And Diurnal Activity of Stomoxys Calcitrans in Thailand

Geographic Distribution of Stomoxyine Flies (Diptera: Muscidae) and Diurnal Activity of Stomoxys calcitrans in Thailand Author(s): Vithee Muenworn, Gerard Duvallet, Krajana Thainchum, Siripun Tuntakom, Somchai Tanasilchayakul, Atchariya Prabaripai, Pongthep Akratanakul, Suprada Sukonthabhirom, and Theeraphap Chareonviriyaphap Source: Journal of Medical Entomology, 47(5):791-797. 2010. Published By: Entomological Society of America DOI: http://dx.doi.org/10.1603/ME10001 URL: http://www.bioone.org/doi/full/10.1603/ME10001

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BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research. BEHAVIOR,CHEMICAL ECOLOGY Geographic Distribution of Stomoxyine Flies (Diptera: Muscidae) and Diurnal Activity of Stomoxys calcitrans in Thailand

VITHEE MUENWORN,1 GERARD DUVALLET,2 KRAJANA THAINCHUM,1 SIRIPUN TUNTAKOM,3 SOMCHAI TANASILCHAYAKUL,3 ATCHARIYA PRABARIPAI,4 PONGTHEP AKRATANAKUL,1,5 6 1,7 SUPRADA SUKONTHABHIROM, AND THEERAPHAP CHAREONVIRIYAPHAP

J. Med. Entomol. 47(5): 791Ð797 (2010); DOI: 10.1603/ME10001 ABSTRACT Stomoxyine ßies (Stomoxys spp.) were collected in 10 localities of Thailand using the Vavoua traps. These localities represented four major ecological settings, as follows: small local dairy farms, large industrial dairy farms, a national park, and one elephant conservation area. Three species of stable ßies were identiÞed in the following proportions: Stomoxys calcitrans (91.5%), Stomoxys indicus (7.9%), and Stomoxys sitiens (0.6%). The number of ßies collected differed signiÞcantly among collection sites (␹2 ϭ 360.15, df ϭ 3, P Ͻ 0.05). The greatest number of stomoxyine ßies was captured in dairy farms. Seasonal and daily activity of S. calcitrans was observed during a 1-yr period at two selected locations (Dairy Farming Promotion Organization of Thailand and Khao Kheow Open Zoo). S. calcitrans was more abundant during the rainy season (MarchÐSeptember), but was not associated with the total rainfall (r2 ϭ 0.0002, P Ͼ 0.05). Peak of daily ßight activity of males S. calcitrans was at 1000 and 1600 h, whereas females showed an increase of activity all along the day until 1600 h. A better understanding of stomoxyine ßy behavior related to patterns of daily activity will facilitate and improve the efÞciency of ßy control measures in private and government sectors.

KEY WORDS Stomoxys spp., distribution, seasonal and diurnal activity, Vavoua traps, Thailand

The genus Stomoxys (Muscidae: Stomoxyinae) con- (LaBrecque et al. 1975, Berry et al. 1976, Smith et al. tains at least 18 described species (Zumpt 1973). They 1985). are obligate blood-sucking insects with some species Stomoxyine ßies can be a severe problem in dairies considered signiÞcant economic pests of livestock and and feedlots, where they breed in moist soil and similar other warm-blooded animals in many parts of the substrates (Meyer and Petersen 1983). Severe biting world (Zumpt 1973, Mullens et al. 1988, Masmeath- activity can result in a reduction in animal weight and athip et al. 2006). Among these, Stomoxys calcitrans milk production. SigniÞcant economic losses as a re- (L., 1758), known as stable ßy, is the most important sult of reduction of anticipated gross weight gain and and cosmopolitan species. Both male and female sto- 30Ð40% decreases in milk yields have been observed moxyine ßies feed on blood, generally once each day. (Hall et al. 1982, Mullens et al. 1988). In the United Adult ßies have a typical ßight range of 1 mile (Ϸ1.6 States, the estimated economic loss to the beef and km). The biology of stable ßies is described elsewhere dairy industry is nearly 400 million dollars annually (Smith et al. 1987). The high number of ßies biting cattle and other affected animals may have a direct inßuence on the epidemiology of communicable 1 Department of Entomology, Faculty of Agriculture, Kasetsart University, Bangkok 10900, Thailand. diseases. Several stomoxyine ßy species have been 2 Centre dÕEcologie fonctionnelle et e´volutive (Unite´ Mixte de implicated as mechanical vectors of anaplasmosis Recherche 5175 Centre dÕEcologie Fonctionnelle et Evolution), Uni- (Anaplasma marginale), trypanosomosis (Trypano- versite´ de Montpellier, 20 Montpellier, France. soma spp.), and different viruses (e.g., bovine leucosis 3 Department of Entomology, Faculty of Agriculture, Kasetsart University, Kamphaengsean, Campus, Nakhon Pathom 73140, Thai- virus, bovine herpesvirus-2, and lumpy skin disease land. virus) (Buxton et al. 1985, Mihok et al. 1995, Torr et al. 4 Division of Biostatistics and Computer, Faculty of Liberal Arts and 2006, Carn 1996). Science, Kasetsart University, Kamphaengsean Campus, Nakhon Stomoxys species have been found to have a wide Pathom 73140, Thailand. 5 Center for Agricultural Biotechnology, Kasetsart University, host range (Warnes and Finlayson 1987). In Egypt, Bangkok 10900, Thailand. domestic donkeys and horses are the preferred hosts 6 OfÞce of Plant Protection Research and Development, Depart- (Hafez and Gamal-Eddin 1959). Warnes (1984) found ment of Agriculture, Ministry of Agriculture and Cooperative, S. calcitrans preferred to feed on cattle and horses in Bangkok 10900, Thailand. 7 Corresponding author: Department of Entomology, Faculty of the United Kingdom. Numerous host factors appear to Agriculture, Kasetsart University, Bangkok 10900, Thailand. inßuence the long-range olfactory responses of stable

0022-2585/10/0791Ð0797$04.00/0 ᭧ 2010 Entomological Society of America 792 JOURNAL OF MEDICAL ENTOMOLOGY Vol. 47, no. 5

Fig. 1. Collection sites of stomoxyine ßies in Thailand.

ßies that include age, size, sex, and nutritional state. dance of Stomoxys species in Thailand, but the study Pheromones produced by cattle also play a role at- was limited to one location and did not compare the tracting some stable ßies (Torr et al. 2006). species diversity among different ecological settings. Surveys of adult stable ßy populations can be as- In this current study, stomoxyine ßy species were sessed using several different techniques. Many stud- surveyed and identiÞed from 10 provinces, represent- ies have employed the use of direct counts or collec- ing four different habitats throughout Thailand. In tions from host animals, especially leg counts to access addition, a monthly stable ßy collection was per- ßy densities (McNeal and Campbell 1981, Berry and formed during a 1-yr period at Dairy Farming Promo- Campbell 1985). Various trapping devices and tech- tion Organization of Thailand, Saraburi Province niques have been developed to collect ßies (Gersa- (SB), and Khao Kheow Open Zoo, Chon Buri Prov- beck and Merritt 1983, Foil and Hogsette 1994). These ince (CB), to study the seasonal and daily activity. The include sticky traps (Williams 1973, Broce 1988) and results of these studies might support more effective also the Vavoua trap (Laveissiere and Grebaut 1990), ßy control programs in private and government sec- originally designed for tsetse ßy collection. This trap tors. has proved very efÞcient at capturing various Stomoxys spp. in many regions of Africa (Holloway and Phelps Materials and Methods 1991, Mihok et al. 1995) and in La Reunion Island (Gilles et al. 2007). Collection Sites. Stable ßy collections were made in Whereas most studies on stomoxyine ßies, particu- 10 geographical locations in Thailand (Fig. 1). Geo- larly S. calcitrans, have been documented in Africa and graphic coordinates and a brief description of collec- the United States, relatively little is known in Thailand tion sites are provided in Table 1. about the presence of different stomoxyine ßy spe- Stable Fly Collection. At each collection site, nine cies, their distribution, and biology. Recently, Mas- Vavoua traps (Laveissiere and Grebaut 1990) were meatathip et al. (2006) described the seasonal abun- randomly placed around sample sites at Ϸ10 m apart. September 2010 MUENWORN ET AL.: DIURNAL ACTIVITY OF Stomoxys calcitrans 793

Table 1. Stomoxyine ﬂy collection sites in Thailand

Dates of Collection site provinces Coordinates Characteristics collection Nong Han, San Sai, Chiang Mai 18Њ 48Ј N, 98Њ 58Ј E Industrial dairy farm, Maejo University; ϳ80 cows April 2007 Wiang Tan, Hang Chat, Lampang 18Њ 17Ј N, 99Њ 28Ј E Thai Elephant Conservation Center, Thung Kwian Forest April 2007 Park; ϳ40 elephants Thai Samakkhi, Wang Nam Khiao, 15Њ 0Ј N, 102Њ 6Ј E Local dairy farm, Wang Nam Khiao District; ϳ40 cows Mar. 2007 Nakhon Ratchasima Mit Taphap, Muak Lek, Saraburi 14Њ 31Ј N, 100Њ 52Ј E Industrial dairy farm: Dairy Farming Promotion Mar. 2007 Organization of Thailand; ϳ200 cows Bang Phra, Si Racha, Chon Buri 13Њ 24Ј N, 101Њ 0Ј E National park, Khao Kheow Open Zoo; a variety of May 2007 natural and resident wild life Pong Kanang, Mueang, Trat 12Њ 13Ј N, 102Њ 30Ј E Local dairy farm: Bo Rai District; ϳ20 cows May 2007 Tha Sao, Sai Yok, Kanchanaburi 14Њ 1Ј N, 99Њ 31Ј E Local dairy farm: Military Development OfÞce; ϳ30 cows April 2007 Huoy Sat Yai, Hua Hin, Prachuap 11Њ 49Ј N, 99Њ 47Ј E Local dairy farm: Dairy Farming Cooperatives; ϳ20 cows April 2007 Khiri Khan Makham Tia Mueang, Surat Thani 9Њ 8Ј N, 99Њ 19Ј E Local dairy farm: Mueang District; ϳ20 cows Sept. 2007 Nam Noy, Hat Yai, Songkhla 7Њ 0Ј N, 100Њ 28Ј E Local dairy farm: Hat Yai District; ϳ20 cows Sept. 2007

For the distribution study, stomoxyine ßies were sam- Elephant Conservation Center (one), and a national pled from 0600 to 1800 h for a 2-d period in each site park (one) (Table 1). during the period MarchÐSeptember 2007 (Table 1). Stomoxyine ßy captures made from March to Sep- A more intensive monthly stable ßy collection was tember 2007 for the distribution study are summarized performed from June 2007 to May 2008 for2dper in Table 2. During this period, a total of 811 specimens, month in two locations, Dairy Farming Promotion representing three different species, was captured. S. Organization of Thailand, SB, and Khao Kheow Open calcitrans (91.5% of captures) was found the most Zoo, CB. For this study, four Vavoua traps were placed prevalent and present in all sites. Stomoxys indicus from 0600 to 1800 h and stable ßies were captured at (7.9%) was present in nine sites, and Stomxys sitiens 2-h intervals. In each study, the ßies were preserved in (0.6%) was found only in three sites (Table 2). 95% ethanol and recorded by date, hour of capture, The greatest proportion of S. calcitrans (30.3%) has and location. The specimens were brought back to the been recorded from the western provinces (Kan- Department of Entomology, Faculty of Agriculture, chanaburi and Prachuap Khiri Khan); 22.9% have been Kasetsart University (Bangkok, Thailand) for the captured from northern provinces (Chiang Mai and identiÞcation according to Zumpt (1973). Air temper- Lumpang); 20.1% from northeast-central (Nakhon ature and relative humidity were also recorded at 2-h Ratchasima and Saraburi); 22.5% from the east (Chon periods. Buri and Trat); and only 4.2% from southern provinces Data Analysis. Spatial and temporal data (time, date, (Surat Thani and Songkhla) (Table 2). place, species, number of specimens, and environmen- Collectively, 81.5% (661) of stomoxyine ßies were tal conditions) were used in the analysis as appropri- captured from dairy farms, 12.9% (105) from the Na- ate. ␹2 test was used to evaluate the differences in tional Park in Chon Buri Province, and 5.5% (45) from number of stable ßies among categories at different the Thai Elephant Conservation Center in Lampang collection sites. The interaction between number of S. Province. Among the eight dairy farms, S. calcitrans calcitrans captured and the yearly total rainfall at in- was found the most abundant (Table 3), representing dividual site was analyzed using variables with biva- 92.4% of the captured ßies, whereas S. indicus repre- riate normal distribution (Kleinbaum et al. 1987). The sented 7% and S. sitiens was relatively rare (0.8%). All GLM procedure in SPSS program package (version 13, three species were recorded from local dairy farms SPSS, Chicago, IL) was used to evaluate the differences between numbers of stable ßies collected at each location. All statistical signiÞcance was set at P Ͻ Table 2. Total numbers of stomoxyine ﬂies collected in 10 0.05. collection sites

Collection sites Stomoxyine ßies Results (province) S. calcitrans S. indicus S. sitiens Total Chiang Mai 129 1 0 130 A survey of stomoxyine ßy species was made in 10 Lumpang 41 4 0 45 locations within six geographical regions of Thailand, Nakhon Ratchasima 9 11 0 20 including the following: 1) Chiang Mai and Lampang Saraburi 140 6 0 146 (north), 2) Nakhon Ratchasima (northeast), 3) Sara- Chon Buri 90 15 0 105 Trat 77 17 0 94 buri (central), 4) Chon Buri and Trat (east), 5) Kan- Kanchanaburi 111 2 1 114 chanaburi and Prachuap Khiri Khan (west), and 6) Prachuap Khiri Khan 114 2 1 117 Surat Thani and Songkhla (south) (Fig. 1). Four po- Surat Thani 28 6 3 37 tential Stomoxys habitats were sought, including in- Songkhla 3 0 0 3 dustrial farms (two sites), local dairy farms (six), Thai Total 742 64 5 811 794 JOURNAL OF MEDICAL ENTOMOLOGY Vol. 47, no. 5

Table 3. Total number of stomoxyine ﬂies among four differ- activity (1000 and 1600 h). The diurnal activity of ent habitats females was less clear, with a continuous increase until 1600 h (Table 5 and Fig. 2). Habitats S. calcitrans S. indicus S. sitiens Total Industrial dairy farm (2) 269 7 0 276 Local dairy farm (6) 342 38 5 385 Discussion National park (1) 90 15 0 105 Elephant conservation 41 4 0 45 There are very few publications on stomoxyine ßy Center (1) species in Thailand (Sucharit and Tumrasvin 1981, Total 742 64 5 811 Masmeatathip et al. 2006). The data obtained from the current study provide better insight on species distribution, bionomics, and diurnal activity of this group of (six), whereas only two species (S. calcitrans and S. biting ßies in Thailand. These data should also prove indicus) were captured from the two industrial dairy useful to help to identify the respective roles of these farms. S. calcitrans was also the predominant species species as nuisance and potential vectors of pathogens. seen at the National Park (85.7%), compared with S. We used the Vavoua trap as a proven method to attract indicus (14.3%). The prevailing species in the elephant and capture these diurnally active ßies. If some sto- center was S. calcitrans (91.1%), followed by very moxyine ßies do have a more nocturnal activity, they small numbers of S. indicus (Table 3). The differences were not captured with this trap. This is perhaps the in numbers of stomoxyine ßies among the different case with S. indicus (Zumpt 1973). In the current habitats were found highly signiÞcant by ␹2 test (␹2 ϭ study, S. calcitrans was found to be the most widely 360.15, df ϭ 3, P Ͻ 0.05). distributed species and strongly associated with both A monthly stable ßy collection was performed at small and large dairy farms in Thailand. Dairy Farming Promotion Organization of Thailand, The comparatively high numbers of stomoxyine SB, and Khao Kheow Open Zoo, CB, from June 2007 ßies, and S. calcitrans in particular, collected in dairy to May 2008. A total of 3,374 and 1,696 S. calcitrans was farms are most likely the consequence of relative high captured at SB and CB, respectively (Table 4). The sex host density for blood-feeding adults and suitable soil ratios (5.6 and 2.1 for SB and CB, respectively) were and environmental conditions for stable ßy larvae to always in favor of males (Table 4). SigniÞcant differ- complete their life cycle. This combination appears to ences in number of stable ßies collected during the 12 play a signiÞcant role in stable ßy abundance. In ad- mo at both locations were obtained (P Ͻ 0.05). In SB, dition, the mixture of manure with silage and spilled stable ßy abundance increased when the collection feed in dairy farms appears a highly favorable medium began, reached a peak in July 2007, and declined from for developing stable ßy larvae (Masmeatathip et al. August 2007 to January 2008. In CB, the seasonal trend 2006). Romero et al. (2006) found that female stable was very similar. The major peak of S. calcitrans abun- ßies are attracted to oviposition sites by stimuli caused dance was observed in May 2008 in SB and in March by bacteria present in manure nearby. 2008 in CB, and the secondary peak was in July 2007 Stable ßy abundance and peak densities are the (Fig. 2). No association between the density of S. consequence of appropriate environmental condi- calcitrans and the yearly total rainfall was observed tions, i.e., moisture, light intensity, rainfall, and tem- (r2 ϭ 0.0002, P Ͼ 0.05). perature to maintain acceptable breeding habitats. The diurnal activity of males and females of S. cal- Mullens and Meyer (1987) observed a single seasonal citrans at SB and CB is shown in Fig. 2. The numbers peak of S. calcitrans, being the most prevalent during of stable ßies collected from different periods of the the summer season from May to June, whereas marked day (0600Ð0800, 0800Ð1000, 1000Ð1200, 1200Ð1400, bimodal and trimodal peaks have been documented in 1400Ð1600, and 1600Ð1800 h) are statistically different other locations in the United States, presumably in- (P Ͻ 0.05). Males show clearly two peaks of diurnal ßuenced by ambient temperatures in more temperate

Table 4. Numbers of male and female S. calcitrans collected at Dairy Farming Promotion Organization of Thailand, Saraburi Province (SB), and Khao Kheow Open Zoo, Chon Buri Province (CB)

No. S. calcitrans at SB No. S. calcitrans at CB Month Males (%) Females (%) Total Males (%) Females (%) Total June 07 235 (84) 46 (16) 281 91 (74) 32 (26) 123 July 07 496 (88) 67 (12) 563 183 (65) 97 (35) 280 Aug. 07 290 (86) 46 (14) 336 129 (69) 58 (31) 187 Sept. 07 96 (65) 52 (35) 148 52 (50) 52 (50) 104 Oct. 07 132 (85) 24 (15) 156 18 (42) 25 (58) 43 Nov. 07 38 (90.5) 4 (9.5) 42 26 (48) 28 (52) 54 Dec. 07 41 (87) 6 (13) 47 18 (49) 19 (51) 37 Jan. 08 17 (61) 11 (39) 28 15 (50) 15 (50) 30 Feb. 08 146 (91) 15 (9) 161 43 (75) 14 (25) 57 Mar. 08 202 (86) 34 (14) 236 420 (75) 143 (25) 563 April 08 516 (83) 109 (17) 625 70 (74) 25 (26) 95 May 08 657 (87.5) 94 (12.5) 751 80 (65) 43 (35) 123 September 2010 MUENWORN ET AL.: DIURNAL ACTIVITY OF Stomoxys calcitrans 795

Fig. 2. Total numbers of S. calcitrans collected at Dairy Farming Promotion Organization of Thailand, SB, and Khao Kheow Open Zoo, CB. climates (Lysyk 1998). In Thailand, Masmeatathip et (1994) reported that the stable ßies can disperse up to al. (2006) reported a wet season peak density of S. 5 km or more in search of blood meals. calcitrans associated with rainfall. In our study, the In Thailand, investigations on the daily activity of majority of adult stable ßies were captured during the Stomoxys species have been limited. Masmeatathip et dry period from March to May. al. (2006) reported a bimodal activity of S. calcitrans, The results from the National Park were conspicu- with the Þrst peak period being between 0800 and ously different from the dairy farm settings. The park 1000 h and another less marked period between 1600 is a large and open area, and most warm-blooded and 1800 h. Besides, there were a number of studies animals may either rest or otherwise be absent during reporting a bimodal activity of S. calcitrans, in Florida the daytime, resulting in poor availability for adult (Simmonds 1944); Mauritius (Kunz and Monty 1976); stable ßies to blood feed. Furthermore, the parkÕs Manaus, Brazil (Charlwood and Lopes 1980); and environment may have lacked sufÞcient and suitable Kansas (Semakula et al. 1989). In contrast, unimodal medium for stable ßy development. In a similar situ- activity pattern on the daily feeding of S. calcitrans was ation, Mihok and Clausen (1996) monitored stomoxy- observed in Uganda (Coaker and Passmore 1958, Har- ine ßies with Vavoua traps on a single day in a forested ley 1965). In our study, there were two distinct daily area of the Nairobi National Park, Kenya, Þnding fairly peaks of male S. calcitrans, occurring at 1000 and robust numbers of species (six), but all in very low 1600 h, and one unclear peak of female at 1600 h. In densities. Brazil, a fairly constant peak of female activity was Similarly, the Thai Elephant Conservation Center reported throughout the day (Charlwood and Lopes appears not to be an ideal ecological setting for sto- 1980). Marked number of S. calcitrans was captured moxyine ßies, considering the low numbers of ßies from the dairy farms compared with the forest setting, captured. Only S. calcitrans was present in any mean- indicating that S. calcitrans is more associated with the ingful number. We also suspect that a feedlot and anthropic ecological setting. livestock area that was located near the elephant cen- Our study has several limitations. First, ßy collec- ter (Ϸ3 km distance) was the primary breeding hab- tions between the different sites were not conducted itat for S. calcitrans in this area. Foil and Hogsette contemporaneously; therefore, seasonal and temper-

Table 5. Numbers of male and female S. calcitrans collected at Dairy Farming Promotion Organization of Thailand, Saraburi Province (SB), and Khao Kheow Open Zoo, Chon Buri Province (CB), from 6 am to 6 pm

No. S. calcitrans at SB No. S. calcitrans at CB Time Males (%) Females (%) Total Males (%) Females (%) Total 0600Ð0800 272 (9.5) 53 (10.4) 325 98 (8.6) 37 (6.7) 135 0800Ð1000 708 (24.7) 70 (13.8) 778 287 (25.1) 57 (10.3) 344 1000Ð1200 448 (15.6) 83 (16.3) 531 180 (15.7) 109 (19.8) 289 1200Ð1400 421 (14.7) 76 (15) 497 153 (13.4) 100 (18.2) 253 1400Ð1600 659 (23) 122 (24) 781 244 (21.3) 144 (26.1) 388 1600Ð1800 358 (12.5) 104 (20.5) 4621 83 (16) 104 (18.9) 287

Percentages are expressed as per day per sex. 796 JOURNAL OF MEDICAL ENTOMOLOGY Vol. 47, no. 5 ature differences between sites may have greatly in- bales: initial associations (Diptera: Muscidae). J. Kansas ßuenced collection numbers. Second, because we Entomol. Soc. 55: 617Ð620. used only one trapping method, it is unclear whether Harley, J.M.B. 1965. Seasonal abundance and diurnal vari- this single method favored one Stomoxys species over ations in activity of some Stomoxys and Tabanidae in another. Third, most of wild mammals are nocturnal Uganda. Bull. Entomol. Res. 56: 319Ð331. and not accessible to diurnal ßies. Perhaps other spe- Holloway, M.T.P., and R. J. Phelps. 1991. The responses of cies of Stomoxys do exist in such areas, but should be Stomoxys spp. (Diptera: Muscidae) to traps and artiÞcial host odours in the Þeld. Bull. Entomol. Res. 80: 51Ð55. captured using light traps during the night or traps in Kleinbaum, D. G., L. L. Kupper, and K. E. Muller. 1987. the canopy (Mavoungou et al. 2007). Use of multiple Applied Regression Analysis and Other Multivariable trapping methods and mark-release-recapture studies Methods, 2nd ed. Duxbury Press, Belmont, CA. of Stomoxys species, particularly S. calcitrans, will be Kunz, S. E., and J. Monty. 1976. Biology and ecology of subjects for further investigation. Stomoxys nigra Macquart and Stomoxys calcitrans (L.) (Diptera: Muscidae) in Mauritius. Bull. Entomol. Res. 66: 745Ð755. Acknowledgments LaBrecque, G. C., D. E. Weidhaas, and T. L. Whitﬁeld. 1975. Graphic models as intermediate steps to computerized This work was supported by the Thailand Research Fund simulations of stable ßy population. Mosq. News 35: 316Ð Organization through the Royal Golden Jubilee Ph.D. Pro- 321. gram (Grant PHD/0037/2549), the Senior Research Scholar Laveissiere, C., and P. Grebaut. 1990. 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