Fireflies in Art: Emphasis on Japanese Woodblock Prints from the Edo, Meiji, and Taishō Periods

Simple Summary Fireflies are beetles (Coleoptera: Lampyridae) famous for their bioluminescence. This study examined artistic representations of fireflies and depictions of how people interacted with these insects in Japan from a historical perspective. Visual information from the artwork was summarized, highlighting themes and connections to firefly biology and cultural entomology. Multiple artists were represented, including several renowned masters, and the artwork highlights the complex interactions between fireflies and humans. Analyzing artwork can enhance awareness of the historical and cultural significance of insects and may help with conservation efforts. Abstract Examining how insects are represented in artwork can provide insight into people’s perceptions and attitudes towards arthropods, as well as document human–insect interactions and how they change through time. Fireflies are well-known bioluminescent beetles (Coleoptera: Lampyridae) of great cultural significance, especially in Japan. A selection of online museum collections, art databases, and dealer websites were used to find artwork featuring fireflies, with an emphasis on Japanese ukiyo-e wood block prints from the Edo, Meiji, and Taishō time periods (1600–1926). Quotes from early twentieth century texts were used to provide additional historical context. Over 90 different artists created artwork featuring fireflies, including several renowned masters. Artists depicted adult fireflies in a variety of ways (e.g., relatively accurately, more generalized, symbolic or abstract, yellowish dots) in the absence and presence of people. Most images were set outdoors during the evening near water, and primarily featured women and children, groups of women, and large parties catching fireflies or observing caged fireflies. ‘Beauties’, geisha, courtesans, kabuki actors, and insect vendors were also common subjects. Various types of collecting tools and a diversity of cages were featured, as well as insect vendors. The artwork highlights the complex connections between fireflies and humans. Insect-related art can contribute to education and conservation efforts, particularly for dynamic insects such as fireflies that are facing global population declines.


Introduction
Art is a means of expression and a valuable communication tool. Examining artistic representations of insects and their relatives can provide insight into people's perceptions and attitudes towards arthropods, as well as document human-insect interactions and how they change through time [1][2][3][4][5][6]. Insect-related artwork can also contribute to educational efforts [7,8], and stimulate discussion about modern societal concerns, e.g., impacts of anthropogenic activities on the environment [9] and conservation of culturally important insects, such as fireflies [10,11].
Fireflies are beetles (Coleoptera: Lampyridae) famous for their bioluminescence. There are over 2500 species of lampyrids [12]. They have a broad geographic distribution and all produce light at some point during their life cycle, although not all adult fireflies bioluminesce [13]. Researchers have used mating behavior signals to group North American fireflies into three categories: diurnal species that rely on pheromones (dark fireflies),

Focus on Fireflies
Artists depicted adult fireflies in the absence or presence of people, with the apparent lack of immatures perhaps related to their more cryptic nature. Representations of adults were diverse, whether free-ranging or contained within cages, and ranged from realisticlooking insects to yellow-colored or golden dots. Research indicates flashes of crepuscular firefly species are yellower than the greener flashes emitted by nocturnal fireflies [40]. 'Fireflies at Ochanomizu' by Kiyochika (T49) is a good example of where the fireflies' light

Focus on Fireflies
Artists depicted adult fireflies in the absence or presence of people, with the apparent lack of immatures perhaps related to their more cryptic nature. Representations of adults were diverse, whether free-ranging or contained within cages, and ranged from realisticlooking insects to yellow-colored or golden dots. Research indicates flashes of crepuscular firefly species are yellower than the greener flashes emitted by nocturnal fireflies [40]. 'Fireflies at Ochanomizu' by Kiyochika (T49) is a good example of where the fireflies' Insects 2022, 13, 775 4 of 29 light was greenish or chartreuse, rather than golden yellow. Some fireflies appeared to be generalized insects (e.g., T54) or resembled butterflies (e.g., T144-146, T175) or were more abstract, such as a blotch or the letter 'X' (T171). However, in other pieces the insects were clearly fireflies, with elongate bodies, well defined elytra, segmented abdomens, pink coloration on the pronotum, and the distal end of the abdomen greenish-yellow or depicted as glowing. Table A1 provides information on how fireflies were represented in the artwork (i.e., letter codes after the title). Of pieces clearly related to fireflies where the insects could be seen (with multiple sheets of a triptych counted as one image), 22% of artworks had relatively accurate depictions of fireflies, 32% had less accurate and more generalized fireflies, 26% had more symbolic or abstract fireflies, and 20% had fireflies represented by yellowish or greenish dots. Additionally, 47% had some indication of firefly bioluminescence (e.g., yellowish or greenish abdomen or dots at the end of the abdomen. Artwork by certain artists, such as Sōzan and Zeshin, had more accurate representations of fireflies. In general, the more accurate depictions of fireflies tended to be on artwork lacking people or on objects. With regard to the latter, fireflies on two carved netsuke (T104-105) had a rounder body and reddish pronotum reminiscent of a less common diurnal firefly from Japan, Cyphonocerus ruficollis Kiesenwetter [17,41,42]. Images of fireflies interacting with other animals were rare, perhaps because fireflies are chemically protected [18,19], although one print showed fireflies above what looks like an interested dragonfly naiad (T167), and another showed a firefly trapped in a spider's web (T170). Lafcadio Hearn (b.1850Hearn (b. -d.1904), an author who wrote extensively about Japan [43] reported, "[ . . . ] the firefly has a very bitter taste, and birds appear to find it unpalatable. (Frogs [ . . . ] do not mind the bad taste: they fill their cold bellies with fireflies till the light shines through them [ . . . ])" [34] (p. 138).
Two-dimensional artwork focused solely on the insects fell into two broad categories. Several pieces, including multiple works by Zeshin, had a few non-glowing fireflies flying or at rest, usually outdoors in the daytime surrounded by white space (T115-116, T161, T166, T168-169, T199-200, T203) or in still-lifes with flat uchiwa fans, decorative trays, or lanterns (T174, T198, T202). A few prints showed fireflies on plants with greenish-gray or dark gray backgrounds (T183, T197). A second group of paintings and prints depicted fireflies-often dozens of them-at night near water. These night scenes had muted gray or black color palettes, which highlighted the fireflies' reddish body parts and luminescence, and included woodblock prints by Kōgyō (T59), Koson (T65), and Sōzan (T148-149) and paintings by Shōnen (T126-127, Figure 2) and Bunrin (T4-8). Bunrin often depicted fireflies [44], and the gallery label for his 'River Landscape with Fireflies' (T4) reads, "[ . . . ] tiny golden flashes of fireflies along a riverbank evoke the charms of summer. Tufts of bamboo, willow trees, and cascading waters convey the cool nocturnal atmosphere. By incorporating naturalistic effects such as the brushwork and ink tones that capture the volume of rocks and water, Bunrin created a sensation of rushing waters and cooling nighttime breezes".
Some night scenes had human elements, such as buildings (T151, T155) and indistinct figures on boats (T49) or behind window shades (T50), although the fireflies were the primary element of interest. Kiyochika's print 'Koromogawa River at Tennōji-shita', (T50) was reproduced in a book based on the experiences of an English woman who lived in Japan for several years, who recounts of her journey to Ikao, "Suddenly, in a lull of the rain, I saw a great white star moving slowly down towards me out of the sky. Only when it floated close to my eyes did I discover that it was the very patriarch of all the fireflies [ . . . ]." [45] (p. 27). Several images had silhouetted figures, including work by Gekkō (T24), Hiroaki (T131), Shōtei (T130, T134), Shōun (T136), Toshihide (T156), and Toshikata (T157), and one had the background landscape and plants in silhouette while the people in the foreground were in vibrant color (T88).

Settings
Watching and hunting fireflies for entertainment has been a popular custom in Japan for centuries, and Hearn [34] (p. 149) indicated that, "anciently it was an aristocratic amusement; and great nobles used to give firefly-hunting parties,-hotaru-gari". Many places were famous for their fireflies, such as the Hotaru-Dani (Valley of the Fireflies) near Ishiyama, the lake ofŌmi, and Uji in Yamashiro Province [34,46,47]. An early travel guide [46] (p. 552) based on [34]  , and the scores of illuminated boats on which there are dancing and singing, geisha, music, and jollity, add to the charm. When the fireflies have assembled in force myriads dart from either bank and meet and cling above the water. At moments they so swarm together as to form what appears to the eye like a luminous cloud, or like a great ball of sparks. [ . . . ] After the Hotaru-Kassen is done, the river is covered with the still sparkling bodies of the drifting insects. Then the natives refer poetically to the stream as the 'Milky Way;, the 'River of Heaven', etc." Some pieces of art mentioned specific locations, many of which were renowned for their fireflies. Two of Shōnen's hanging scroll paintings depicted fireflies over the Uji River (T126-127), and Tokuriki had a piece featuring the Uji River in his '15 Views of Kyoto' series (T155). Kunisada I created a print of women catching fireflies by the Uji River (T88), a print listed as, 'Catching Fireflies at Sekiya', (T71), which is a village by the Sumida River, and one featuring actors titled, 'Catching Fireflies by the Sumida River' (T79). The Sumida River was also referenced in a print by Kiyonaga (T56). Other places referenced in the artwork included: Sahô River (T117), Koromogawa river at Tennōji-shita (T50), Ochanomizu (T23, T49), Ochiai (T89), Higashiyama and Yoneyama (T102), Sekiguchi (T132), Negishi Village in Toyko (T128), Mount Dōkan (T34), Chiyoda Castle (T12), Byodo-In Temple in Kyoto (T151), and Ichinose Bridge (T1, T172). The swarms of fireflies in the Valley of the Fireflies near Ishiyama and the lake ofŌmi were considered a natural marvel prior to 1703, but by 1903 people had noticed their populations declining [34] (pp. 143-144).
Image settings were primarily outdoors and featured water, most often streams or small rivers, and less frequently large rivers or lakes. Common plants featured in images included grasses, iris flowers (e.g., T14, T16, T29, T32, T44, T52, T54, T90), dwarf bamboo (T8), and willow trees (e.g., T9, T12, T112, T186). One 18th century garment (furisode kimono) at the National Museum of Japanese History with irises and fireflies was thought to be based on a 13th century poem by the shogun Minamoto no Sanetomo [44]. Hearn wrote, "Fireflies frequent the neighbourhood of water, and like to circle above it; but some kinds are repelled by impure or stagnant water, and are only to be found in the vicinity of clear streams or lakes. The Genji-firefly shuns swamps, ditches, or foul canals; while the Heiké-firefly seems to be satisfied with any water. All fireflies seek by preference grassy banks shaded by trees; but they dislike certain trees and are attracted by others. They avoid pine trees, for instance; and they will not light upon rose-bushes. But upon willow trees-especially weeping willows-they gather in great swarms. Occasionally, on a summer night, you may see a drooping willow so covered and illuminated with fireflies that all its branches appear 'to be budding fire'," [34] (pp. 151-152).
In modern-day Japan, watching adult fireflies is especially popular in early summer from May to July [30,48], and many prints referenced summer or a specific month in their titles, e.g., fifth month (T156), seventh month or July (T13, T96, T116), including a calendar print (T13) from July 1910 with an advertisement for 'Deer and Stag' pure silk from the Kawamata Silk Refining Company, Yokohama, Japan.
Several woodblock prints showed people collecting fireflies during a partial or full moon (e.g., T43, T46, T56, T68, T69, T102, T109, T132, T169, T171). Ambient light levels influence firefly behavior [49] although firefly abundance is similar during full and new moon phases [50,51]. However, artificial light pollution negatively impacts flashing activities and mating success of some firefly species and is considered a threat to firefly populations and conservation [52][53][54]. Hearn [34] (p. 152) indicated, "During a bright moonlight night fireflies keep as much as possible in shadow […]. Lamplight, or any strong artificial light, drives them away; but small bright lights attract them. They can be lured, for example, by the sparkling of a small piece of lighted charcoal, or by the glow of a little Japanese pipe, kindled in the dark. But the lamping of a single lively firefly, confined in a bottle, or cup, of clear glass, is the best of all lures." Some images, especially larger triptych prints, had both an outdoor and indoor component, and included open air balconies or patios (e.g., T19, T56, T68-69, T96), gardens (e.g., T41, T109), or other open-air spaces adjoining houses (e.g., T9, T144-146, T182). These images often show people engaged in multiple types of activities, such as collecting fireflies, watching others collect, observing caged insects, or enjoying refreshments. It was less common to encounter images that were set solely indoors (T185), and occasionally the setting was unclear (T2), especially when only people and cages were shown (T26, T28, T45). In one Kunisada I print (T72), the only reference to fireflies was a picture of them hanging on a wall.
In modern-day Japan, watching adult fireflies is especially popular in early summer from May to July [30,48], and many prints referenced summer or a specific month in their titles, e.g., fifth month (T156), seventh month or July (T13, T96, T116), including a calendar print (T13) from July 1910 with an advertisement for 'Deer and Stag' pure silk from the Kawamata Silk Refining Company, Yokohama, Japan.
Several woodblock prints showed people collecting fireflies during a partial or full moon (e.g., T43, T46, T56, T68, T69, T102, T109, T132, T169, T171). Ambient light levels influence firefly behavior [49] although firefly abundance is similar during full and new moon phases [50,51]. However, artificial light pollution negatively impacts flashing activities and mating success of some firefly species and is considered a threat to firefly populations and conservation [52][53][54]. Hearn [34] (p. 152) indicated, "During a bright moonlight night fireflies keep as much as possible in shadow [ . . . ]. Lamplight, or any strong artificial light, drives them away; but small bright lights attract them. They can be lured, for example, by the sparkling of a small piece of lighted charcoal, or by the glow of a little Japanese pipe, kindled in the dark. But the lamping of a single lively firefly, confined in a bottle, or cup, of clear glass, is the best of all lures."

People Represented in Artwork
Japanese children frequently spend a great deal of time learning about, and observing or playing with insects, or mushi, and often hunt fireflies in the summer [34,48,[55][56][57]. "Girls follow the chase with paper fans; boys, with long light poles to the ends of which wisps of fresh bamboo-grass are tied." [34] (p. 150). This is not restricted to Japan, as Liu [58] notes, "The fireflies are still one of the best evening entertainments the Chinese children have today. Mothers are generally requested by their children to save their empty egg-shells in which the youngsters house their catch and watch the flashing in the dark when they go to bed." Catching fireflies is also a common pastime for children in the United States [59][60][61][62][63]; Carter's image 'Fireflies' showed two boys standing in water looking at fireflies trapped in a glass jar [64]. Some Japanese artwork only featured children (T39, T48, T195-196, T201), including Shuntei's print with five girls, where the artist captured a sense of vigorous movement and excitement (T143). Most of the images with children showed one child collecting fireflies with one woman who was likely their parent (e.g., T16, T31, T43, T89, T138) or walking home after collecting (T132). There were also multiple women with one child (T42, T90) and multiple women with multiple children (e.g., T9, T11, T44, T109, T140-141), including 'Catching Fireflies (Hotaru gari)' by Utamaro (T186, Figure 5).
Images with larger groups of people showed social gatherings where the evening's entertainment focused on collecting and observing fireflies. Most images had women and men, although sometimes only members of one sex were present, and occasionally children were also in attendance (e.g., T19-20, T56, T70, T88, T112, T128, T144-146). Lanterns, blankets, food, beverages, pipes, musical instruments, and pets were often pictured, emphasizing the recreational aspect. Artists rarely pictured individuals wading in the water while collecting (T18, T186). However, small groups in boats (T10, T60, T133) and larger boating parties were common (T107, T154, T181). 'Firefly Viewing Party' shows a boating party near a bridge where other people were also catching fireflies, and the museum commentary reads, "Hotarugari (firefly viewing) is a popular summer pastime in Japan. These small insects produce flashes of light-which can be seen at night-during the hot months when they breed. Since they live near water, firefly viewing had an added attraction: the cool night breezes off the water brought relief from the heat. The people depicted in this print have hired a boat to take them out on the water. Those in the bow reach towards the fireflies with their fans, attempting to sweep them closer. Those in the center of the boat are drinking and chatting convivially. An attendant blows on a portable stove, attempting to keep a small fire alive so that he can prepare a snack for the group. Twenty-nine haiku poems on the theme of summer are printed in the upper portion of the print." (Minneapolis Museum of Art, T171). This artwork is considered a surimono, or genre of non-commercial woodblock prints that pair illustration with text, which were often used as private announcements for special events [67]. Other surimono featuring fireflies included: T41 (Figure 7), T43-44, T138, and T171.
Kabuki Actors, with July featuring Lady Kasane,' which showcased an onnagata (female impersonator); during that time period in kabuki theaters female roles were played by men [35,66]. It was uncommon to see images that only featured men that were not linked to the Genji story or actors (T34, T117).
Images with larger groups of people showed social gatherings where the evening's entertainment focused on collecting and observing fireflies. Most images had women and men, although sometimes only members of one sex were present, and occasionally children were also in attendance (e.g., T19-20, T56, T70, T88, T112, T128, T144-146). Lanterns, blankets, food, beverages, pipes, musical instruments, and pets were often pictured, emphasizing the recreational aspect. Artists rarely pictured individuals wading in the water while collecting (T18, T186). However, small groups in boats (T10, T60, T133) and larger boating parties were common (T107, T154, T181). 'Firefly Viewing Party' shows a boating party near a bridge where other people were also catching fireflies, and the museum commentary reads, "Hotarugari (firefly viewing) is a popular summer pastime in Japan. These small insects produce flashes of light-which can be seen at night-during the hot months when they breed. Since they live near water, firefly viewing had an added attraction: the cool night breezes off the water brought relief from the heat. The people depicted in this print have hired a boat to take them out on the water. Those in the bow reach towards the fireflies with their fans, attempting to sweep them closer. Those in the center of the boat are drinking and chatting convivially. An attendant blows on a portable stove, attempting to keep a small fire alive so that he can prepare a snack for the group. Twenty-nine haiku poems on the theme of summer are printed in the upper portion of the print." (Minneapolis Museum of Art, T171). This artwork is considered a surimono, or genre of non-commercial woodblock prints that pair illustration with text, which were often used as private announcements for special events [67]. Other surimono featuring fireflies included: T41 (Figure 7), T43-44, T138, and T171. A few images were humorous and involved physical comedy. Hirokage's 'Catching Fireflies at Mount Dōkan' (T34) showed fireflies flying around four men drinking alcoholic beverages. One sumizuri-e print (T180) pictured a standing woman reaching for a firefly and an upended bench with a man tumbling unceremoniously to the ground. 'Thirty-six Amusing Views of Famous Places in Tokyo: Negishi Village' by Ikkei (T128) A few images were humorous and involved physical comedy. Hirokage's 'Catching Fireflies at Mount Dōkan' (T34) showed fireflies flying around four men drinking alcoholic beverages. One sumizuri-e print (T180) pictured a standing woman reaching for a firefly and an upended bench with a man tumbling unceremoniously to the ground. 'Thirty-six Amusing Views of Famous Places in Tokyo: Negishi Village' by Ikkei (T128) depicted a man falling into a stream and fireflies escaping from his airborne cage.
Artwork showing fireflies being used for practical purposes was rare. The design on a kozuka (T147) showed a man reading by the light of a suspended bag of fireflies, and Hearn [68] (p. 459) mentioned, "[ . . . ] story of that Chinese student who, being too poor to pay for a lamp, imprisoned many fireflies in a paper lantern, and thus was able to obtain light enough to study after dark, and to become eventually a great scholar." The museum description of a Chinese painted folding fan featuring multiple men reads, "A firefly lamp hangs from a branch to light the scrolls upon the table." [69].

Tools to Collect and Cage Fireflies
Folding sensu fans and flat uchiwa fans were used to collect fireflies; some were plain but many had ornate designs, including what looks like an actor's face (T77). Uchiwa fans on long poles (T9-10, T44, T88, T109, T112, T128, T140-141, T143-146), or long bamboo poles with leaves at the top are also commonly pictured (T9, T11, T25, T60-61, T110, T128, T138,  T140, T154, T157), although nets of any kind were rare (T29) and may not always be linked to fireflies (T152). People, especially children, also used their hands to capture the insects (e.g., T37, T39, T90, T156). One print (Gesso, T25) showed collection tools and fireflies in a cage in the absence of people. The museum description of the Shinsui print 'Firefly' says, "A young woman is about to bat a firefly with her round fan. She will then place the paralyzed firefly into a cage and collect more to release them all at once later." (Minneapolis Institute of Art, https://collections.artsmia.org/art/62344/firefly-ito-shinsui (accessed on 4 January 2022), a different print of T119).
A wide variety of firefly cages were represented, both in shape, size, and style (Figure 8), and one lacquer box was designed so that it looked like a cage full of fireflies, complete with faux mesh and glowing insects (T176). Cage shapes ranged from cubes, rectangles, tall hexagonal cages, and various types of cylinders, including domes. Figure 8c shows a cylindrical firefly cage made of porcelain with an interlocking pattern of circles (T165). Hearn [34] (p. 148) noted, "The cheapest kind of cage, containing only three or four fireflies, is scarcely more than two inches square; but the costly cages-veritable marvels of bamboo work, beautifully decorated-are as large as cages for song-birds. Firefly cages of charming or fantastic shapes-model houses, junks, temple-lanterns, etc.-can be bought at prices ranging from thirty sen up to one dollar." Cages occurred in many colors, e.g., black, brown, red, yellow, white. Most cages had legs on the bottom and a door to add and release fireflies, and people carried them by a cord attached to the top. The cage in Kuniyoshi's 'Catching Fireflies (T93-94) was incredibly large and ornate, with multiple designs and red tassels. In contrast, an image from the 1950s (T48) showed boys with a cage fashioned from a gourd, although this may be a non-specific insect cage. Firefly cages were typically shown with a tight-weave mesh that occasionally had a floral design (e.g., T89, T95, T113-114). The fine mesh appeared to distinguish firefly cages from other insect cages, such as those for 'singing' insects such as orthopterans, perhaps because the latter can typically chew through fabric mesh (Figure 9).
Based on the morphology of the insects in the images, the cage style, and the surrounding vegetation, artwork labeled as relating to fireflies might actually involve Odonata or Orthoptera. The insects in 'Mother and Children Enjoying Fireflies' (Utamaro, T187) appear to be damselflies or dragonflies, and the child is swatting at them with a thin rod rather than the tools typically used to collect fireflies. Misidentifications were probably due to misinterpretations by people other than the artists, possibly when English titles were added to the artwork. The museum description for a sake dish (T46) reads, "[ . . . ] seven women among autumn plants under a partial moon with three holding an insect cage each trying to catch insects, most probably fireflies, while a group of three men, including two samurai, walk towards the women [ . . . ]." However, the insect pictured in the upper left corner looks like a tree cricket, and there is a distinctive shrub that is pictured in other images where it seems likely that people are hunting Orthoptera instead of fireflies (T21, T30, Figure 10). In some cases, it's difficult to discern if insects or cages relate to Orthoptera or fireflies, e.g., Utamaro's print 'Picture of the Upper Class' (T188), Kuniyoshi's 'Woman with Fan and Insect Cage (T98), and Zeshin's 'Fan and Insect Cage' (T205). Artwork with a likely or definite connection to insects other than fireflies is distinguished in Table A1 by letter codes after the title (see Table A1's footnote 1 for more details). Based on the morphology of the insects in the images, the cage style, and the surrounding vegetation, artwork labeled as relating to fireflies might actually involve Odonata or Orthoptera. The insects in 'Mother and Children Enjoying Fireflies' (Utamaro, T187) appear to be damselflies or dragonflies, and the child is swatting at them with a thin rod rather than the tools typically used to collect fireflies. Misidentifications were probably due to misinterpretations by people other than the artists, possibly when English titles were added to the artwork. The museum description for a sake dish (T46) reads, "[…] seven women among autumn plants under a partial moon with three holding an insect cage each trying to catch insects, most probably fireflies, while a group of three men, including two samurai, walk towards the women […]." However, the insect pictured in the upper left corner looks like a tree cricket, and there is a distinctive shrub that is pictured in other images where it seems likely that people are hunting Orthoptera instead of fireflies (T21, T30, Figure 10). In some cases, it's difficult to discern if insects or cages relate to Orthoptera or fireflies, e.g., Utamaro's print 'Picture of the Upper Class' (T188), Kuniyoshi's 'Woman with Fan and Insect Cage (T98), and Zeshin's 'Fan and Insect Cage' (T205). Artwork with a likely or definite connection to insects other than fireflies is distinguished in Table A1 by letter codes after the title (see Table A1's footnote 1 for more details).

Selling Fireflies
People collected fireflies for personal entertainment, but fireflies were also collected and sold en masse as a commodity [27]. The majority of fireflies for sale in the Japanese modern monarchical period (1868-1945) were field collected and were primarily the larger Genji-boturu (Luciola cruciata) [34,47]. Hearn [34] (pp. 144-146) described in detail how professional firefly-catchers obtained hundreds to thousands of fireflies each night near the Lake ofŌmi to supply large cities such as Kyōto andŌsaka: "Immediately after sunset the firefly-hunter goes forth with a long bamboo pole upon his shoulder, and a long bag of brown mosquito-netting wound, like a girdle, about his waist. When he reaches a wooded place frequented by fireflies,-usually some spot where willows are planted, on the bank of a river or lake,-he halts and watches the trees. As soon as these begin to twinkle satisfactorily, he gets his net ready, approaches the most luminous tree, and with his pole strikes the branches. The fireflies [ . . . ] drop helplessly to the ground [ . . . ] the catcher, picking them up with astonishing quickness, using both hands at once, deftly tosses them into his mouth-because he cannot lose the time required to put them, one by one, into the bag. Only when his mouth can hold no more, does he drop the fireflies, unharmed, into the netting. Thus the firefly-catcher works until about two o'clock in the morning,-the old Japanese hour of ghosts,-at which time the insects begin to leave the trees and seek the dewy soil. There they are said to bury their tails, so as to remain viewless. But now the hunter changes his tactics. Taking a bamboo broom he brushes the surface of the turf, lightly and quickly. Whenever touched or alarmed by the broom, the fireflies display their lanterns, and are immediately nipped and bagged. A little before dawn, the hunters return to town." Fireflies were sorted by the intensity of the light they produced, then several hundred stored in gauze-covered boxes or cages along with moistened grass or turf sprinkled with fresh water [34,71].
Mass-collecting fireflies was not restricted to Japan. In the United States in the 1950s-1980s, children and young biologists collected tens of millions of fireflies for professors, universities and chemical companies, with children recruited to the Sigma Firefly Scientists Club receiving a penny apiece for their bounty [72][73][74].
Laurent [55] indicated that around 1685 in Kyoto insect sellers would carry their wares, specifically singing crickets, in baskets suspended from poles worn across their shoulders. This is akin to Zeshin's print 'Insects Seller' featuring an orthopteran (T204), where two large cages, from which small cages are suspended, hang from a padded pole. Pushcarts that sold insects and cages appeared around 1820 followed by "mushiya, or shops that sold singing insects, fireflies, and jewel beetles as well as cages and trapping devices," in the Meiji period (1868-1912) [55].
Prints or photos of insect sellers included a variety of cages, with and without insects, displayed in what appear to be portable stalls with poles or straps (T3, T57), folding stalls (T28, T76, T189) and indoor or outdoor stands (T114, T137, T190). Several of the stalls or stands had a checkerboard design (T28, T76, T114, T131, T190). Images showed children (T190), women and children (T28, T114, T137), actors as vendors (T57, T76), or lone vendors (T3). Shōtei's 'Insect Seller' (T131) is particularly poignant, with a child in silhouette holding out her firefly cage to a vendor sitting in front of outdoor stall as bats fly overhead.
Hearn [75] (pp. 86-87) described an insect seller at the Market of the Dead, "Hotaru-nikirigisu! [ . . . ] A little booth shaped like a sentry-box, all made of laths, covered with a red-and-white chess pattern of paper; [ . . . ] there are also beautiful little cages full of fireflies,-cages covered with brown mosquito-netting, upon each of which some simple but very pretty design in bright colours has been dashed by a Japanese brush. One cricket and cage, two cents. Fifteen fireflies and cage, five cents." Hoshina [47] reported that based on information in newspapers, the price of a firefly was extremely inexpensive: 0.1 Japanese sen in 1886, and 5 sen in 1917, with 100 sen equal to 1 yen. Hearn [34] (pp. 147-148) wrote, "the wholesale price of living fireflies ranges from three sen per hundred up to thirteen sen per hundred, according to season and quality. Retail dealers sell them in cages; and in Tokyo the price of a cage of fireflies ranges from three sen up to several dollars." However, in 2017, the price of a firefly was much higher, perhaps due to their declining populations, and was listed as 400 yen [47].
Fireflies were sold to individuals, restaurants, hotels, and wholesale and retail insectmerchants [27,34,47]. Fireflies were released at events honoring military victories and royalty and were given out by retailers as free gifts to entice customers [47,59]. "In the famous Dōtombori ofŌsaka, there is a house where myriads of fireflies are kept in a large space enclosed by mosquito-netting; and customers of this house are permitted to enter the enclosure and capture a certain number of fireflies to take home with them." [34] (p. 147).
People often purchased fireflies at pet shops or summer festivals that were kept in cages until the insect died [47]. Fireflies that died in insect-shops still had value and were used in the formulation of drugs and ointments or firefly grease (Hotaru-no-abura) used by woodworkers [34]. Some individuals purchased large quantities of fireflies to release at evening parties or events so that guests could enjoy their sparkling lights [34]. However, one woman wrote, "May 18th 1889. I went to a night fair [ . . . ] there was one stall full of winged lights, tiny stars of green fire clustering all over it. I bought about a hundred Princess Splendours in a black horsehair cage, and brought them home with me. [ . . . ] But the keeper of the strange stall at the fair (and I could hardly see it for the darkness) had captured scores of the winged lights, and sold them by ones and twos in a dainty cage two inches long, with a green leaf for provisions, for two rin, a sum so small that we have no equivalent for it. I stood for a minute before the firefly stall, and then told the interpreter to say that I must have all the fireflies in all the cages. [ . . . ] I carried them all home in the horsehair box; and when everybody had gone to bed, I crept out into the balmy darkness of my garden, opened the box, and set all the lovely creatures free." [76] (pp. [38][39][40][41].

Discussion
This study summarizes the diversity in representations of fireflies and insect-human interactions by a multitude of Japanese artists. Watching and catching fireflies was and continues to be a recreational pastime in many parts of the world, although selling fireflies is rare [47,56]. In Japan, selling insects declined in the 1930s and mushiya were rare by the end of World War II [55], although there is still interest in insects as a commodity [77,78]. Insects such as fireflies remain an important part of Japanese culture [31,47,56], and are protected by legal and social policies [27]. Firefly ecotourism (e.g., celebrations, festivals, tours of firefly sanctuaries, firefly 'villages'), has emerged in multiple countries, which can serve as a potential tool to educate the public about firefly conservation and threats to their populations, including habitat degradation, water and light pollution, and insecticides [10,23,29,[79][80][81][82][83][84]. Ensuring that these revenue-generating activities do not negatively impact fireflies or their habitat is an important component of sustainable firefly ecotourism [10,[82][83][84][85]. The concept and definition of environmental art, or eco-art, has evolved throughout time, but in general eco-art education is a multidisciplinary approach that integrates art, science, and education with a focus on the environment, including biodiversity, conservation, sustainability, and restoration [86][87][88]. Using art to highlight the historical and cultural significance of fireflies may also help with conservation efforts. Acknowledgments: Thank you to the art museums and the people associated with the ukiyo-e database (ukiyo-e.org) for making artwork accessible online.

Conflicts of Interest:
The author declare no conflict of interest.