Foraging Polyethism in Odontotermes Formosanus Shiraki

Journal of Advanced Agricultural Technologies Vol. 1, No. 1, June 2014

E. Soleymaninejadian1, B. Z. Ji1, S. W. Liu2, J. J. Yang1, and X. W. Zhang1 1College of Forest Resources and Environment, Institute of Forest Protection Nanjing Forestry University, Nanjing 210037, China 2The Administration Bureau of Dr.Sun Yat-sen，s Mausoleum, Nanjing 210014, China Email: [email protected]; [email protected]; {liushuwen2006, brocadeyang} @126.com; [email protected]

Abstract—South of China is always invaded by showed that workers with different sizes have different Odontotermes formosanus Shiraki. In this paper Polyethism roles in construction behavior. He divided workers into of foragers in O.formosanus Shiraki have been investigated. two small workers (SW1, SW2) and three large workers Foragers of Thirty nests in Nanjing Forestry University (LW1, LW2. LW3) based on a research was done by have been studied, 4782 workers and soldiers captured. We McMahan (1970) [14]. Building behavior of all of them realize that workers with head width of medium (MW) are the main foragers. According to frequency of head width had been analyzed by him. He found that SW1 and LW3 and resulted peaks, foragers have been divided into Very are moderate in abundance at the gallery repair site, they small workers (VSW), Small Workers (SW), Medium are vigorous yet. In spite, the SW2 are the active builders. Workers (MW), Large Workers (LW), and Very Large Also, LW3 are taking part in grooming [15]. As well Workers (VLW). Studying each nest separately showed that McMahan et al. (1984) attested that 96% of small all the above groups are not available in foraging behavior workers are attending in queen chamber on the contrary of one nest. We found three groups of soldiers among 99% of large workers take part in foraging and 70% of foragers, Small Soldiers (SS), Medium Soldiers (MS), and them can be found around swarming sites exit holes [16]. Large Soldiers (LS). In this case, MS were the main groups Campora, and Grace (2004) mentioned that the size plays of soldiers. In addition, foraging polyethism in different tree species has been investigated Cinnamonum camphora was a critical role in tunneling galleries. They analyzed their the main target for this species. At the end it became data using ANOVA and regression analysis. The research obvious that however some of other groups taking part in confirmed that large workers can make wide with less foraging in different nests, but workers and soldiers with segmentation tunnels, while the small worker are good medium head width have a key role in foraging behavior by enough to make tunnels with more segmentation still less this kind of termite. width [9]. Size polyethism is not involved only among the workers, small and large soldiers of N. exitiosus act Index Terms—termite, isoptera, odontotermes formosanus differently in facing with enemies. In this species small shiraki, size polyethism, foragers soldiers are more attracted to the intruders and they use their frontal gland to secret and repel the invaders. In contrast, large soldiers are non combative and flee at the I. INTRODUCTION time of danger. Non combative large soldiers may serve During the evolution of social insects polyethism is as messenger and make the nest ready by raising the one of the key phenomenons [1]. Polyethism makes the danger signals [17], [18]. Furthermore, one study on N. colony works harmoniously. To put it in a nutshell, costalis has shown that soldiers have a key role in polyethism means division of duties among the insect foraging by recruiting other soldiers and large workers to colony member. As Evans (2006) revealed in his paper, the source of food [19]. This phenomenon is called in Nasutitermes exitiosus, worker shifted their work from chemical regulation of polyethism during foraging [20]. foraging to building behavior and vice versa very rarely. It was cleared that large instar workers have bigger This research has shown that duties are somehow divided sternal glands than soldiers, subsequently produce more in different individuals in a fix manner [2]. Recent trail pheromone to recruit more foragers [21], [20]. studies have shown that age [3]-[7] size [8]-[11] and sex In this paper O.formosanus Shiraki has been studied to [12] or combination of them [13] are the fundamental see polyethism among foragers. Foraging polyethism of criteria in polyethism. 30 nests has been studied. Furthermore, polyethism has Size of the termites especially workers is one of the been studied among different tree species. To indicate important morphological characters that confirm the this phenomenon, the head width of foragers measured activity of the workers. Jones (1980) studied the and put into different groups. Then, major groups construction behavior of Nasutitermes costalis. He involving in foraging have been demonstrated.

II. MATERIALS AND METHODS Manuscript received March 12, 2014; revised May 22, 2014.

Nanjing Forestry University (NFU), located in the east To measure head width (exactly after antenna toward of Nanjing, at the foot of Zijin Mountain and east of back side of the head Fig. 1) of the samples, JSZ6S Xuanwu Lake (32° 4′ 50.66″ N, 118° 48′ 41.06″ E), was (20030568) microscope (manufactured by Nanjing the study site. All the termites captured in a sunny day Jiangnan Novel Optics Co.,Ltd) with a camera (JIFEI) between 10Am-2 Pm with temperature of 20, humidity of (manufactured by Nanjing JIFEI Technology Co.,Ltd) 50 and a breeze blowing (0.5 Km/H). Hot damp summers assembled on it has been used. The microscope was and cold damp winters with short spring and fall are the connected with a computer. JIFEI software helped us to main characteristic seasons in Nanjing. Finding shelter measure and capture photos in the computer. Microsoft tubes on the trees considered as a sign of foragers of a Excel has been used to calculate Average and Standard nests. Making a breach in the tube gently (tweezers can Error. After measuring the head width, the data has been be used) and then by the means of smooth painting brush loaded into Microsoft Excel and column charts made samples collected into a plastic container (11 cm in using head width and frequency of them as x and y axis diameter, 4 cm high) with a moist filter paper in it. respectively. Based on the peaks, workers and soldiers Foragers of each nests collected into one plastic container. have been classified into different groups [10], [22]-[24]. After collecting the samples, name of the invaded species For division of the groups we consider a gradual rise recorded. ended with a peak and a gradual down. So a sudden rises Samples taken to the lab and mud particles removed or falls did not consider as groups. Also, lowest peak is from them, then transferred into glass vials containing reflected as starting of one group. FAA (37% formalin: acetic acid: ethanol=6:1:16). In the case of studying polyethism among trees, the Foragers kept into FAA over night. Then replace the nests divided based on the tree they had been hosted. FAA with 70% alcohol. Then, head width measured and similar criteria used to divide the foragers.

Figure 1. The black line is stretched from side to side of head is the place was used to measure head width

foragers with medium head width are the most abundant III. RESULTS ones in the 30 nests. Foragers with Head width of 1.42 are the majority of the foragers. Average of head width Foragers of 30 nests have been studied in this paper. and Standard Error for 30 nests was calculated in Total number of foragers caught was 4782, 4747 workers Microsoft Excel and they were 1.45 and 0.001 and 35 soldiers. As it can be seen in the Fig. 2 head width respectively. of workers vary from 0.4-1.72 millimeters (mm). All the sizes in this paper are based on mm. It is cleared that

Figure 2. Distribution of foragers based on head width measurement. X axis is the head width and Y axis is the number of foragers. Foragers with head width of 0.4-1.26 were classified as Very Small Workers (VSW), 1.27-1.38 Small Workers (SW), 1.39-1.48 Medium Workers (MW), 1.49-1.59 Large Workers (LW), and 1.6-1.72 Very Large Workers (VLW) Borders of each group are highlighted in black.

Based on the peaks in the graphs foragers have been Medium Workers (MW), 1.49-1.59 Large Workers (LW), classified into different groups. Foragers with head width and 1.6-1.72 Very Large Workers (VLW). Percentage of of 0.4-1.26 were classified as Very Small Workers each group can be seen in the Fig. 3. (VSW), 1.27-1.38 Small Workers (SW), 1.39-1.48

Figure 3. Percentages of different groups of workers among foragers of 30 nests in total. As it can be seen less than 1% of foragers are VSW. Most abundant workers among foragers are MW with 41% followed by LW (30%), SW (21%), and VLW with 8%.

Only 0.75 % of all the foragers were soldiers. From 35 they also can be classified as Small Soldiers (SS), 1.24- foraging soldiers, soldiers with head width of 1.34 are the 1.33, Medium Soldiers (MS), 1.34-1.44, and Large most abundant ones (Fig. 4a). Foraging soldiers also Soldiers (LS), 1.45-1.51. Percentages of each instar have were divided into different groups based on peaks in Fig. come in the Fig. 4b. 3, 1.24-1.33, 1.34-1.44, and 1.45-1.51. Based on this,

a b

Figure 4. a) Soldiers head width distribution during Foraging behavior. Border of each group has been illustrated by the number above the columns. b) Percentages of different soldier instars based on head width. As it can be seen MS with 59% are the main soldiers, then LS with 23% and at the end SS with 18 %.

Rest of the nests has been analyzed in same manner Robinia pseudoacaia, Magnolia denudata, Metaseuoia and the results have come in Table I. Result of Studing glyptostroboide, Prunus mume, Catalpa speciosa, Ulmus polyethism among different tree species brought in Table pumila, and Castanea sativa were highly infested by O. II; Cinnamonum camphora, Sophora japonica, formosanus Shiraki (Table II). Liriodendron chinensis, Cunninghamia lanceolata,

TABLE I. STUDY THE POLUETHISM AMONG 30 NESTS SEPERATLY

Nests Total Workers Workers Standard Main Group Number of Soldiers Standard Main Number Number Average Error of Workers Soldiers Average Error Group of Head Width Workers Head Soldiers Soldiers Width 1st 167 167 1.58 0.002 LW - - - - 58 % VLW 42%

2nd 122 120 1.37 0.003 SW 2 1.29 0.01 SS 58% MW 42% 3rd 152 147 1.56 0.003 MW 5 1.48 0.037 SS 4% 20% LW, MS 70% 40% VLW LS 26% 40% 4th 95 95 1.50 0.004 MW - - - - 36% LW 64% 5th 200 200 1.4 0.002 SW - - - - 38% MW 61% LW 1% 6th 128 127 1.52 0.004 SW 1 145 - LS 0.8% MW 15.7% LW 81.1% VLS 2.36% 7th 183 182 1.52 0.004 VSW 1 1.41 - MS 0.5% MW 20% LW 73% VLW 6.5% 8th 111 111 1.42 0.003 SW - - - - 14% MW 84.% LW 2% 9th 55 55 1.44 0.006 SW - - - - 13% MW 71% LW 16% 10th 61 61 1.47 0.01 SW - - - - 24% MW 18% LW 56% VLW 2% 11th 161 159 1.44 0.002 SW 2 1.31 0.03 SS 3% MS MW 83% LW 14% 12th 202 200 1.35 0.02 SW 2 1.35 0.01 MS 76% MW 24% 13th 72 72 1.52 0.005 MW - - - - 17% LW 74% VLW 9% 14th 80 80 1.54 0.004 MW - - - - 10% LW

86% VLW 4% 15th 189 189 1.34 0.003 VSW - - - - 6% SW 92% MW 2% 16th 105 105 1.52 0.003 MW - - - - 10% LW 88% VLW 2% 17th 200 200 1.44 0.002 SW - - - - 1.5% MW 87% LW 11.5% 18th 203 202 1.4 0.005 VSW 1 1.27 - SS 0.5% SW 26% MW 71% LW 2.5% 19th 200 200 1.43 0.003 SW - - - - 15% MW 71% LW 14% 20st 206 199 1.4 0.002 SW 7 1.37 0.011 MS 28% MW 70% LW 2% 21rd 201 201 1.4 0.002 SW - - - - 13% MW 76% LW 11% 22th 180 179 1.5 0.004 MW 1 1.34 - MS 31% LW 62% VLW 7%

23th 208 202 1.48 0.003 SW 6 1.42 0.011 MS 2% 67% MW LS 46% 33% LW 52% 24th 200 200 1.45 0.002 SW - - - - 6% MW 68.5% LW 25.5% 25th 159 157 1.49 0.003 MW 2 1.44 0.015 MS 35.5% LS LW 63.5% VLW 1%

26th 213 212 1.35 0.003 VSW 1 1.32 - SS 1.5%

SW 79% MW 19.5% 27th 234 233 1.61 0.003 SW 1 1.5 - LS 0.5% LW 28.5% VLW 71% 28th 203 202 1.43 0.002 SW 1 1.5 - LS 10.5% MW 77.5% LW 12% 29th 173 172 1.56 0.004 SW 1 1.24 - SS 0.5% MW 7.5% LW 60% VLW 32%

30th 118 117 1.35 0.003 SW 1 1.37 - MS 85% MW 15%

TABLE II. POLYETHISM AMONG DIFFERENT TREE SPECIES

Trees Foragers Number Average Standard Percentages Number Average Standard Percentages Number 0f Head Size Error of Different of Head Error of Different Workers (Workers) (Workers) Groups Soldiers width (Soldiers) Soldiers (Workers) (Soldiers) Groups (Soldiers) C. camphora 1814 1790 1.46 0.001 VSW 24 1.39 0.012 SS 0.2% 16.5% SW MS 19% 62.5% MW LS 40% 21% LW 34% VLW 6.8% S. japonica 1068 1064 1.44 0.001 SW 4 1.36 0.07 SS 15.5% 25% MW MS 59% 50% LW LS 25% 25% VLW 0.5% L. chinensis 623 622 1.45 0.005 VSW 1 1.5 - LS 2% SW 40.5% MW 20% LW 11% VLW 26.5% R. psuedoacaia 288 287 1.52 0.003 VSW 1 1.41 - MS 0.5% MW 16.5% LW 78.5% VLW 4.5% M. denudota 203 202 1.4 0.005 VSW 1 1.27 - SS 0.5%

SW 26% MW 71% LW 2.5% M. 55 55 1.44 0.006 SW - -- - - glyptostroboides 13% MW 71% LW 16% C. lanceolata 202 200 1.35 0.002 SW 2 1.35 0.015 MS 76% LW 24% P. mume 80 80 1.54 0.004 MW - - - - 10% LW 86% VLW 4% C. speciosa 203 202 1.43 0.002 SW 1 1.5 - LS 10.5% MW 77.5% LW 12% U. pumila 173 172 1.53 0.004 SW 1 1.24 - SS 0.5% MW 7.5% LW 60% VLW 32% C. sativa 72 72 1.52 0.005 MW - - - - 17% LW 74% VLW 9%

a b Figure 5. a) Percentage of working foragers is shown in this photo. MW (1.4-1.5) with 41% were the main workers during foraging, LW1 ranked second, then SW2 and finally LW2 and SW1. b) LS1 and SS2 were highly available during foraging behavior.

Another kind of classification was possible without and 1.6-1.7 Large worker 2 (LW2). Considering the looking at peaks and only considering head width. percentage of each group can be seen in the Fig. 5a. If we close our eyes on the two groups with These 9 groups have not seen among foragers of only one percentages of less than one (Very Small Workers nest. In usual only three largest instars have seen in (VSW) ,0.4-1, and Very Large Workers (VLW),1.7-1.8), foragers of one nest. Same criteria has been used to Foragers with 1.2-1.3 can be classified as Small Worker divide soldiers, Foraging soldiers also can be divided into 1 (SM1), 1.3-1.4 Small Worker 2 (SW2), 1.4-1.5 different instars based on head width, 1.2-1.3, 1.3-1.4, Medium Worker (MW), 1.5-1.6 Large Worker 1 (LW1), 1.4-1.5, and 1.5-1.6. Based on this, they also can be

classified as Small Soldiers 1 (SS1), 1.2-1.3, Small of curve. At least two groups can be seen in one nest. As Soldiers (SS2), 1.3-1.4, and Large Soldiers 1 (LS1), 1.4- it can be seen in Fig. 6a, to see small, medium and large 1.5, Large Soldiers 2 (LS2), 1.5-1.6. Percentages of each worker in one nest is 44%. instar have come in the Fig. 5b. As it can be seen in the However foraging polyethism among tree species chart 1.4-1.5 contains 40% of the population. showed that there is no especial relation between a tree Looking at each nest closely is another story; not all of species and foraging polyethism in O.formosanus Shiraki, the groups are available in one nest. For example, in the but in this case also the majority of workers were the first nest only LW and very large workers can be seen ones with medium head width. Chart in Fig. 6b can tell and other groups are not available or have not caught. It us that there was not special group in foragers that can be was interesting that although all the groups of workers seen only in one tree species, the groups randomly are not available in foraging behavior, but there was a distributed among them. According to Table II, the more similarity between column charts of them; when head foragers attack one species the more possibility to see width used as ‘x’ axis and frequency of them as ‘y’ axis different groups. For example, C.camphora is mostly nearly all the charts were bell curve in structure. It means attacked by O.formosanus Shiraki and more groups of always the bulkiest numbers of foragers are in the middle workers can be seen among foragers.

a b Figure 6. a) As it can be seen in 44% of the nests small, medium, and large workers are available, as well as M, L, and VL were available in 17% of the nests. b) Different groups of foragers among the tree species; there is no special pattern in distribution of foragers.

Figure 7. As it can be seen C.camphora with 33% is in top of most favorite food by O.formosanus Shiraki, as well as S.japonica with 21%, L.chinensis and C.lanceolata each one 11%, R.pseudoacaia and M.denudata each one 5%, M.glyptostroboide, P.mume, C.speciosa, U.pumila, and C.sativa each of them 3%

C.camphora (12), S.japonica (8), L.chinensis (4), tubes were empty or the number were a few. C.lanceolata (4), R.pseudoacaia (2), M.denudata (2), O.formosanus Shiraki raid C.camphora in highest M.glyptostroboide, P.mume, C.speciosa, U.pumila, and percentage (Fig. 7). C.camphora is the most favorite trees C.sativa were highly infested by O. formosanus Shiraki. (33% of all the infestations) for O. formosanus. As Appel The numbers in the parentheses are frequencies of et al (2012) said in their paper that 93% of camphor trees infestation by this kind of species. In this case shelter were invaded by this species [25]. However we found tubes were the sign of infestation, however some of the that Magnolia denudate is overrun by O. formosanus

only in 5% of the cases, but Kasseney et al (2011) have noticed that the bigger head width of the foragers is mentioned that most favorite trees for them was M. the more polyethism can be seen in the foraging behavior. denudate (75%) followed by Elaeocarpus It was interesting that nest with big head widths, glabripetalus Merr (41%) [26]. Our result highly Sophisticated and Old nests (1.4-1.7), the division of task suggests that O. formosanus has a very vast range of diet. can be seen more than Naïve and Young nests. This Polyethism analysis among different plant species has result has proven the result released by Holbrook et al done. In contrast with above studies that only test a (2011). They found a significant correlation between limited numbers of plant species, we analyze their diet in colony size and division of labor [27]. Interchanging the foot of Purple Mountain with variety of species ,621 tasks among workers of older, larger colonies were plant species from 383 genera and 118 different families, tremendously low. Moreover, the bigger the size of one including 78 cultivated species. colony, the more division of labor could be and it was Moreover, we classified the nest into four instars based completely independent of colony age. We did the same on the head width, Naïve (1.2-1.4), Young (1.3-1.5), classification of workers among different tree species Sophisticated (1.4-1.6), and Old (1.5-1.7) (Fig. 8a). We (Fig. 8b).

a b Figure 8. a) Division of nests based on foragers head width showed that 40% of nests in NFU are young nests followed by sophisticated ones with 36%, Naïve 17%, and Old 7%. b) Based on the columns C.camphora has hosted by all kinds of nests from Naïve to old ones. However S.japonica is invaded four times by young and two times by sophisticated nests, but L.chinensis is in worst position in the case raiding by variety of nests with different head width.

workers into 5 groups based on peaks. Soldiers of thirty IV. DISCUSSION nests divide into three groups based on peaks. In this case also soldiers with medium head width were the most Our results are showing that size is one of the abundant one in foraging behavior. The results are not important priorities in polyethism or at least in foraging supporting paper published in 1974 by McMahan that LS behavior. As it can be interpreted from Fig. 2 that, are the most active soldiers in the case of foraging [17]. workers with medium head width can be seen in large Studying polyethism in other species such as ant and numbers at site of foraging, but others also are available. bees are more to age and physiological differences than This outcome supports the idea by Crosland et al 1997 size. For example, a paper published by Calderon (1998), that each group of workers do not do only one task in age polyethism had been studied in honey bees and it was Reticulitermes fukienensis [6]. On the other hand, we cleared that physiological and environmental effects are found that workers with larger head width are not the more important than age in polyethism [30]. Cameron most active ones in all the tasks. The result in Fig. 2 and Robinson in 1990 studied effects of Juvenile made it was clear that in the case of foraging groups with hormone (JH) in division of labors in bumble bees. They head width 1.39-1.48 (MW) are the most active workers said however JH has a key role in polyethism in honey (41%) among foragers of thirty nests together. Our data bees but there is no effect on bumble bee. It was due to are matching with a research has done by Miura and weak age polyethism in bumble bees [31]. Ants as well Matsumoto (1995); they said workers with medium head have studied to see polyethism in them. Calabi et al width are the main foragers in the case of gnawing and (1983) found that age the most important criteria in carrying the food balls [28]. As environment has a very Pheidole horten.sis. The polyethism was continues and critical role in instar formation of termites [29]. It was they could not seen any overlap in the tasks of different cleared also that workers with head width of 0.4 can take groups [32]. Further study is needed to focus on part in foraging. In previous paper workers were divided physiological changes in different groups of foragers and into three groups, minor, medium, and major workers and soldiers in the nest. the head width was started at 0.8 [25], [28]. It is reported in this paper that even smaller than this head width can ACKNOWLEGMENTS take part in foraging. In addition, we classified the

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A. L. Watson, and T. Eisner, “Non combative “Immunomodulatory properties of Msenchymal Stem Cells: Cytokine behavior of large soldiers of nasutitermes exitiosus (Hill) an and Factors,” American Journal of Reproductive Immunology, vol. 67: analytical study,” Insectes Sociaux, vol. 1, pp. 103-111, Oct. 1997. pp.1 -8. 2012.

Mr. Soleymaninejadian currently working on polyethism in Mr. Ehsan Soleymaninejadian is reviewer of Biomedical Journals O.formosanus Shiraki. He got Jiangsu Governmet Scholoarship for his (OMICS publication groups). work. His major interests are immunology, immunotherapy, drug delivery using liposom, stem cell therapy.