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The Rarity and Overexploitation Paradox: Stag Beetle Collections in Japan

Article  in  Biodiversity and Conservation · June 2012

DOI: 10.1007/s10531-012-0253-y

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ORI GIN AL PA PER

The rarity and overexploitation paradox: stag beetlecollections in Japan

Pierline Tournant • Liana Joseph • Koichi Goka • Franck Courchamp

Received: 18 October 2011 / Accepted: 27 January 2012� Springer Science+Business Media B.V. 2012

Abstract For some wildlife commodities, rare species are especially sought after. The

tendency for rare commodities to be of higher value can fuel their exploitation and as

numbers dwindle, the demand can increase. Consequently, this can precipitate these rare

species into an overexploitation vortex where they become increasingly rare, valued and

exploited until eventual extinction. We focus here on the hobby of collecting stag beetles,

to ascertain if the market value of these items is driven by rarity and if, consequently, these

species are vulnerable to this overexploitation vortex. Stag beetle collections fuel a large

and lucrative market in Japan, involving more than 700 species from all over the world,

with over 15 million specimens imported a year. Some particularly valued species fetch

more than US$5,000 a piece. We assessed the importance of species rarity as an acquisition

criterion in this market using two methods: an Internet online questionnaire responded to

by 509 participants and through examining the quantities imported in Japan and prices paid

by collectors. We discovered that species rarity is one of the main choice criteria for

acquisition by collectors: rare stag beetles are valued more than the common species and,

consequently, stag beetles are vulnerable to the anthropogenic Allee effect in this market.

Because of the sheer size of the market and the pervasive nature of this rarity paradox, the

P. Tournant � L. Joseph � F. Courchamp (&)Ecologie, Systematique & Evolution, UMR CNRS 8079, Universite Paris Sud,91405 Orsay Cedex, Francee-mail: franck.courchamp@u-psud.fr

P. TournantTHEMA (UMR CNRS 6049) and Chrono-environnement (UMR CNRS 6249),Universite Franche-Comte, 25030 Besancon Cedex, France

L. JosephWildlife Conservation Society, 2300 Southern Boulevard, The Bronx, NY 10460, USA

K. GokaNational Institute for Environmental Studies, 16-2 Onogawa, Tsukuba, Ibaraki 305-0053, Japan

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attraction to rarity equates to a potential extinction threat for many rare stag beetles

species.

Keywords Anthropogenic Allee effect � Market price � Questionnaire �Extinction risk � Wildlife trade � Collector

Introduction

The overexploitation of species is among the chief factors responsible for the current

decline of biodiversity, affecting more than one-third of the animals that are threatened

with extinction (Rhodin 1999 cited in Gibbons et al. (2000); Rosser and Mainka 2002). In

many cases, the threat of overexploitation can be amplified and accelerated when species

become rare through a process known as the Anthropogenic Allee Effect (Angulo et al.

2009; Brook and Sodhi 2006; Courchamp et al. 2006). This paradox is analogous with the

Allee effect where the state of being in a small population is correlated with high extinction

risk. The anthropogenic Allee effect occurs because of the tendency for consumers to value

rare commodities over common varieties; this can fuel their disproportionate exploitation

and as numbers dwindle, the demand further increases. Consequently, this can precipitate

these rare species into an overexploitation vortex where they become increasingly rare,

valued and exploited until eventual extinction (Courchamp et al. 2006; Hall et al. 2008).

The anthropogenic Allee effect is proposed to exist in a number of economic markets

including collections, trophy hunting, traditional medicine, exotic pets, luxury goods and

even ecotourism (Courchamp et al. 2006). The clearest example is probably hobby col-

lection of wild plants and animals. Hobby collectors target a wide variety of wildlife;

familiar collections include orchids, butterflies, seashells and bird eggs. Like all types of

collections, the objective is often to acquire a large diversity or a full set of specimens and

the rarest types are the most valued. For example, the price of butterflies harvested for

hobby collections has been shown to be highly correlated to species rarity (Slone et al.

1997). The collecting value of birds eggs is also associated with their rarity: the Taylor’sStandard American Egg Catalogue listed all North American bird eggs together with their

price, giving the Condor egg at $350; the Bald Eagle at $8; the Horned Lark at $1; and the

Blue Jay at 10 cents (Taylor 1904). The hobby collections market is known to trade in both

legal and illegal commodities and is implicated in overexploitation of wildlife throughout

the world (Brook and Sodhi 2006; Kameoka and Kiyono 2004). Collections for other

purposes, for example with scientific objectives, can also trigger an extinction, as exem-

plified by the case of three rare freshwater fish endemic to Mexico, and all collected to

extinction within a few years of their discovery (Rodriguez-Estrella et al. 2006).

One example of this hobby is the hugely popular collection of stag beetles, for display

or as pets, in Japan (Kameoka and Kiyono 2004). Stag beetles are a group consisting of

four subfamilies of the Lucanidae family characterized primarily by large sized mandibles

of males resembling the antlers of stags. Among the 1,200 species described so far, only 35

are found in Japan. The stag beetle mania originated with a an arcade game called Mushi

(insect) King, in which players collect cards of virtual stag beetles as fighters in tourna-

ments. From card and virtual beetles, the passion has led to collection of real insects. Their

immense morphological variety, and the highly visual combats performed by males makes

them both interesting collecting items and valued pets.

Over one million specimens of more than 700 species are imported annually into Japan

from all over the world (see origin map in Goka et al. 2004), some of them illegally

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(New 2005). Trade in stag beetles is hugely lucrative—some individuals selling for more

than US$5000 (Goka et al. 2004). Prices approaching US$100,000 for a single individual

have been recorded (New 2005). This trade is responsible for a destructive illegal trade in

many countries, especially in Asia (Kameoka and Kiyono 2004; New 2005). It is believed

that collecting pressures may be leading to increasing scarcity, and contributing to the

listing of several species or subspecies as ‘Rare’ in the local Red Data Books (New 2005).

Although some species are protected, and their importation illegal, many such species are

found are customs controls or during shop surveys (Goka et al. 2004; New 2005, 2007). In

addition, probably due to the enormous economic market this trade represents (over a

hundred million US$ each year), an increasing number of species are becoming legally

imported to Japan (over 500 species in 2004, Goka et al. 2004).

Despite a blatant lack of protection of this group in such an international trade, the

importance of stag beetles in conservation is real, as illustrated both by their status as

charismatic species, and by their major ecological functional roles in ecosystem. Because

of the enormous scale of the stag beetle trade (both live and dead) and the potential for rare

species to be disproportionately collected, stag beetles may be vulnerable to the anthro-

pogenic Allee effect and be more threatened with extinction than is currently anticipated.

In this context, we aimed to test the hypothesis of such a process occurring in stag beetle

hobby collections in Japan. We used two approaches to assess whether rarity was a

characteristic of interest to collectors, and thereby a threat to the concerned species. First,

we used a market survey to assess if stag beetle collectors have a preference for rarity, by

examining the volumes and the prices paid for rare and common species that are imported

into Japan. Second, we used a web-questionnaire to examine the Japanese stag beetle

collectors’ acquisition criteria and, specifically, their potential preference for rarity.

Materials and methods

Market survey

We collected information about the identity, morphological characteristics, total quantity

imported, selling price, and rarity of species imported to Japan in 2008. The selling price

and the specialists’ estimate of rarity were recorded for 107 species belonging to 17

different genera, the most specious being Dorcus (44 species) and Prosopocoilus (23

species). The selling prices were recorded for a pair of stag beetles (male and female) in

Euros. The morphological measurements were collected for 102 of these species and the

volumes imported were available for 79 of them. Four morphological characteristics were

documented: one qualitative parameter representing the color (black/colored) and three

quantitative parameters comprising the body length (from the end of the abdomen to the

top of the head), thorax width (at the first leg insertion level) and mandible length (from the

end of the mandibles to the top of the head). These characteristics were measured from

the digitalization of reference photographic plates (Mizunuma and Nagai 1994).

The values for rarity were estimated by surveying expert opinions. Rarity is an elusive

concept having a variety of meanings in difference contexts; for example, a species can be

geographically restricted, have a low density or be difficult to find on the market (Gaston

1994). In this study, we used a definition of rarity that represented the consumer’s

understanding of rarity, which is influenced by the availability on the market, the number

of individuals in the wild and the species’ threat status. We asked nine scientists—spe-

cialists of stag beetle collection and trade—to allocate to each species a relative score

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between one, for very rare, to five, for very common. Rarity scores for each species were

calculated as the mean of the nine scores.

Statistical analysis

A correlation matrix was calculated to look for possible correlations between the char-

acteristics that defined species and to identify redundant variables in the database. Cor-

relation tests (Pearson method) were performed for both databases to analyze relationships

between body length, mandibles length, thorax width, color, rarity, selling price and

volumes imported. The many correlations between all parameters allowed us to keep fewer

parameters in the subsequent analyses; we used the following uncorrelated characteristics:

‘‘mandibles length’’, ‘‘thorax width’’, ‘‘rarity’’, ‘‘import’’ and ‘‘color’’.

Following the selection of parameters provided by the correlation matrix, the selling

price variations were analyzed with generalized linear models (GLM) using a set of

explanatory variables: rarity, mandibles length and the color. GLMs were constructed

assuming normal distribution and a stepwise selection procedure allowed to select the best

fitting model (based on the Akaike Information Criterion, AIC). Analyses of variance

(ANOVA) of the GLMs were made using a type 3 ANOVA and associated P values were

calculated. In the same way, the variations of the volumes imported were analyzed with

GLM using the same parameters as explanatory variables. The statistical analysis were

performed with R� version 2.9.0 (2009-04-17).

Questionnaire

Using a specifically designed web site, we developed a questionnaire with the aim to

determine which parameters (and their relative importance) influence decisions to acquire

stag beetles. The stag beetle parameters included in this questionnaire were morphological,

behavioral and reproductive criteria, as well as rarity as perceived by the respondent. The

questionnaire was composed of six web pages and contained four different experiments.

The questionnaire (http://www.webbeetlechoice.fr.st/) was presented both in English and

in Japanese. The link to the questionnaire was published on the professional website of the

Japanese laboratory of invasive species and was sent to Japanese stag beetle collectors and

advertised on websites dedicated to stag beetle collecting.

Amongst scientists who use social surveys to examine consumer preferences, concern may

exist about the reliability of Internet based studies. However, a study specifically aimed at

comparing the potential differences in the use of stated preference surveys when conducted

either over the Internet or through face-to-face interviews showed Internet surveys did not seem

to lead to significantly different or biased results (Lindhjem and Navrud 2011). In addition,

Internet surveys can provide a much larger sampling size, especially when the focus group is

relative specialized. For this reason, we chose to conduct our study over the Internet, targeting

Japanese stag beetle collectors through advertising on specialised websites. The disadvantages

of not being able to select the participants are thought to be greatly outweighed by the advantage

of the number of respondents, thereby reducing the potential limitations of this method

(Lindhjem and Navrud 2011).

The Internet survey consisted of six pages: The first page provided a description of the

questionnaire and a brief survey to record the participant’s demographic details. In an

attempt to limit the potential bias in the responses, the description did not reveal the true

purpose of the experiments—which was to examine preference for rarity—but, instead, it

indicated that the purpose was to examine choice-considerations in stag beetle collection in

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general. The demographic details included the participant’s sex (man or woman), age

(\13 years old, 13–25, 26–50, [ 50), collector status (collectors of stag beetles, collectors

of other animals/plants, collectors of objects such as coins or stamps, non collectors), and

their pet owners status (stag beetles, domestic animals such as dogs or cats; owners of

exotic animals; non-owners). The last webpage invited participants to send the link of the

website to friends and colleagues including a prepared message and requested that they did

not to disclose information about the experiments. The four intervening pages contained

our experiments and are described below.

Experiment 1

The first experiment was designed to measure value attributed to stag beetles because of

perceived rarity (i.e., fewer individuals in an image). This webpage contained an image with 50

stag beetles; the individuals were, in fact, from one single picture colored in three different

shades (red, black or green) and positioned randomly in the image (see Fig. 1a). Of the three

colors presented, one was relatively abundant (33 individuals), one was less abundant (15

individuals) and one was rare (two individuals only). The participants were asked to choose

their preferred stag beetle in the image by clicking on it. For 472 participants, no information

was provided to help with the choice. For 37 additional participants, information was provided

describing the relative abundance of the stag beetles. For example, in one scenario, the green

stag beetle was represented by 2 images and the description stated that one type of stag beetle

was rare. Six pictures with different combinations of relative abundance and color were pre-

sented randomly. A program recorded the color and rarity of the beetle that had been chosen (by

the participant clicking on their preferred choice). We worked under the hypothesis that, all

other things being equal, if participants chose the rarer color more often than expected by

chance (i.e., the number expected proportional to the number of individuals of the color in the

image), it showed a preference for rarity.

Experiment 2

The second experiment was designed to simultaneously test preference for rarity, color and

mandible size and to ascertain their relative importance. The participants were asked to

choose, in order of preference, three individuals from eight stag beetles in a picture (see

Fig. 1b). Each individual was a modification of a single photo and a unique combination of

a set of three variables with two categories each: color (red or black), mandibles size (small

or large) and rarity status (rare/abundant). Rarity was explicit in this experiment, indicated

by a label below the photo (with the other variables). The combination of these three

criteria was placed randomly each time to avoid a bias due to the position of the word or

the place of the picture in the webpage.

Experiment 3

The third experiment was designed to determine if knowledge of the rarity of a species will

influence the consumer’s choice in stag beetles when compared with other factors,

including mandible size, color and size. We asked participants to choose one of four

species of stag beetles, this time represented by photographs of real collected species.

Similarly to the other experiments, the photos appeared in a random order for each par-

ticipant. Each stag beetle was characterized by one outstanding attribute among the four

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Fig. 1 a Picture presented for the first experiment of the questionnaire is composed of stag beetles with threedifferent colors: black is the rarer here, green is moderately abundant and red is very abundant. b Picture presentedfor the second experiment where people were asked to choose among beetles with three different attributes (color,mandible size, rarity). c Picture showing the four species of stag beetles presented to the participants for the thirdexperiment. The first one from the left side is rarer than the three others, the second has larger mandibles, the third ismore colorful and the fourth is larger

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following: large size, colorful, large mandibles and rarity (see Fig. 1c). Only Dorcusgracilicornis is rare, the other ones are common but Dorcus curvidens has a large size,

Phalacrognatus muelleri is very colorful and Cyclommatus metallifer has large mandibles.

Except for rarity, each characteristic is visible on the photos; the rarity value should be

only known by people with a good knowledge of stag beetles. We compared four groups of

people: non-collectors and collectors of objects or of animals and plants—these groups

were expected to be unable to identify that one of the species is rare—and collectors of stag

beetles only—some which were expected to be able to identify one of the species as rare

and the others as common. We hypothesized that the participants will select the beetle

according to a criterion important for them; if rarity is highly valued and they have a high

understanding of the market, they will select the rare species in preference. Through a

comparison of the two groups, we expected to be able to demonstrate that participants are

selecting stag beetles for their rarity value only when they are aware of this rarity.

Experiment 4

With the objective of determining if rarity is chosen among a set of potential choice

criteria, the fourth experiment invites participants to choose in order of preference three out

of twenty characteristics: rare, popular, familiar, exotic, easy to breed, cheap, colorful,

large size, abundant, difficult to breed, large mandibles, small size, unpopular, unusual,

non-aggressive, aggressive, expensive, small mandibles, black, and Japanese native. To

avoid the bias that may occur because of the position of the words, the words appeared in a

unique and random order for every participant. This experiment highlights the role of new

characteristics (e.g., aggressiveness, popularity) that had not been included in the other

experiments to assess whether rarity would still be favored among a larger number of

potential attributes.

Statistical analysis

The analysis of the distribution of the answers to the four experiments was done by v2 tests

(the H0 hypothesis being the homogeneity of the data distribution). The ordination of the

responses was examined using Kendall tests (the H0 hypothesis being the independence of

the order of data). The statistical analysis were performed with R� version 2.9.0 (2009-04-

17).

Results

Market survey

Price explanation

The best explanatory model of the variations of the selling price is provided by the size of

the mandibles and rarity of the species, without interaction. Rarity is significantly corre-

lated to the selling price of the individuals (LR v2 = 12.13; P = 0.0005). A negative

correlation was demonstrated between the estimation of rarity and the selling price of the

individuals (r = -0.25; P = 0.0013, see Fig. 2a); that is, the rarer the species, the more

expensive it is.

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Volume of importation explanation

The best explanatory model of the variations of the volumes imported is provided by the

size of the mandibles and rarity of the species, without interaction. Rarity is significantly

correlated to the volumes imported (Likelihood Ratio v2 = 9.99; P = 0.0016). A positive

correlation was found between the estimation of rarity and the volumes imported

(r = 0.37; P = 0.0003, see Fig. 2b); that is, fewer individuals of rare species are imported

than common ones.

Questionnaire

The questionnaire was put on line the 15th of May. We collected answers until the 28th of

November 2009. In total, 509 persons participated to the questionnaire (i.e., participated to

the four experiments), among which 153 were not collectors, 51 were collectors of objects,

70 collected plants or animals other than stag beetles and 235 were stage beetles collectors.

Fig. 2 Anthropogenic Allee effect demonstrated by a a negative correlation between estimation of rarityand the selling price and b a positive correlation between estimation of rarity and the volumes imported. Inboth cases, 1 is for the rarer species

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The distribution of the number of participants relative to the different categories is pre-

sented on Fig. 3. In all results below, n is therefore 509.

Experiment 1

The first experiment demonstrated that there is a significant difference in the frequency that

participants selected the rare, medium and abundant type of stag beetle. When no infor-

mation was provided to the participant about the abundance of each type, significantly

fewer people choose the most abundant type compared with rarest (v2 = 9.99;

P = 0.0406; see Fig. 4a) and marginally fewer people also selected the medium compared

with the rarest (v2 = 9.02; P = 0.0605; see Fig. 4a). Furthermore, when we provided

information about the relative abundance of the types, participants significantly choose the

rarest compared to the medium and abundant types (v2 = 22.52; P = 0.0001; see Fig. 4b).

Experiment 2

When participants were asked to rank a set of criteria in order of preference, we found that

the order was significantly different from random (v2 = 165.12; P \ 2.2e-16): the

majority selected the criteria ‘‘large mandibles’’, ‘‘rarity’’, and ‘‘colored’’ (see Fig. 5a). We

found that for each rank position, a criterion was significantly favored more than the

others; i.e. there was preference for a particular criterion at the first (v2 = 487.60;

P \ 2.2e-16), second (v2 = 301.60; P \ 2.2e-16) and third (v2 = 58.39; P = 2.607e-11)

positions. In each of rank position, ‘‘large mandibles’’ was favored the most, followed by

‘‘rarity’’ and then ‘‘colored’’. This is supported by the Kendall ordination tests, which

showed that the criteria order of the first choice is the same as the second choice (s = 1;

P = 0.0085) and as the third choice (s = 0.867; P = 0.0242). Similarly, the criteria order

of the second choice is the same as the third choice (s = 0.867; P = 0.0242).

When we repeated the analysis for different categories of collectors (non collectors,

collectors of stag beetles, and other collectors), the results revealed that the choice order of

Fig. 3 Ventilation of the number of participants of the survey in each category (A1: age class \13 yearsold, A2: age class 13–25 years old, A3: age class 26–50 years old, A4: age class [50 years old; C1: nocollector, C2: plants or animals collector, C3: objects collector, C4: stag beetles collector; O1: no owner,O2: domestic animal owner, O3: exotic animal owner, O4: stag beetle owner)

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collectors was even more specific than the choice order of non-collectors: first

(v2 = 24.81; P = 0.053), second (v2 = 26.80; P = 0.030) or third (v2 = 21.83;

P = 0.1121) rank position.

Experiment 3

When we compared the frequency of selection of four different stag beetles by all par-

ticipants (i.e., not sorting the participants into the four categories), we found that most

participants preferred the large bodied species (D. curvidens) followed by the species with

the large mandibles (C. metallifer), the colorful species (P. muelleri) and the rare species

(D. gracilicornis; see Fig. 6a) and that the difference in the frequency in which each of the

species was selected was highly significant (v2 = 99.56; P \ 2.2e-16).

When we categorized the participants into groups (i.e. non-collectors, collectors of

objects, collectors of animals and plants, and collectors of stag beetles), we found that each

group had a different preferred species (v2 = 33.97; P = 9.053e-05). The rare species was

Fig. 4 Number of answers (%, ±SD) according to the status of the species on the picture of Experiment 1(rare, medium or abundant), regardless of colors: a rarity was not specified to the participants andb participants are informed that one species is rare

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selected more frequently first by the collectors of plants and animals (v2 = 19.28;

P = 0.0002), and then by collectors of stag beetles (see Fig. 6b). The non-collectors

preferred the large bodied species (D. curvidens) and the collectors of objects preferred the

colorful species (P. muelleri).Notably, the participants that were not expected to have knowledge of the rarity value of

the stag beetles (i.e., non-collectors and collectors of objects) selected the rare species

much less often than the participants that were expected to have some knowledge

Fig. 5 Occurrence (%, ±SD) of the best criteria defining stag beetles, according to the order of choice ofparticipants a when rarity was proposed among only six criteria in Experiment 2 and b when rarity wasproposed among 20 criteria in Experiment 4

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(i.e., collectors of animals and plants, and collectors of stag beetles; Fig. 6b). Knowing that

a species is rare thus influences preference for that species.

Experiment 4

When we compared the frequency in which criteria were selected from list of twenty, we

found that some criteria were selected more than others (v2 = 162.62; P \ 2.2e-16): the

criteria selected the most frequently were (in order) ‘rare’, ‘large size’, ‘large mandibles’,

‘colored’ and ‘easy to breed’ (see Fig. 5b). We found that for each rank position, a

criterion was significantly favored more than the others; i.e., there was preference for a

Fig. 6 a Choice of participants for the four stag beetles species presented in Experiment 3, showing that inabsence of explicit information, the rare species, Dorcus gracilicornis, is the less often chosen; b choice ofthat rare species for each collectors class (%, ±SD), showing that animal (including stag beetle) collectorschose this species more often than other participants

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particular criterion at the first (v2 = 797.99; P \ 2.2e-16), second (v2 = 429.66;

P \ 2.2e-16) and third (v2 = 230.76; P = 2.607e-11) positions. Figure 5b reveals that in

each of rank position, ‘‘rare’’ was favored the most, followed by ‘‘large size’’ and then

‘‘large mandibles’’. This is supported by the Kendall ordination tests, which showed that

the criteria order of the first choice is the same as the second choice (s = 0.667;

P = 7.4912e-05) and as the third choice (s = 0.613; P = 0.0002). In the same way, the

criteria order of the second choice is the same as the third choice (s = 0.809;

P = 1.1917e-06).

When we repeated the analysis for different categories of collectors (non collectors,

collectors of stag beetles, and other collectors), the results revealed that everyone

responded in the same way regardless of their category. Specifically, we found that there

was no significant difference among the rank orders selected by each category of partic-

ipant for the first (v2 = 84.32; P = 0.2403), second (v2 = 96.48; P = 0.0565) or third

(v2 = 106.90; P = 0.0112) rank position.

Discussion

We discovered that rarity is highly valued in stag beetles collections. Preference for stag

beetles is influenced by rarity conveyed in a variety of different ways including through (1)

visual cues of rarity (Experiment 1), (2) being informed that a type is rare (Experiments 1

and 2), and (3) knowing that a species is rare (Experiment 3). We also found that people

will specify rarity as a key criterion of the choice in stag beetles when asked to rank a short

list of criteria (Experiment 2) and select from a longer list (Experiment 4). Additionally,

we were able to demonstrate that rarity in the wild is correlated with rarity on the market

and consumers pay more for rare species (market survey). These finding provide com-

pelling evidence that there is a strong and pervasive desire for rarity in stag beetles and

suggests that this phenomenon may pose a threat to endangered stag beetles in the wild.

Furthermore, our results show that preference for rarity is found in collectors and non-

collectors alike. That rarity value is not restricted to consumers of specific economic

markets but is more general, as was already found in the context of luxury food con-

sumption (Gault et al. 2008) and trophy hunting (Palazy et al. 2011a, b). This may be

considered a disturbing result as it implies that the attraction to rarity may potentially exist

among society as a whole and is not restricted to a certain set of consumers.

The exaggerated value that is placed on rare stag beetles is likely to pervade the entire

hobby collection market, which is a form of materialistic luxury consumption driven by the

need to acquire either (or both) ‘distinction’ through having an extensive set and dem-

onstrating expertise, and ‘closure’ through completing a set, or a series, or filling in gaps in

a collection (Danet and Katriel 1990; Belk 1995 both cited in Long and Schiffman 1997).

Stag beetle and other wildlife products—butterflies and birds eggs for instance—make

ideal collector item as they belong to a large superordinate category (i.e. an Order)

comprised of neat subcategories (i.e. Genera and Families) which allows for manageable

sets to complete while offering seemingly endless opportunities; all of which are key

characteristics of collectables (Danet and Katriel 1990 in Long and Schiffman 1997).

Collectors of all types of objects are attracted to rarity as the scarcer an item, the more

likely the opportunity for distinction from possessing it and its ability to complete a set

(Danet and Katriel 1990; Belk 1995 in Long and Schiffman 1997). Within the wildlife

collection market, the desire for rarity has been documented for butterfly collections

(Courchamp et al. 2006) and trophy hunting (Palazy et al. 2011a, b).

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Unlike other objects that collectors desire (i.e., stamps, coins, art), collections of

wildlife commodities can have dire negative consequences by depleting the species in the

wild. Through the process of the Anthropogenic Allee Effect, rare species that are cher-

ished by collectors can paradoxically be driven to extinction by them. Hall et al. (2008) and

Gault et al. (2008) demonstrated with ecological-economic models that the desire for rare

wildlife commodities could theoretically result in the extinction of the species through this

process. These results have been corroborated by examples in the wild where hobby

collectors have been implicated as a driver of the extinction or near extinctions: for

example, the great auk (Fuller 1999), the lady’s slipper orchid (Cypripedium calceolus;

Ramsay and Stewart 1998) and large copper butterfly (Lycaena dispar; Slone et al. 1997).

The interest in rare species as a leisure good is especially worrying as the general level

of world wealth increases. If these general trends translate to more interest in collecting

wildlife such as stag beetles, it will result in increased pressure on rare species that are

already threatened with over exploitation and the anthropogenic Allee effect.

These words of alarm can be tempered by the positive implications of wildlife hobby

collections. Benefits may include an increased awareness and appreciation of nature and

the increase in information gathering and sharing by enthusiasts (Lee and Trace 2009).

Recently, Sir David Attenborough—a very high profile nature documentary producer and

conservationist—publicly announced his opinion that children should be allowed to collect

common species to encourage an appreciation of nature and conservation (Gray 2010;

Wilkes 2010). It is indeed indisputable that increased public interest in insects is likely to

enhance the public’s positive perceptions of nature in general and invertebrates in par-

ticular (New 2005). However, our results and those of others (e.g., Hall et al. 2008; Gault

et al. 2008; Palazy et al. 2011a) suggest that encouraging leisure exploitation of wildlife

can promote a demand for rare species, which could be devastating for them if not bal-

anced by appropriate information and regulation. The threat over-collection can represent

in Japan for rare species of stag beetles has already been emphasized (Kameoka and

Kiyono 2004; New 2005). As stated by New (New 2007), ‘‘beetlephilia can be a two-edgedsword in conservation’’. The increased interest of collectors is likely to drive the rare

species to extinction in the wild before the improved knowledge developed within the

hobbyist’s community can improve their chances of survival (Stuart et al. 2006) but see

Kratochvil (2006). Regrettably, our understanding of consumer behavior in wildlife col-

lecting and in natural resource extraction in general—described aptly as ‘‘Tragedy of the

Commons’’ (Hardin 1968)—suggests that rare species are often acutely threatened by

collectors and policies that encourage hobby collections may risk hastening their extinc-

tion. In the case of stag beetles hobby collection in Japan, it is noteworthy however that

ongoing effort to breed some species may result in significant lower harvest as ecologically

rare species become common on the market and, thus, less valued.

Given the potential threat to wildlife presented by hobby collectors on some taxonomic

groups, it might seem the straightforward policy recommendation should be to ban all trade

on demonstrated cases of overexploitation. However, evidence suggests that trade bans can

also stimulate illegal trade (Rivalan et al. 2007) and strictly regulated trade can potentially

maintain viable and healthy populations in the wild (Hutton and Leader-Williams 2003).

Similarly, withholding information about rarity can hinder conservation programs and

must therefore be used with circumspection (Courchamp et al. 2006; Rivalan et al. 2007).

We recommend that to protect endangered stag beetles and other wildlife collector items,

management should be supported at all levels including: improving enforcement of already

existing bans in trade in the wild endangered species (Messer 2010), improving monitoring

(Phelps et al. 2010), setting scientifically and informed quotas, encouraging captive

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breeding (Hutton and Leader-Williams 2003; Kratochvil 2006), as well as educating and

informing the consumers (Jepson et al. 2008; Robbins 2003) who, paradoxically, may

inadvertently harm the very species they appreciate the most.

Acknowledgments We would like to thank Maxime Guinard for the website conception and Dr Yuya Watarifor translations of the site into Japanese. We also thank Kunio Araya, Luca Bartolozzi, Hiroshi Fujita, Paschoal C.Grossi, Tadatsugu Hosoya, Jen-Pan Huang, Hiroshi Kojima, Jean-Michel Maes and Eva Sprecher who all kindlyaccepted to rankall the stag beetles species in the database according to its rarity. We also wholeheartedly thank allparticipants for completing the survey and/or forwarding it to other participants. This work was supported by anANR Grant and a David H. Smith Conservation Research Fellowship.

References

Angulo E, Deves AL, Saint Jalmes M, Courchamp F (2009) Fatal attraction: rare species in the spotlight.Proc R Soc Lond Biol Sci 276:1331–1337

Belk RW (1995) Collecting as luxury consumption—effects on individuals and households. J Econ Psychol16:477–490

Brook BW, Sodhi NS (2006) Conservation biology—rarity bites. Nature 444:555–557Courchamp F, Angulo E, Rivalan P, Hall RJ, Signoret L, Bull L, Meinard Y (2006) Rarity value and species

extinction: the anthropogenic Allee effect. PLoS Biol 4:2405–2410Danet B, Katriel T (1990) Glorious obsessions, passionate lovers, and hidden treasures: collecting, meta-

phor, and the romantic ethic. In: International conference on the socio-semiotics of objects: the role ofartifacts in social symbolic processes. Toronto

Fuller E (1999) The great auk. Harry N. Abrams Inc. Publishers, New YorkGaston KJ (1994) Rarity. Chapman & Hall, LondonGault A, Meinard Y, Courchamp F (2008) Consumers’ taste for rarity drives sturgeons to extinction.

Conserv Lett 1(5):199–207Gibbons JW, Scott DE, Ryan TJ, Buhlmann KA, Tuberville TD, Metts BS, Greene JL, Mills T, Leiden Y,

Poppy S, Winne CT (2000) The global decline of reptiles, deja vu amphibians. Bioscience 50:653–666Goka K, Kojima H, Okabe K (2004) Biological invasion caused by commercialization of stag beetles in

Japan. Glob Environ Res 8(1): 67–74Gray R (2010) Let children collect flowers and fossils says Sir David Attenborough (in The Telegraph-28/

04/10)Hall R, Milner-Gulland E, Courchamp F (2008) Endangering the endangered: the effects of perceived rarity

on species exploitation. Conserv Lett 1(2):75–81Hardin G (1968) Tragedy of commons. Science 162:1143–1248Hutton JM, Leader-Williams N (2003) Sustainable use and incentive-driven conservation: realigning human

and conservation interests. Oryx 37:215–226Jepson P, Prana M, Aksenta S, Amama F (2008) Developing a certification system for captive-bred birds in

Indonesia. TRAFFIC Bull 22:1–9Kameoka S, Kiyono H (2004) A survey of the rhinoceros beetle and stag beetle market in Japan. TRAFFIC

East Asia-Japan 36–74Kratochvil L (2006) Captive breeding and a threatened gecko. Science 313(5789): 915–915Lee CP, Trace CB (2009) The Role of information in a community of hobbyist collectors. J Am Soc Inf Sci

Technol 60:621–637Lindhjem H, Navrud S (2011) Are internet surveys an alternative to face-to-face interviews in contingent

valuation? Ecol Econ 70:1628–1637Long MM, Schiffman LG (1997) Swatch fever: an allegory for understanding the paradox of collecting.

Psychol Mark 14:495–509Messer KD (2010) Protecting endangered species: when are shoot-on-sight policies the only viable option to

stop poaching? Ecol Econ 69:2334–2340Mizunuma T, Nagai S (1994) The lucanid beetles of the world. Mushisha, TokyoNew TR (2005) Inordinate fondness: a threat to beetles in South East Asia? J Insect Conserv 9(3):147–150New TR (2007) Beetles and conservation. J Insect Conserv 11:1–4Palazy L, Bonenfant C, Gaillard J-M, Courchamp F (2011a) Cat dilemma: too protected to escape trophy

hunting? PLoS One 6(7): e22424

Biodivers Conserv

123

Author's personal copy

Palazy L, Bonenfant C, Gaillard J-M, Courchamp F (2011b) Rarity, tropy hunting and ungulates. AnimConserv. doi:10.1111/j.1469-1795.2011.00476.x

Phelps J, Webb EL, Bickford D, Nijman V, Sodhi NS (2010) Boosting CITES. Science 330:1752–1753Ramsay MM, Stewart J (1998) Re-establishment of the lady’s slipper orchid (Cypripedium calceolus L.) in

Britain. Bot J Linn Soc 126:173–181Rhodin A (1999) Celebrate the turtle: perception and preservation. In: Powdermill 1999, 4th occasional

freshwater turtle conference. Laughlin, NVRivalan P, Delmas V, Angulo E, Bull LS, Hall RJ, Courchamp F, Rosser AM, Leader-Williams N (2007)

Can bans stimulate wildlife trade? Nature 447:529–530Robbins C (2003) Eco-labelling as a conservation tool for American Ginseng. TRAFFIC Bull 19:153–156Rodriguez-Estrella R, Carmen M, Moreno B (2006) Rare, fragile species, small populations, and the

dilemma of collections. Biodivers Conserv 15:1621–1625Rosser AM, Mainka SA (2002) Overexploitation and species extinctions. Conserv Biol 16:584–586Slone TH, Orsak LJ, Malver O (1997) A comparison of price, rarity and cost of butterfly specimens:

implications for the insect trade and for habitat conservation. Ecol Econ 21:77–85Stuart BL, Rhodin AGJ, Grismer LL, Hansel T (2006) Scientific description can imperil species. Science

312 1137–1137Taylor HR (1904) The standard American egg catalogue: a true basis for the exchange of nests and eggs.

H.R. Taylor, New York 32Wilkes D (2010) Let children collect bugs again to boost interest in science, says Sir David Attenborough (in

MailOnline)

Biodivers Conserv

123

Author's personal copy

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