nut orchards in Hawaii. Reductions in feral honeybee populations in Hawaii attributable to invasions by varroa mites (Varroa destruc- tor) and small hive beetle (Aethina tumida) have resulted in growers becoming dependent on managed bees, requiring increased understanding of the role they play as pollinators. Several parameters determining the contributions of insect pollinators in macadamia nuts were measured: (1) species richness and abun- dance of insects visiting macadamia flowers, (2) the effects of insect pollination in regards to fruit set, fruit retention, fruit size, and weight, and (3) insect pollen removal efficacy based on the mean number of pollen grains an individual insect removed from the stigma while foraging on a macadamia flower. The results from data collected in the study orchard showed that while the order Diptera ranked highest in species richness (9 species), A. mellifera was the most abundant species visiting the macadamia inflorescences (62.7% in abundance, with an average of 17 honeybees seen per 15 minutes compared to 8 flies per 15 minutes). Inflores- cences that were accessible to insects for pollination produced higher fruit sets and yield compared to inflorescences from which insect visitation was excluded. Abundance, foraging behavior and stigma contact, suggested that honeybees are the greatest contributors to macadamia nut pollination over other insects observed in the orchard. The hoverflies (Syrphidae) observed in the orchard may have con- tributed to pollination, but likely to a lesser extent than the honeybees due to the low abundance of the flies. The macadamia nut (Macadamia inte- the history of putative mutual benefits grifolia, Proteaceae) was ranked amongst between the grower and beekeeper, the Hawaii’s top agricultural commodities in importance of honeybees, A. mellifera, to 2011(USDA 2012), producing $38.2 mil the macadamia has not been quantified. In farm gate value in 2011 (NASS 2012). the family Proteaceae, pollination may be The macadamia nut industry is among complex, with various pollinators being the few agricultural industries in Hawaii key for different species, e.g. insects, birds, with a connection between farmers and rodents and wind may all be significant the local beekeepers. The growers of large pollen vectors (Rourke and Delbert 1977, macadamia orchards have long relied Hattingh and Giliomee 1989, Coetzee and on beekeepers to bring in managed bee Giliomee 1985, Wright et al. 1991, Roubik hives for pollination services. Despite 1995), and it is thus relevant to quantify the 36 tavares et al. relative importance of insect pollinators in It is essential to account for various macadamia nuts. factors to determine pollination efficacy The recent onslaught of honeybee pests of an insect, such as the amount of pol- previously absent from Hawaii (Varroa len transferred onto stigmas, and insect destructor (varroa mite) and Aethina behavior when foraging on flowers (Abe tumida (small hive beetle)) have reduced et al. 2011, Wallace et al. 1996). The maca- feral and managed bee colonies dramati- damia stigma is covered in pollen when cally, causing probably in excess of 90% it opens. However, to improve chances of reduction in feral colonies. There are no cross-pollination it is necessary that the previous studies of the density of insect pollen be removed from a newly emerged pollinators required to provide effective stigma (rather than from anthers, as is fruit set in macadamia nut orchards, the case in most plants). Assessing the because feral bees could simply augment ability of insects to remove pollen from pollination by managed hives. Under- the stigma is thus critical. This paper standing the importance of honeybees focuses on evaluating the contribution in macadamia orchards is underscored of insect pollination on macadamia nuts because of the recent impacts to bees in production in Hawaii. Specific objectives the state. of this study were to examine: (1) species Previous research on macadamia nuts richness and abundance of potential insect has highlighted several significant aspects pollinators visiting macadamia flowers, regarding pollination. Macadamia flowers (2) the impacts of insect pollination on are protandrous; a biological feature that macadamia fruit quantity and quality, and is often seen in insect pollinated plants (3) the pollen removal efficacy of insects (Willmer 2011) and thus consistent with while foraging on macadamia flowers. macadamia nut being entomophilous. Quantifying insect abundance and spe- Materials and Methods cies richness in macadamia orchards Study site. This study was conducted is therefore a key aspect to address in at the University of Hawaii Waimanalo macadamia pollination (Heard and Exley Research Station (21°33’ N 157°71’ W) 1994). Comparing fruit set and yields on the island of Oahu, between February from inflorescences visited by insects to 2010 and January of 2011 in Year I and insect-excluded inflorescences can show between March 2011 and March 2012 in the importance of insect pollination in Year II. The study orchard was relatively macadamia nuts (Wallace et al. 1996). small (155 x 74 m), consisting of 91 maca- Macadamia nut fruit quality (mean kernel damia nut trees in seven rows consisting weight) has been shown to improve with of cultivars ‘Kau 344’ and ‘Keaau 660’. cross-pollination by hand (Wallace et al. As is typical of commercial orchards, the 1996, Trueman and Turnbull 1994), and two varieties are interspersed to promote is therefore another important variable cross-pollination. In the study plot ‘Keaau influenced by pollen transfer. The move- 660’ plants were dispersed as every third ment of pollen among plants is a key fac- tree within every third row (Figure 1). tor, with crops that need or benefit from ‘Kau 344’ was used for all experiments, cross-pollination. Urata (1954) reported with the exception of quantifying species that Syrphidae and honeybees were the richness and abundance, in which both most frequently observed flower visitors cultivars were included. Fifty managed on macadamia, but did not quantify their beehives were located on the northeast role in pollination. side of the orchard, between trees in the 494 Insect PollInatIon of Macadamia In Hawaii 37 Hive placement is within rectangle Row 1 Row 2 Row 3 Row 4 Row 5 Row 6 Start of transect census Row 7 ‘Kau 344’ = ‘Keaau 660’ = 50 m (164ft) 495 Figure 1. Map of orchard at the University of Hawaii Waimanalo Research Station 496 Figure 1. Map of orchard at the University of Hawaii Waimanalo Research Station showing transect route. 497 showing transect route. 498 first and second rows. bud turns completely white and the style Monitoring insect foragers for species emerges from the perianth, creating a 499 richness and abundance. Monitoring of loop with the style (Figure 3A). Since the 500 insect foragers was conducted on clear stigma is still enclosed at this time, pollen sunny days, between 6:00 a.m. and 5:00 from that particular floret is unavailable p.m. when trees were in bloom in both to insects; however, insects are able to seasons. Assessment of species richness reach the nectar at the base of the florets (the total number of different species) was by inserting their mouthparts into the slit done through collection and identification in the perianth, which was created by the of insects observed foraging on macada- emerging style. The flower opens by the mia inflorescences. Insects were identified next day making pollen available to insect to species and a reference list compiled. foragers (Figure 3 B). On the second day Voucher specimens were deposited in the of flowering, florets remain attractive to University of Hawaii Insect Museum. In insects, still producing nectar (Figure 3 addition to flower visitor diversity, honey- C), but by day three petals darken to tan bee abundance was also recorded in Year (Figure 3 D) becoming less attractive to I. In Year II, after becoming familiarized insects. By day four petals turn brown and with the insect visitors during Year I, data will soon drop (Figure 3 E). on diversity and abundance of all insect During observations, notes were made pollinators was gathered. on whether the insect being observed was Only inflorescences that were attractive actively collecting pollen or foraging on to insects were included in the abundance the flower’s nectar. Determining whether 18 analysis. This included inflorescences a honeybee was foraging for pollen or one day before opening, and those with nectar was based on the presence of pollen open flowers (Figure 2 B and C). Before on the bee’s pollen baskets and the forag- individual macadamia florets open, the ing location of the bee, coupled with the 38 tavares et al. A 501 B C 502 FigureFigure 2. 2. A) (A) Immature Immature macadamia macadamia inflorescence, inflorescence, B) macadamia (B) macadamia infloresce inflorescencence one day one day before opening with a syrphid, Ornidia obesa, and (C) macadamia inflorescence before opening with a syrphid, Ornidia obesa, and C) macadamia inflorescence that has 503 that has flowers completely open with a honeybee, Apis mellifera. 504 flowers completely open with a honeybee, Apis mellifera. position of the pollen bearing structures in sect abundance: (1) transect insect count, 505 each flower. With flies, the determination where the observer walked briskly down of pollen or nectar foraging was made on the orchard rows and identified and re- the foraging location of the insect alone.
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