Name: evaluation of the productivity of social wasp colonies (vespinae) and an introduction to the traditional japanese vespula wasp hunting technique
Uploaded: 2019-09-11T13:00:00
Description: Watch this Scientific Journal Video about Evaluation of the Productivity of Social Wasp Colonies Vespinae and an Introduction to the Traditional Japanese Vespula Wasp Hunting Technique at JoVE.com

The author would like to thank Katsuyuki Takahashi, Hiroo Kobayashi, Fumihiro Sato, Daikichi Ogiso, Toshihiro Hayakawa, and Hisaki Imai for teaching him the traditional wasp hunt method. The author would like to offer special thanks to Kevin J. Loope and Davide Santoro for carefully proofreading the manuscript. The author is grateful to Masato Abe, Yasukazu Okada, Yuichiro Kobayashi, Masakazu Shimada, and Koji Tsuchida for their discussion. The author wants to thank Yuya Shimizu and Haruna Fujioka for their technical assistance with evaluating colony productivity. The author would like to thank Tsukechi black bee club for supporting the video shooting. The author wishes to thank three anonymous reviewers for their comments on an early version of this paper. This study was supported in part by Takeda Science Foundation, Fujiwara Natural History Foundation, Funding of the Nagano Society for The Promotion of Science, Shimonaka Memories Foundation, Takara Harmonist Fund, and the Dream Project by Come on UP, Ltd.

1. Evaluation of Colony Productivity
<ol>
	<li>Estimation of the number of cells per comb
	<ol>
		<li>Separate the combs one by one. Sweep away all adult wasps from the comb and pull out all larvae and pupae from the cells with tweezers.</li>
		<li>Measure the square measures of 10 randomly chosen cells per a comb, by using imaging software (e.g., Image J version 1.48, see http://imagej.nih.gov/ij/).
		<ol>
			<li>Take a picture with the scale reference so that all the cells are pictured from ...

<ol>
	<li>Davies, N. B., Krebs, J. R., West, S. A. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=.">.</a> An introduction to Behavioural Ecology. , (2012).</li><li>Tibbetts, E. A., Reeve, H. K. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Benefits+of+foundress+associations+in+the+paper+wasp+Polistes+dominulus:+increased+productivity+and+survival,+but+no+assurance+of+fitness+returns.">Benefits of foundress associations in the paper wasp Polistes dominulus: increased productivity and survival, but no assurance of fitness returns.</a> Behavioural Ecology. 14, 510-514 (2003).</li><li>Mattila, H. R., Seeley, T. D. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Genetic+Diversity+in+Honey+Bee+colonies+Enhances+Productivity+and+Fitness.">Genetic Diversity in Honey Bee colonies Enhances Productivity and Fitness.</a> Science. 317, 362 (2007).</li><li>Weber, N. 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An Account of the Biology and Natural History of Social and Solitary Wasps, with Particular Reference to Those of the British Isles. , (1973).</li><li>Matsuura, M., Yamane, S. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=.">.</a> Comparative Ethology of the Vespine Wasps. , (1984).</li><li>Greene, A. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Production+schedules+of+vespine+wasps:+an+empirical+test+of+the+bang-bang+optimization+model.">Production schedules of vespine wasps: an empirical test of the bang-bang optimization model.</a> Journal of Kansas Entomological Society. 57 (4), 545-568 (1984).</li><li>Cole, B. J. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Multiple+mating+and+the+evolution+of+social+behavior+in+the+Hymenoptera.">Multiple mating and the evolution of social behavior in the Hymenoptera.</a> Behavior Ecology Sociobiology. 12, 191-201 (1983).</li><li>Goodisman, M. A. D., Kovacs, J. L., Hoffman, E. A. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=The+significance+of+multiple+mating+in+the+social+wasps+Vespula+maculifrons.">The significance of multiple mating in the social wasps Vespula maculifrons.</a> Evolution. 61 (9), 2260-2267 (2007).</li><li>Greene, A., Ross, K. G., Matthews, R. W. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Dolichovespula+and+Vespula.">Dolichovespula and Vespula.</a> The Social Biology of Wasps. , 263-305 (1991).</li><li>Yamane, S., Yamane, S. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Investigating+methods+of+dead+vespine+nests+(Hymenoptera,+Vespidae)+(Methods+of+taxonomic+and+bio-sociological+studies+on+social+wasps.+II).">Investigating methods of dead vespine nests (Hymenoptera, Vespidae) (Methods of taxonomic and bio-sociological studies on social wasps. II).</a> Teaching Materials for Biology. 12, 18-39 (1975).</li><li>Loope, K. J. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Matricide+and+queen+sex+allocation+in+a+yellowjacket+wasp.">Matricide and queen sex allocation in a yellowjacket wasp.</a> The Science of Nature. 103 (57), 1-11 (2016).</li><li>Matsuura, M. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=.">.</a> Social Wasps of Japan in Color. , (1995).</li><li>Foster, K. R., Ratnieks, F. L. W., Gyllenstrand, N., Thoren, P. A. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Colony+kin+structure+and+male+production+in+Dolichovespula+wasps.">Colony kin structure and male production in Dolichovespula wasps.</a> Molecular Ecology. 10 (4), 1003-1010 (2001).</li><li>Loope, K. J., Chien, C., Juhl, M. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Colony+size+is+linked+to+paternity+frequency+and+paternity+skew+in+yellowjacket+wasps+and+hornets.">Colony size is linked to paternity frequency and paternity skew in yellowjacket wasps and hornets.</a> BMC Evolutionary Biology. 14 (1), 1-12 (2014).</li><li>Nonaka, K. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Cultural+and+commercial+roles+of+edible+wasps+in+Japan.">Cultural and commercial roles of edible wasps in Japan.</a> Forest Insects as Food: Humans Bite Back. Proceedings of a workshop on Asia-Pacific resources and their potential for development. , 123-130 (2010).</li><li>Yamane, S., Funakoshi, K. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=The+unique+ecology+of+Vespula+shidai+amamiana+and+the+origin+of+distribution.">The unique ecology of Vespula shidai amamiana and the origin of distribution.</a> Ecological Society of Japan. Biodiversity of the Nansei Islands, its formation and conservation. , (2015).</li><li>. R: The R Project for Statistical Computing Available from: <a target="_blank" href="https://www.R-project.org/">https://www.R-project.org/</a> (2018)</li><li>Saga, T., Kanai, M., Shimada, M., Okada, Y. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=Mutual+intra-+and+interspecific+social+parasitism+between+parapatric+sister+species+of+Vespula+wasps.">Mutual intra- and interspecific social parasitism between parapatric sister species of Vespula wasps.</a> Insectes Sociaux. 64 (1), 95-101 (2017).</li><li>Van Huis, A., et al. <a target="_blank" href="http://www.ncbi.nlm.nih.gov/entrez/query.fcgi?db=PubMed&cmd=Search&doptcmdl=Citation&defaultField=Title+Word&term=.">.</a> Edible insects: future prospects for food and feed security. , (2013).</li></ol>

The colony productivity of bees, ants, and wasps has been estimated previously by the number of workers and cells in nests or by the weight of the nests3,9,10. This study shows that the estimate of the number of meconia provides a better estimate of the overall number of individuals produced (i.e., a better indicator of colony productivity). In fact, it was found that, for both worker and queen combs, the number of meco...

Every species is thought to develop an optimal strategy for survival and reproduction among a vast array of possible strategies. In natural selection, individuals with traits that maximize an individual&#39;s reproductive success will leave more offspring (and genes) to the next generation. Therefore, the number of offspring produced by an individual can be used as an indicator of the individual&#39;s relative evolutionary fitness. In a given ecological context, the comparison of the number of offspring produced relative to alternative behavioral strategies can help researchers predict the best strategy for optimizing fitness1.
Social Hymenoptera (such as wasps, bees, and ants) have a system of three different castes, which are workers (sterile females), queens (gynes), and males1. Only new queens (gynes) and males count toward fitness in social Hymenoptera. Worker production does not directly contribute to fitness since the worker is infertile. On the other hand, a queen that can produce a higher colony productivity (such as a higher number of total cells or a heavier nest) is considered to have a higher fitness in social Hymenoptera, regardless of the number of actually produced new queens and males (see, e.g.,Tibbetts and Reeve2 and Mattila and Seeley3). In general, it is difficult to precisely count the number of offspring produced by a colony of social Hymenoptera. In fact, the queens of many social insects live for more than 1 year (e.g., leaf-cutter ant queens can live &#62;20 years4 and honeybee queens may live for 8 years5). In addition, one queen may produce thousands of reproductive offspring over the course of several weeks or months, even in annual species of genera Vespa and Vespula6,7,8. Furthermore, the lifespan of workers is shorter than that of their mother queen, and workers often die away from their nests. Hence, even if one could accurately count all adults in a nest at any given point in time, such a count would not accurately depict the number of offspring produced. Therefore, the number of offspring produced has been roughly estimated from the size of the nest, the number of workers in the nest, or the weight of the nest at a given point in time3,9,10. The number of larval cells could result in an overestimation of the offspring production when some cells are empty. The same method could also result in a potential underestimation of the offspring production because combs of small cells that contain worker brood can produce two or three cohorts of larvae6,7,11.
The first aim of this work is to provide an improved method for estimating vespine wasp colony productivity in terms of the number of adults produced. Yamane and Yamane suggested that the best way to estimate the number of offspring produced by a colony is to count the meconia in the nest12. The meconia is the fecal pellet comprising larval cuticle, gut, and gut contents that a larva leaves in its cell when pupating (Figure 1A). The total number of meconia produced per comb is calculated by multiplying the total number of cells present by the average number of meconia per cell. There are often multiple layers of meconia in a cell, and each meconia indicates that an individual successfully pupated in that cell6,11 (Figure 1B). When estimating the mean number of meconia per cell, if the number of cells examined is small (a small sample size), the standard error (SE) increases, and as a result, the error for the total number of meconia per comb becomes higher than if the sample size was larger. The SE of the mean (SEM) is a measure of the dispersion of sample means around the population mean. Therefore, in this study, I&#160;focus on the SEM of the number of meconia per cell to estimate the population (the number of adults produced) from the sample mean (the average number of meconia per cell). This study attempts to determine how many samples are required to obtain an SE rate of less than 0.05 per cell. To do this, a numerical simulation is performed with real data on the number of meconia per comb, to determine the minimum sample size (for both worker and queen combs) needed to estimate this value accurately within the defined SE of 0.05.
Vespine wasp colonies live in concealed nests (underground or aerial) composed of multiple horizontal combs, built in series from top to bottom6,7,11. The average size of the cells increases from the first (top) to the last (bottom) comb. In the bottom combs, a sudden shift in the average cell size can be seen. These wider cells are built for the development of new queens. Hence, a more accurate estimate of colony productivity (i.e., the number of individuals produced) can be obtained when the total number of meconia in the worker cells (small cells) and queen cells (large cells) are considered. In order to estimate fitness at the colony level, researchers could estimate the number of queens produced and focus on the meconia in the queen cells alone. As for reproductive males, these are reared either in worker or queen cells, depending on the species. Thus, it may be difficult to estimate the male production of a colony, except in species where males have a third, unique cell size13 (e.g., Dolichovespula arenaria).
The second aim of this work is to present a useful technique for locating wild vespine wasp colonies in the field and transplanting them into laboratory nest boxes. Although some researchers obtain wasp nests from pest control calls (i.e., people reporting them as pests14,15), this method is not always possible or desirable. Researchers might need to collect nests in wild and inhabited areas where pest controllers do not operate, or to conduct their research by more flexibly obtaining nests at specific times. Interestingly, people living in the mountainous areas of central Japan traditionally collect and rear wasps (Vespula shidai, Vespula flaviceps, and Vespula vulgaris) for food. Therefore, collecting and artificial rearing techniques for these wasps are well developed in those areas17.
This paper also summarizes the methods employed to rear Vespula wasps. The experimental organism for this study was V. shidai, a social, ground-nesting wasp inhabiting western Asia and Japan. V. shidai possesses the largest colony size among all Japanese vespine wasps, with a total of 8,000 to 12,000 cells per nest, with a maximum of 33,400 cells14,18. Workers of V. shidai have an average wet weight of 67.62 &#177; 9.56 mg. Males are usually reared in worker cells; in contrast, new queens are reared in specially constructed, wider queen cells14.
<img alt="Figure 1" class="xfigimg" src="/files/ftp_upload/59044/59044fig1.jpg" /> 
Figure 1: Meconium in a larval cell. (A) Cross section of a comb of Vespula shidai. Meconia is indicated by red arrows. (B) Two meconia are layered. Each blue arrow indicates one meconium. <a href="https://www.jove.com/files/ftp_upload/59044/59044fig1large.jpg" target="_blank">Please click here to view a larger version of this figure.</a>

<table border="0" cellpadding="0" cellspacing="0" style="width:572px;" width="573">
	<tbody>
		<tr height="21">
			<td height="21" style="height:21px;width:216px;">cuttlefish</td>
			<td style="width:71px;">Any</td>
			<td style="width:119px;"></td>
			<td style="width:167px;">fresh/ as a bait</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:216px;">dace</td>
			<td style="width:71px;">Any</td>
			<td style="width:119px;"></td>
			<td>fresh/ as a bait</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:216px;">chichken heart</td>
			<td style="width:71px;">Any</td>
			<td style="width:119px;"></td>
			<td>fresh/ as a bait</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:216px;">plastic bag (polyethylene)</td>
			<td style="width:71px;">Any</td>
			<td style="width:119px;"></td>
			<td>as a flag</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:216px;">bamboo skewer</td>
			<td style="width:71px;">Any</td>
			<td style="width:119px;"></td>
			<td></td>
		</tr>
		<tr height="42">
			<td height="42" style="height:42px;width:216px;">industrial sewing thread</td>
			<td style="width:71px;">FUJIX Ltd.</td>
			<td style="width:119px;">King polyester, No.100</td>
			<td></td>
		</tr>
		<tr height="42">
			<td height="42" style="height:42px;width:216px;">paint marker pen</td>
			<td style="width:71px;">Mitsubishi pencil</td>
			<td style="width:119px;">UNI, POSCA, PC5M</td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:216px;">fishing rod</td>
			<td style="width:71px;">ANY</td>
			<td style="width:119px;"></td>
			<td></td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:216px;">carrying box</td>
			<td style="width:71px;"></td>
			<td style="width:119px;"></td>
			<td>made of wood</td>
		</tr>
		<tr height="21">
			<td height="21" style="height:21px;width:216px;">nest box</td>
			<td style="width:71px;"></td>
			<td style="width:119px;"></td>
			<td>made of wood</td>
		</tr>
	</tbody>
</table>

evaluation of the productivity of social wasp colonies (vespinae) and an introduction to the traditional japanese vespula wasp hunting technique

For vespine wasps, colony productivity is typically estimated by counting the number of larval cells. This paper presents an improved method that enables researchers to estimate more accurately the number of adults produced, counting the number of meconia (the stools left in the cells by wasp larvae when pupating into adults, per 100 cells) in each comb. This method can be applied before or after colony collapse (i.e., in active or inactive nests). The paper also describes how to locate wild Vespula wasp colonies by &#34;flagging&#34; wasp baits and chasing the wasp collecting them, using a method traditionally performed by local people in central Japan (as illustrated in the associated video). The Vespula chasing method described has several advantages: it is easy to reinitiate the chase from a point where the forager flying back to the nest was lost, and it is easy to pinpoint the nest location as marked wasps often lose their flag at the nest entrance. These methods for estimating colony productivity and collecting nests can be valuable for researchers studying social wasps.

One goal of this study was to determine how many samples are required to obtain an SEM of the number of meconia per cell which is less than 0.05. In this study, a comb with an average cell size of &#60;20 mm2 was defined as a worker comb, whereas larger combs were defined as queen combs. I&#160;counted the number of cells for queen combs and worker combs (in this study, counts were made of six queen combs and six worker combs from five V. shidai colonies). The actual n...

Watch this Scientific Journal Video about Evaluation of the Productivity of Social Wasp Colonies Vespinae and an Introduction to the Traditional Japanese Vespula Wasp Hunting Technique at JoVE.com

Evaluation of the Productivity of Social Wasp Colonies Vespinae and an Introduction to the Traditional Japanese Vespula Wasp Hunting Technique

This methodological paper evaluates the productivity of a social wasp colony by examining the number of meconia per 100 cells of comb, to estimate the total number of adults the wasps produced. The associated video describes how to search for Vespula wasp nests, a method developed by amateur wasp chasers.

Evaluation of the Productivity of Social Wasp Colonies (Vespinae) and an Introduction to the Traditional Japanese Vespula Wasp Hunting Technique

For vespine wasps, colony productivity is typically estimated by counting the number of larval cells. This paper presents an improved method that enables ...

Since Vespula wasp colonies cannot be lured from busts, nests must be collected from the wild. Here I demonstrate traditional wasp hunting as has been developed in Japanese culture. When following wasps to their nests, this technique can help in resuming nest location from the point at which a wasp has escaped detection.To attract vespula, hang 10-gram pieces of unflagged bait on a tree branch at a height that can be easily reached by hand along a transect at 500 to 100 stations, with at least 5 meters between each station. While waiting for the wasps, use scissors to cut plastic polyethylene bags into three to five millimeter wide by 15 centimeter long strips. And cut cuttlefish, freshwater fish or chicken hearts into 10 milligram pieces.Next, tie one end of extremely fine polyester thread to one plastic strip and the other end to one piece of meat. Then cut any loose thread above the knots. before mounting the bait onto individual bamboo skewers, When wasps begin to approach the hanging bait, hold out the flagged bait on the bamboo skewers to allow easier access to the feeding wasps.While the wasps are biting the bait, use a water-based paint pen to paint a unique mark on the thorax of each wasp. Orient the flag with the thread under the wasp while it bites the flagged bait so that the thread passes under the wasp's abdomen from below its thorax. After identifying a wasp that returns to the bait repeatedly within four minutes of leaving gather the baits from the surrounding areas so that the returning wasp is more likely to return to the same spot and follow the marked wasp of interest.When a followed wasp lands somewhere on the way to its nest, gently lift the wasp with a long stick or fishing rod and watch the wasp until it resumes flight. If a wasp escaped detection while being followed, wait for the wasp to return to the bait station on the transect before resuming the chase and carry the bait stick and wasp to the point where the wasp had last escaped detection. Foraging wasps do not let go of their baits easily and do not sting if handled gently and therefore can be moved to the desired location by holding the flag without the wasp escaping.Upon the location of a nest, put on the appropriate protective clothing and collect as many workers into the nest as possible by vigorously stomping on the ground around the nest for 10 to 20 minutes so that the workers remain inside to protect the nest. When all of the wasps have returned to the nest shine a light directly into the nest entrance to determine the direction in which the nest entrance runs and use a finger to confirm the orientation of the nest hole, while gently excavating soil from around the nest. When the wasps have calmed, place the excavated nest into a wooden custom made carrying box and cover all but the top of the nest with branches and newspaper.To estimate the number of cells per comb, separate the combs one by one, and sweep away all of the adult wasps from the comb. Use tweezers to pull out all the larvae and pupae from the cells, and place a scale reference onto the comb. Take a picture of the comb from above and open the image in ImageJ.Measure the square areas of ten randomly chosen cells per comb in pixels and convert the measurements to the actual areas. Then divide the area of each comb by the average cell area per comb to estimate the number of worker and queen cells. To count the number of miconia per 100 cells for each comb carefully break the comb and examine the miconia.The miconia may have solidified into two or more layers in the cells. In this study, a comb with an average cell of less than 20 millimeter squared was defined as a worker comb and larger combs were defined as queen combs. After counting the number of cells for queen and worker combs, the actual number of cells per comb can be estimated via extrapolation.Using real data, the mean and standard deviation of the number of miconia per cell can be calculated with the number os samples set up 1000 for each sample size. This analysis showed that the standard error of the mean was less than 0.05 when the sample size was 100 cells for both the worker and queen combs. The actual number of miconia in the cells of worker combs which represents the number of workers produced ranged from 1.96 times more than the number of estimated larval cells to 0.89 times less than the estimated number of cells.In the queen combs, the actual number of cells was often less then the estimated number of cells. The number of miconia in the queen combs which can represent a component of fitness was 0.69 to 1.97 times the estimated number of cells. If a wasp escapes detection while being followed, the bait stick can be carried to the point where the wasp was last observed.The flags can also be bound directly onto the bodies of wasps with a large body size to aid in the location of the nest. This method not only makes it easier to find wasp nests, but it can also help researchers in the analysis of wasp homing behaviors. Always make sure no one else is around the nest before approaching, to prevent other visitors from being attacked.Remove the workers after collecting the nest.

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