Original article Expression of body and hair color in three adult castes of the red honeybee Apis koschevnikovi von Buttel-Reepen, 1906 in Sabah, Borneo J Woyke Bee Division, Agricultural University - SGGW, 166 Nowoursynowska, 02-787 Warsaw, Poland (Received 10 December 1996; accepted 15 July 1997) Summary — Apis koschevnikovi bees were collected in Sabah, Borneo. A total of 303 workers from ten colonies were measured in 1995, and 500 workers, 200 drones and ten queens were inves- tigated in 1996. Measurements were made of the width of the dark bands on terga II, III and IV. Drawings of the different body color patterns were made. The workers are light orange with dark brown bands on abdominal terga. Body color patterns were assigned to five groups. The successive groups are gradually darker. Queens and drones have different color patterns and are darker than the work- ers. The upper part of the head and the scutum are black, and the dark brown bands on terga are similar in all three bee castes. However, the light colored body parts such as clypeus, scutellum, and the light abdominal bands are light orange or orange in workers and light brown or reddish brown in the sexuals: queens and drones. The major gene responsible for body color in A koschevnikovi is proposed to be designated as Ko. Hairs covering the thoraces of workers, queens and drones are light orange, gold and gray, respectively. The gene responsible for the color of these hairs is proposed to be designated as Kh. Apis koschevnikovi / honeybee body color / color differences in castes INTRODUCTION Earlier papers concerning the heredity of body color in honeybees were reviewed by Body color is a striking character of any Woyke (1977) and Tucker (1986). Heredity honeybee. The pattern of light (yellow, of body color in A mellifera, and especially orange) and dark (black, brown) color varies the bi-modal frequency distribution was between species, within species and between explained by Woyke (1977, 1978). Woyke the three castes of adults: workers, queens (1995, 1997) also described major body and drones within the same honeybee color genes and their expression in the three colony. castes of adults of four Asian honeybee Tel: (48) 22 471561; fax: (48) 22 471562; e-mail: [email protected] species. A koschevnikovi workers were RESULTS described Koschevnikov (1900) as A by color indica komerunana, by Buttel-Reepen Body (1906) as A koschevnikovi, and by Maa In A koschevnikovi, dense hairs mask the (1953) as A vechti. This bee was rediscov- body color of live bees. Variation in body ered by Tingek et al (1988) as A vechti, and color of worker bees was found. Five dif- Mathew and as A cer- by Mathew (1988) ferent body color types presented in fig1 ana sabahana. A koschevnikovi and queens (and photos on the Internet) were established drones remain undescribed. Therefore, the from the continuous variations. The five expression of body and hair color in the types present the most characteristic repre- three bee castes in a population of A sentative of each group. The names used in koschevnikovi from Tenom, Sabah, was this the color investigated. paper present only popular name, which was the closest to the proper color. Exact determinations of the colors are presented in tables I and II according to MATERIALS AND METHODS Pantone (1995, 1996) color-code. Worker heads are black with some Live A koschevnikovi workers, and drones queens orange-colored parts (fig 1a). The clypeus, were examined in the apiary of the Agricultural Research Station, Tenom, Sabah, Malaysia in genae, labrum and mandibles are orange. March 1995 and October 1996. Together 303 However, in the lightest groups some of the worker bees were sampled from ten colonies in area around the clypeus is orange. In the 1995 and 500 from ten colonies in 1996. All bees darkest group, half of the clypeus and the were in 75% alcohol. Exact measure- preserved are dark. The antennal are ments were made on workers collected in 1995. genae scapes in workers from 1 to The rest served as control for the general color orange groups 3, appearance of worker bees. Two hundred drones orange-brown in group 4 and brown in from nine colonies were investigated. Only ten group 5. The flagella are brown in all work- were examined in detail, 20 queens although ers. Most of the light areas of thorax and more were observed on combs in bee colonies, and in photos. abdomen are light orange or rufus and are lighter than the clypeus. The thorax is cov- Measurements of the width of the dark bands ered by hairs masking the body color. The on terga II, III and IV were made under a stereo- pronotum of the light orange prothorax has microscope with the aid of an eye-piece microm- a darker area on each side (fig 1b and g). eter in arbitrary units equal to 0.0625 mm. Some- Most of the mesothorax is dark (fig 1 g). times the dark hairs of the tomentum had to be The scutum is black in all five How- scratched off to enable exact measurement of the groups. the scutellum is width of the bands. The sum of the width of the ever, light orange, and the dark bands on the three terga was calculated. small plate between it and the scutum, the Color scale code (Pantone, 1995, 1996) was used mesotergum, is a little darker. The two small to define the and hair color. body Representa- pleural plates below the wings, the upper tive specimens of each worker group were drawn. and lower are brown. The Only one specimen of queens and drones was epimera, light is brown with a round dark drawn. Bees under the stereo-microscope were episternum drawn with the aid of a Zeiss drawing appara- brown area in its lower part. The propodeum tus. The drawings were scanned into a computer, is light orange with a distinct marked oval and were with the aid of the Corel- processed curved area. The area is of the same color as DRAW6 These as well as program. drawings the in the three first and some photos of A koschevnikovi workers and propodeum groups, queens are available on the Internet; brown in groups 4 and 5, being a little darker http://alpha.sggw.waw.pl/∼woyke/. in the last group (fig 1b). The legs are differently colored in work- light brown in all other groups. Tarsi are ers from different groups. The outside sur- dark in workers of all the groups. faces of the coxae, trochanters and femurs The abdominal terga are covered with are light orange and the inside surfaces are dense hairs. The terga are light orange with lighter. The femurs of workers from group dark bands on their posterior margins. In 5 are light orange-brown. The tibia is light alcohol-preserved specimens, the light bands orange in workers from groups 1 and 2 and become yellow. The color of the dark bands, according to CMYK coding is about 80, 90, (table III). The mean sums of the three terga 100, 30%. It can be named as black-brown, ranged in the five groups from 1.06 to 3.33 dark walnut or dark brown; however, it is mm. Thus the dark bands in the darkest not rufus (Rinderer et al, 1989). No distinct group were three times wider than in the difference was noticed in the intensity of lightest group. that color in workers of particular groups. However, the pattern (width) differed. The Workers of group 2 were the most numerous 40% of the band on tergum I in group 1 is interrupted. (about population, fig Figure 1b and table III show that the dark 2). Workers of that group were the most common A koschevnikovi at Tenom. The bands are wider in successive terga towards the end of the abdomen (mean tergum II-IV; mode value for that population divided into five was 1.59 mm. The 0.48-0.79 mm, respectively). However, the groups frequency distribution is skewed as well last tergum VI is light orange, except in slightly (fig 2) group 5 where it is of the same dark brown color as the dark bands on the other terga. Older workers, collected at the hive entrance, lost the dark hairs of the tomentum to different degrees. Thus the light areas of the terga were uncovered, and such bees looked very light in color. The sterna are light orange-gold, being a little lighter than the light tergal bands. The sum of the width of the dark bands on terga II, III and IV ranged from 0.688 to 3.688 mm (table III). The mean was 1.88 ± 0.65 mm (x ± SD, n = 303), the variance 0.43 and the median 1.81 mm. The total range was divided into five groups 0.6 mm wide (fig 2). The width of the band of the same tergum was wider in successive groups as the values: for the mean (1.88 mm), and abdomen are much wider than in workers the median (1.81 mm) are both below the (fig 1e). Some variation was observed in center of the range (2.19 mm), being shifted the width of the dark bands. The abdomen of into the direction of the lighter group. The some live queens appeared entirely dark skewness was 0.493. The frequencies of the brown. However, when the queen curved five groups fit to a lognormal distribution; the abdomen, then the light bands were vis- 1.885 ± 0.692 mm (x ± SD), χ2 = 4.305, P = ible.