Luciola

Luciola is a genus of flashing fireflies in the family Lampyridae. They are especially well known from Japan and are often called Japanese fireflies, but their members range farther into Asia and reach southern Europe and Africa. This genus is traditionally held to extend to Australia, but these species do not seem to belong herein.

Biological description

Unlike some other fireflies, the females of Luciola are fully winged. Ecologically, the genus is remarkable in that the larvae of several species are aquatic. The adults of the typical Luciola are similar to those of Atyphella which was formerly included in this genus. They can be easily distinguished by the males' aedeagus however, which in Luciola proper has large lateral lobes that do not taper and have elongated, slender and flattened smaller lobes along their ventral margin. The middle lobe of the males' aedeagus is very slim and has a point before the tip. By contrast, the Australian group resembles Pyrophanes in its lateral lobes of the aedeagus being small and not visible from beneath. But unlike in Pyrophanes, the sheath of the aedeagus lacks paraprocts, and the seventh ventrite of the abdomen lacks the hairy lobes and points at its hind margin.
Two Japanese species of Luciola, collectively known as hotaru, are significant in Japanese culture and folklore. They are symbols of the hitodama, the souls of the newly-dead. See also the explanations at the article on the movie Hotaru no Haka for a discussion of the cultural significance of the hotaru.

Systematics

Luciola in the narrow sense appears to be most closely related to a rather basal group of Luciolinae, including Bourgeoisia and Lampyroidea and Hotaria. Their relationships are not well resolved and Hotaria is sometimes merged with Luciola. On the other hand, the genera Atyphella and Pygoluciola, which for some time were included in Luciola, is now again recognized as distinct.
The internal systematics of Luciola are somewhat in need of revision, and it is not certain whether these species are all correctly assigned to this genus. For example, the Australian species appear to be separate and not very closely related to the other members of this genus, but rather to Colophotia, Pteroptyx and Pyrophanes. Whether the larvae of these can be recognized by the share soft terga lacking paranota, or whether this is a plesiomorphic trait also present in other Luciolinae, remains to be determined.
L. deplanata might be worthy of separation in a monotypic genus Photuroluciola.
But considering the fact that many species remain essentially unstudied, a 2008 review of the genus concluded that:

"A phylogenetic analysis may give indications for subdivisions within Luciola. However, there are elements of the cart and the horse here - which should come first? Break up Luciola based on narrow analyses, or wait until more analyses are available? It is our contention that those Luciolinae species which do not fit elsewhere should either be described as Luciola sp. or remain undescribed until such time as phylogenetic analyses indicate a repeatable pattern of subdivisions with Luciola. Any other action would be, in our opinion, premature."

In 2010, leii, ficta, hydrophila and lateralis were transferred to the genus Aquatica.

Species

These species belong to the genus Luciola:Luciola affinis Ritsema, 1875 Luciola amplipennis Fairmaire, 1880 Luciola angusticollis E. Olivier, 1886 Luciola antennalis Bourgeois, 1905 Luciola antica Boisduval, 1835 Luciola antipodum Bourgeois, 1884 Luciola apicalis Luciola aquilaclara Ballantyne, 2013 Luciola atriceps Pic, 1916 Luciola atrimembris Pic, 1933 Luciola atripes Pic, 1929 Luciola atritarsis Pic, 1916 Luciola atriventris Pic, 1925 Luciola aurantiaca Pic, 1927 Luciola auritula E. Olivier, 1910 Luciola basipennis Pic, 1958 Luciola bicoloriceps Pic, 1924 Luciola bimaculata Klug, 1855 Luciola bimaculicollis Luciola binhana Pic, 1927 Luciola binotata Pic, 1951 Luciola biscutellata Fairmaire, 1896 Luciola bomansi Pic, 1956 Luciola bourgeoisi E. Olivier, 1895 Luciola burgeoni Pic, 1931 Luciola caffra Luciola candezei E. Olivier, 1902 Luciola capensis Luciola carinaticollis Pic, 1952 Luciola cavifrons Fairmaire, 1900 Luciola chapaensis Pic, 1923 Luciola christyi Pic, 1931 Luciola cincta Luciola cincticollis Klug, 1855 Luciola cingulata E. Olivier, 1885 Luciola cisteloides Klug, 1855 Luciola clara E. Olivier, 1907 Luciola collarti Pic, 1931 Luciola complanata Gorham, 1895 Luciola completelimbata Pic, 1952 Luciola concreta E. Olivier, 1907 Luciola congoana E. Olivier, 1895 Luciola consobrina Hicker, 1942 Luciola costulata Fairmaire, 1886 Luciola coxalis E. Olivier, 1883 Luciola cruciata Motschulsky, 1854 genji-botaru Luciola curticollis Pic, 1927 Luciola curtithorax Pic, 1928 Luciola davidis E. Olivier, 1895 Luciola delauneyi Bourgeois, 1890 Luciola deleta Fairmaire, 1896 Luciola deplanata Pic, 1929 Luciola dilatata Pic, 1929 Luciola dilecta E. Olivier, 1908 Luciola discicollis Laporte, 1833 Luciola discobrunnescens Pic, 1951 Luciola discoidalis Pic, 1931 Luciola dregei Luciola errans E. Olivier, 1902 Luciola exigua Luciola exoleta Motschulsky, 1854 Luciola exstincta E. Olivier, 1886 Luciola extricans Luciola falsa E. Olivier, 1907 Luciola fasciiventris Fairmaire, 1898 Luciola filiformis E. Olivier, 1913 Luciola fissicollis Fairmaire, 1891 Luciola flava Pic, 1929 Luciola flavescens Luciola flebilis E. Olivier, 1909 Luciola fukiensis Pic, 1955 Luciola fusca Luciola fuscula Luciola gaiffei Alluaud, 1899 Luciola galactopyga Fairmaire, 1884 Luciola gaudens E. Olivier, 1910 Luciola gemina E. Olivier, 1905 Luciola geniculata E. Olivier, 1902 Luciola ghesquierei Pic, 1934 Luciola gigantea Pic, 1916 Luciola gigas E. Olivier, 1888 Luciola gowdeyi Pic, 1933 Luciola grandjeani Pic, 1931 Luciola guerini Laporte, 1833 Luciola hirticeps E. Olivier, 1888 Luciola horni Bourgeois, 1905 Luciola hypocrita E. Olivier, 1888 Luciola ibukiyamana Nawa, 1921 Luciola imbellis E. Olivier, 1908 Luciola imerinae Bourgeois, 1899 Luciola immarginata Bourgeois, 1890 Luciola impedita E. Olivier, 1902 Luciola incerta Luciola infuscata Luciola inlateralis Pic, 1952 Luciola innotatipennis Pic, 1916 Luciola insignis E. Olivier, 1883 Luciola intermedia Fairmaire, 1896 Luciola intricata Luciola italica Luciola japonica Luciola jeanvoinei Pic, 1927 Luciola jengai Nada, 2019 Luciola judaica E. Olivier, 1884 Luciola kagiana Matsumura, 1928 Luciola kervillei E. Olivier, 1909 Luciola klapperichi Pic, 1955 Luciola krugeri E. Olivier, 1905 Luciola kuroiwae Matsumura, 1918 Luciola laeta Gerstaecker, 1871 Luciola lambkinae Keller & Ballantyne, 2021 Luciola latediscoidalis Pic, 1952 Luciola latelimbata Pic, 1956 Luciola laticollis Gorham, 1883 Luciola leroyi Pic, 1952 Luciola limbalis Fairmaire, 1889 Luciola limbatipennis Pic, 1923 Luciola longula E. Olivier, 1914 Luciola lucernula Reiche, 1850 Luciola lusitanica Luciola maculiscuta Fairmaire, 1884 Luciola madagascariensis Luciola magambae McDermott, 1963 Luciola maindroni Pic, 1927 Luciola major Pic, 1952 Luciola mareei Pic, 1952 Luciola mediodivisa Pic, 1952 Luciola mediojuncta Pic, 1952 Luciola melancholica E. Olivier, 1913 Luciola melanura Laporte, 1833 Luciola mendax E. Olivier, 1905 Luciola mocquerysi E. Olivier, 1902 Luciola mongolica Motschulsky, 1860 Luciola multicostulata Pic, 1927 Luciola mutata Pic, 1939 Luciola niah Jusoh, 2019 Luciola nicolleri Bugnion, 1922 Luciola nigripes Gorham, 1903 Luciola nigritarsis Pic, 1927 Luciola nitidicollis Pic, 1916 Luciola nitidula Luciola noctivaga E. Olivier, 1907 Luciola notaticollis Pic, 1914 Luciola novaki Müller, 1946 Luciola obscura Pic, 1928 Luciola obscuripennis Klug, 1855 Luciola ochracea Gorham, 1895 Luciola oculofissa Ballantyne, 2013 Luciola olivieri Bourgeois, 1908 Luciola overlaeti Pic, 1931 Luciola owadai Satô & Kimura, 1994 - Kumejima-botaru - Luciola pallida Kolbe, 1883 Luciola pallidipes Pic, 1928 Luciola papariensis Doi, 1932 Luciola parvula Kiesenwetter, 1874 Luciola pattersoni Pic, 1935 Luciola perpetiuscula Kolbe, 1887 Luciola perspicua E. Olivier, 1907 Luciola picea Gorham, 1882 Luciola pici McDermott, 1966 Luciola picticollis Kiesenwetter, 1874 Luciola pieli Pic, 1939 Luciola pouilloni Pic, 1931 Luciola praestans E. Olivier, 1896 Luciola praetermissa E. Olivier, 1909 Luciola pumila Luciola puncticollis Laporte, 1833 Luciola quadrimaculata Pic, 1952 Luciola recticollis E. Olivier, 1900 Luciola robusticeps Pic, 1928 Luciola roseicolllis Pic, 1933 Luciola ruficollis Luciola rugiceps E. Olivier, 1886 Luciola satoi Jeng & Yang, 2003 Luciola scheitzae Pic, 1952 Luciola semiventralis Fairmaire, 1899 Luciola semivittata Pic, 1952 Luciola septemmaculata E. Olivier, 1888 Luciola singapura Jusoh & Ballantyne, 2021 Luciola sordida E. Olivier, 1905 Luciola spectralis Gorham, 1880 Luciola spilodera Fairmaire, 1898 Luciola stanleyi E. Olivier, 1913 Luciola stigmaticollis Fairmaire, 1887 Luciola subunicolor Pic, 1931 Luciola sudra Gorham, 1903 Luciola syriaca Luciola tabida Gorham, 1880 Luciola taeniaticollis Fairmaire, 1886 Luciola tenuicornis E. Olivier, 1885 Luciola testaceipes Pic, 1916 Luciola testaceiventris Pic, 1916 Luciola tetrasticta Fairmaire, 1888 Luciola tincticollis Gorham, 1895 Luciola tiomana Ballantyne, 2019 Luciola trimaculata Pic, 1916 Luciola trivandrensis Raj, 1947 Luciola tsushimana Nakane, 1970 Luciola tuberculata Yiu, 2017 Luciola turneri Pic, 1934 Luciola umbratica E. Olivier, 1907 Luciola unmunsana Doi, 1931 Luciola varia E. Olivier, 1908 Luciola vittata Laporte, 1833 Luciola vleeschouwersi Pic, 1952 Luciola xanthochroa Fairmaire, 1904 Luciola xanthura Gorham, 1880