Oculogryphus, A Remarkable New Genus of Fireflies from Asia (Coleoptera: Lampyridae)

2007·4.6 MB·English

by  M.-L. Jeng

PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY
CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024
Number 3600, 19 pp., 13 figures, 1 table December 12, 2007
Oculogryphus, A Remarkable New Genus of Fireflies
from Asia (Coleoptera: Lampyridae)
M.-L. JENG,1 MICHAEL S. ENGEL,2 AND P.-S. YANG3
ABSTRACT
Anunusualnewlampyridgenus,OculogryphusJeng,Engel,andYang,newgenus,anditstype
species,OculogryphusfulvusJeng,newspecies,fromVietnam,isdescribedandfigured.Thegenus
ischaracterizedbyitsfiliformantennae,largeandventrallyapproximatecompoundeyesthatare
deeply emarginate posteriorly, largely exposed head (i.e., not covered by the pronotum), eight
abdominalventrites,rectangularabdominaltergites,absenceofphotogenicorgans,andsymmetric
aedeagalsheathandgenitaliainthe male (thefemaleis presentlyunknown). Thegenusdoesnot
adequatelyfitintoanysubfamilyastheyarecurrentlydefined,highlightingthedifficultiesofthe
subfamilial and tribal classification presently in use. Though most likely allied to Ototretinae or
Luciolinae,thesystematicpositionofOculogryphuscannotbedeterminedascurrentlythereisno
comprehensivephylogenyofLampyridaeandrelatedgroups.Thespeciesmayrelyonphoticcues
for purposes of mating. A key to the genera of Lampyridae (sensu Crowson) occurring in
PalaearcticEastAsia,Oriental,andAustralianregionsisprovided.Sometaxonomicproblemsof
Ototretinaeare commented upon.
INTRODUCTION associated with human beings. Biolumines-
cence is an attractive and useful feature of
The family Lampyridae, commonly known these beetles, though it is not unique to
as lightning bugs, fireflies, or glow worms in Lampyridaeandnotalllampyridspeciesglow
English, is one of the beetle groups closely or flash (McDermott, 1964; Lloyd, 1971).
1Division of Entomology, Natural History Museum, and Department of Ecology & Evolutionary Biology, 1501
CrestlineDrive–Suite140,UniversityofKansas,Lawrence,Kansas66049-2811([email protected]).
2DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory;DivisionofEntomology(Paleoentomology),
Natural History Museum, and Department of Ecology & Evolutionary Biology, 1501 Crestline Drive – Suite 140,
UniversityofKansas,Lawrence,Kansas66049-2811([email protected]).
3Department of Entomology, National Taiwan University, #1, Sec. 4, Roosevelt Road, Taipei 106, Taiwan
([email protected]).
CopyrightEAmericanMuseumofNaturalHistory2007 ISSN0003-0082
2 AMERICAN MUSEUMNOVITATES NO. 3600
Glowing ability can also be found in several 2002, 2003b; Fu et al., 2005a). About 2000
elateroid families such as Elateridae, species have been described (Nakane, 1991),
Omalisidae, Phengodidae, and Rhagophthal- and a considerable portion of them are active
midae (Lloyd, 1971, 1979; Crowson, 1981; in the daytime (McDermott, 1964; Lloyd,
Branham and Wenzel, 2003; Grimaldi and 2002; Branham and Wenzel, 2003). Though
Engel, 2005). Confirmed bioluminescent attractive,diverse,anduseful,Lampyridaedid
display for the purpose of mating is known nothaveacleardefinitionuntilrecently.Their
in Rhagophthalmidae and Lampyridae family-group compositions and boundaries
(Branham and Wenzel, 2003), and precise were altered frequently and dramatically
control over bioluminescence is unique to the during the past century. The modern classifi-
latter(Trimmer etal.,2001).Thebrilliantand cationofLampyridaewasfundamentallybuilt
delicate bioluminescent behavior has made on the basis of Olivier (1907, 1910), then
fireflies common subjects in cultural activities modifiedandimprovedbyGreen(1948,1959),
andworksofartsinceearlytimes,especiallyin Crowson (1955, 1972), McDermott (1964,
Eastern Asian countries (Kanda, 1935; 1966), Wittmer (1979), Nakane (1991),
Harvey, 1957; Yamamoto, 1968; Konishi, Lawrence and Newton (1995), Jeng et al.
1997;Yang,1998).Nowadaysfirefly-watching (1998a, 2006a, 2006b), Lawrence et al. (1999),
isapopularandseasonallyroutineactivityfor and Branham and Wenzel (2001). Olivier
ecological tourism in several Asian countries (1907) published the first lampyrid catalogue
(Othman and Othman, 1998; Chen, 1999, in which he constructed a nine-subfamily
2003; Ho et al., 2002; Nallakumar, 2003; system, largely based on antennal and head
Ohba, 2004). Fireflies are also good subjects morphology. McDermott’s seven-subfamily
for education, conservation, and ecological system (1964, 1966) reduced several of
and biodiversity research (e.g., Furuta, 1991; Olivier’s subfamilies to subordinate units
Kobayashi, 1991; Murakami, 1991; Lloyd, of Lampyrinae and added Matheteinae,
1997, 1998, 1999, 2000, 2001, 2003; Faust, Rhagophthalminae, and Pterotinae. Ototre-
2004; Ohba, 2004). In a practical application, tinae was first treated as a subfamily in
the firefly luciferase gene and the enzyme it McDermott (1964) but later demoted to
codes for has been used as an indicator or a lucioline tribe in McDermott (1966).
probe in biological and medical sciences, McDermott (1964) also provided the latest
thereby further highlighting the broad impor- comprehensive accounts on the family-group
tance of lampyrids for human endeavors definitionsandcompositions.Crowson(1972)
(Grimaldi and Engel, 2005). clarified the separation of Lampyridae from
Lampyridae are small to medium-sized the other cantharoid families. He removed
beetles, usually not longer than 3 cm, though Rhagophthalminae to Phengodidae and
somelarviformfemalescanbeaslargeas8cm Matheteinae to Omethidae, transferred many
inlength.Theyarenearlygloballydistributed, genera of Drilidae to Ototretinae, and estab-
particularly thriving and diverse in warm, lished Cyphonocerinae and Ototretadrilinae
humid environments (McDermott, 1964). based on former drilid genera. Crowson did
The larvae are mostly terrestrial and active not address much on the content of each
huntersoropportunisticscavengersfeedingon subfamilybutgaveatentativekeytotheeight
living or dead small invertebrates and/or lampyrid subfamilies he suggested. In total,
vertebrates (Lawrence and Britton, 1994; 23 family-group taxa based on different type
Lloyd, 2002; Ho and Chu, 2002; Chen, genera have been proposed across the
2003). Some unusual habitats including an three major classifications (Olivier, 1907;
intertidal spray zone, ant nests, marshes, McDermott, 1964; Crowson, 1972) (table 1).
running water, seepage environments, hot The classification of Lampyridae currently
springs, and decaying tree trunks have also used is essentially a mix of McDermott’s and
been reported (Annadale, 1900, 1906; Blair, Crowson’ssystems:adoptingCrowson’seight-
1927;Imms,1933;BallantyneandBuck,1979; subfamily system and following McDermott’s
Buschman, 1984; Ohba and Goto, 1991; generic composition of each subfamily (e.g.,
Sivinski et al., 1998; Lloyd, 2002; Jeng et al., Lawrence and Newton, 1995; Lawrence et al.,
2007 JENGET AL.: NEWVIETNAMESE FIREFLY GENUS 3
TABLE1
Modern Family-group Classifications ofLampyridae
Olivier,1907 McDermott,1966 Crowson,1972a
(9subfamilies,48genera) (7subfamilies,85genera) (8subfamilies)
SubfamilyLampyrinaeLatreille(9)b SubfamilyLampyrinaeLatreille(53) SubfamilyLampyrinaeLatreille
SubfamilyLamprocerinaeOlivier(14) TribeLampyriniLatreille(11) SubfamilyAmydetinaeOlivier
SubfamilyPhotininaeLeConte(6)c TribePleotominiGreen(5) SubfamilyPhoturinaeLacordaire
SubfamilyLucidotinaeLeConte(7) TribeLamproceriniOlivier(5) SubfamilyLuciolinaeLacordaire
SubfamilyDadophorinaeOlivier(1) TribeCratomorphiniGreen(5) SubfamilyOtotretinaeMcDermott
SubfamilyMegalophthalminae TribePhotininiLeConte(27) SubfamilyPterotinaeLeConte
Olivier(2) SubtribePhotininaLeConte(13) SubfamilyCyphonocerinaeCrowson
SubfamilyAmydetinaeOlivier(1) SubtribeLuciodotinaLacordaire(10) SubfamilyOtotretadrilinaeCrowson
SubfamilyPhoturinaeLacordaire(1) SubtribeDadophorinaOlivier(1)
SubfamilyLuciolinaeLacordaire(7) SubtribePhosphaeinaMcDermott(2)
SubtribeLamprigerinaMcDermott(1)
SubfamilyAmydetinaeOlivier(12)
TribeAmydetiniOlivier(2)
TribeVestiniMcDermott(5)
TribePsilocladiniMcDermott(5)
SubfamilyPhoturinaeLacordaire(4)
SubfamilyLuciolinaeLacordaire(9)
TribeLucioliniLacordaire(6)
TribeCurtosiniMcDermott(1)
TribeOtotretiniMcDermott(2)
SubfamilyMatheteinaeLeConte(2)
SubfamilyPterotinaeLeConte(1)
SubfamilyRhagophthalminaeOlivier(4)
aThemorerecentclassificationsofNakane(1991)andBranhamandWenzel(2001)recognizedthesameeightsubfamilies
although with modified generic compositions for Lampyrinae, Amydetinae, and Cyphonocerinae (the synonymy of
CyphonocerinaebyJengetal.[1998a]wasbasedonNakane’s[1991]redefinitionofthesubfamily).BranhamandWenzel’s
cladisticstudyindicatedthatLampyrinae,Amydetinae,andOtotretinaewerepolyphyletic;thatLuciolinaeandperhaps
Photurinaeweremonophyletic;thatOtotretinaeandPterotinaeshouldbeconsideredincertaesedisinElateroidea;whilethe
statusoftwosubfamiliescouldnotbecommentedupon(OtotretadrilinaewasexcludedandCyphonocerinaewasrepresented
byasinglegenus).Kazantsev(2006)createdCheguevariniandtentativelyplacedthetribeasincertaesedisinLampyridae.
bValueinparenthesesindicatesthenumberofexplicitlyincludedgenera.
cAnamehomonymouswiththesubfamilialnamePhotininaeGiglio-Tosi,intheMantidae(Mantodea).
1999),orwithsomeminormodifications(e.g., classification ofthefamily. Hereinwe provide
Wittmer, 1979; Nakane, 1991; Jeng et al., a description of this enigmatic new group,
1998a). However, Branham and Wenzel’s making its name available for forthcoming
(2001) phylogenetic analysis revealed that all contributions to the familial classification, to
ofthesubfamiliescurrentlyrecognized,except alert coleopterists to its presence and discuss
Luciolinae and perhaps Photurinae, are para- its systematic position among other fireflies.
phyletic or polyphyletic, or not lampyrids at
all. The definitions of Lampyridae and their
MATERIAL AND METHODS
subordinate units will need a comprehensive
revision (underway by the senior author). The material was from the late Dr. M.
Recently we identified a remarkable new Satoˆ’spersonalcollection.Thegluedspecimen
genus of Lampyridae from Vietnam. Its was first softened and released in hot water
unique combination of characters does not and then preserved in 80% ethanol solution
fit the definition of any subfamily or tribe/ for dissection and illustration. Male genital
subtribe currently recognized, further high- segments were dissected and soaked in 10%
lighting the difficulties of the present higher potassiumhydroxide(KOH)solutionatroom
4 AMERICAN MUSEUMNOVITATES NO. 3600
Fig.1. HabitusofholotypemaleofOculogryphusfulvus,newgenusandspecies,dorsal(left)andventral
(right) aspect.
temperature for 24 hours prior to examina- ‘‘ventrite’’ for the purpose of visual diagnosis
tion.TheabbreviationsBL,BW,EL,EW,PL, without difficulty, and ‘‘sternite’’ for the
and PW are employed for ‘‘body length’’, identification of morphological homologues
‘‘body width’’, ‘‘elytral length’’, ‘‘elytral across taxa. The nomenclature of hind-wing
width’’, ‘‘pronotal length’’, and ‘‘pronotal venation follows that of Kukalova´-Peck and
width’’, respectively. Body length is the Lawrence (1993).
distance between the anterior head margin
and elytral apex; body width is the greatest SYSTEMATICS
distanceacrosstheelytraortwicethewidthof
an elytron (BW 5 2EW). The term ‘‘ventrite’’ Oculogryphus Jeng, Engel, and Yang,
is used for the visible abdominal sternite; T# new genus
andS#replacethetrue#thtergiteorsternite
of the abdomen, respectively; the last abdom- TYPE SPECIES: Oculogryphus fulvus Jeng,
inal tergite is T8; ‘‘aedeagal sheath’’ is new species (fig. 1).
composed of a syntergite (T9 + 10) and DIAGNOSIS: The new genus can be readily
sternite IX [S9 5 ventrite 8 (V8)]. It should diagnosed by the following combination of
be noted that Ballantyne and Lambkin (2001, characters:partiallyexposedhead;11-articled,
2006) used ventrite to reflect the sternite filiform antennae; large compound eyes that
of true segmentation (e.g., V8 5 S8). We are emarginate posteriorly and approximate
have followed the typical method of using ventrally; strongly curved and crossing man-
2007 JENGET AL.: NEWVIETNAMESE FIREFLY GENUS 5
DuPorte, 1960) weakly convex; antennal
socket elongate elliptical in shape, moderately
distant from labrum, with antennifer in lower
margin of socket; space between antennal
sockets slightly convex, about as broad as
one-halfwidthofsocket;antennae11-articled,
filiform, reaching elytral base when in repose;
scape and pedicel elongate, longer in former;
flagellararticlescylindricalanddenselysetose.
Lowermarginofclypeusbroadlyandroundly
emarginate.Labrumsclerotized,notchedmed-
ioapically. Mandibles well developed, strong
and nearly uniform in diameter to near tip,
curved and pointed apically. Maxillae with
cardines approximate each other at base;
maxillary palpi with four palpomeres, labial
palpiwiththreepalpomeres,bothwithslightly
dilated, thick, terminal palpomeres and bud-
like apices, without thin ridge or tooth on
inner side. Ventral margin of occipital fora-
men emarginate, reaching basal one-third of
head length. Gula very narrow. Posterior
tentorial pits immediately behind labrum, at
about middle of head. Margins of hypostoma
convergent toward base (fig. 4).
Figs. 2–3. Oculogryphus fulvus, new genus and
Pronotum (fig. 2) transverse, subparallel-
species,male,headandpronotum.2.Dorsalaspect.
sided, punctate, and pubescent throughout;
3.Lateral aspect.
mostly opaque although translucent on mar-
gins; anterior margin broadly rounded, with-
dibles with pointed apex; narrowly explanate
out clear anterolateral angles; central disc
pronotal margins and close pronotal hypo-
evenly convex (fig. 3), with median sulcus;
meron; epipleuron restricted to basal half of
apical and lateral explanate margins very
elytra; eight abdominal ventrites (including
narrow; posterolateral angles large and nearly
exposed sternite of aedeagal sheath); abdom-
orthogonal; posterior margin weakly sinuate,
inal tergites not lobed; absence of photogenic
broadly impressed on inner base of posterior
organs and tibial spurs; and progressively
shortened tarsomeres 1–4. These characters angles. Hypomeron close in frontal aspect,
intermingle diagnostic features of Luciolinae, with inner margin attending anterolateral
Rhagophthalminae,andOtotretinae,butnone margin of pronotum. Prosternum (fig. 4) with
of the three subfamilies possess all of these an inverted Y-shaped, short and broad inter-
characters. coxal process. Mesoventrite separated from
DESCRIPTION: Male. Body shape elongate mesopleurites by a clear suture (fig. 4).
oval, somewhat depressed and fully winged. Metaventrite notched medioapically. Elytra
Head (fig. 2) largely exposed from pronotum elongate oval, well paired along midline;
when retracted, intermediate between progna- carinae weak; lateral explanate margins nar-
thous and hypognathous. Compound eyes row; surface covered with fine setae. Humeral
very large, occupying most of head laterally area of elytra (fig. 5) roundly folded toward
andwithhindmarginsremarkablyemarginate marginal costa; epipleura narrow, deeply
(fig. 3); compound eyes separated from each folded, barely visible laterally, extending from
otherindorsalaspectbyaboutone-thirdhead humeral base to basal one-third of elytra.
width and approximate ventrally (fig. 4). Venation of hind wing (fig. 8) with reduction
Vertex flat or slightly depressed. Antennal of crossveins; radial cell incomplete; MP
3+4
calli (convexity above antennal sockets: see and CuA not connected by crossveins and
1+2
6 AMERICAN MUSEUMNOVITATES NO. 3600
Figs. 4. Oculogryphus fulvus,new genus andspecies, male, head andthorax, ventral aspect.
not branched. Legs long and slender. Front Abdomen with eight ventrites (fig. 11, S2–
trochantins setiferous but glabrous in middle 9), reaching elytral apices. Spiracles in lateral
legs. Mesocoxae moderately separated from folded, membranous pleurites, not visible
each other; metacoxae narrowly separated ventrally. Tergites invisible in ventral aspect,
from each other. Tibial spurs absent. Tarsi not lobed and with rounded posterior angles.
(fig. 9) about as long as their tibiae in middle Apical margin of ventrite 7 (5 S8) simple;
and hind legs; tarsomeres progressively short- exposed portion of V8 (5 S9) semirounded.
er from 1–4, cylindrical and slender in 1–3; Aedeagal sheath basically symmetric, broad
tarsomere 4 with tarsal pulvilla and lobed, andshort;T9andT10individuallydistinct;S9
widely open in dorsal cleft (fig. 9); pretarsal broad at base. Male genitalia modified tri-
claws simple. lobed, bilaterally symmetric; much smaller
2007 JENGET AL.: NEWVIETNAMESE FIREFLY GENUS 7
Oculogryphus fulvus Jeng, new species
Figures 1–13
TYPE LOCALITY: Tam Dao, Province Vinh
Phu, Vietnam.
DIAGNOSIS: As for the genus (see above).
DESCRIPTION: Male. BL 6.0 mm, BW 2.5
mm; body form elongate oval and somewhat
depressed; overall coloration brown except
vertex black; antennal scape and pedicel
brown, flagellar articles dark brown; elytra
brown, looking darker from basal fifth to
apices due to shadows of hind wings; venter
brown throughout. Head with vertex and
beyond exposed from above, not hidden by
pronotum; vertex flat between eyes, surface
glabrous, densely punctate and pubescent.
Distance between compound eyes about two-
fifths of greatest width of head in dorsal
aspect, one-fifth in frontal aspect (fig. 7), and
one-sixth in ventral aspect. Antennae (fig. 6)
withscape elongatequadrate in dorsal aspect,
about as long as first flagellar article (FA1);
FA1–3 each subequal in length, 4–8 each
subequalinlengthandshorterthan1–3;FA9
(terminal antennomere) spindle-like.
Pronotum subparallel-sided; posterior an-
gles sharply cornered; punctures on disc
Fig. 5. Oculogryphus fulvus, new genus and separated by about their own diameter
species, male,basal half of elytra,lateralaspect. (fig. 2). Scutellum large and triangular in
shape. Elytra with well-defined lateral mar-
than aedeagal sheath; median lobe uniformly gins; surface more or less shagreened, densely
broad, arched upward in apical half; para- setose, costae insignificant except sutural
meresbroadinlateralaspectandformingaV- costae. Elytral epipleuron (fig. 5) deeply
shapedbandindorsalaspect;basalpiecelarge folded, lying in ventral side of elytra and
and symmetric. nearlyinvisiblefromlateralaspect,aboutone-
Female. Unknown. sixth of elytral length. Mesoventrite (fig. 4)
ECOLOGY: Unknown; see Discussion, be- broadlyV-shaped.Central longitudinal sulcus
low. of metaventrite obsolete. Abdominal T8
ETYMOLOGY: The new genus-group name (fig. 10) broadly rounded apically and slightly
is a combination of the Latin terms oculus emarginatemedioapically,S8(fig. 11)subtrap-
(meaning ‘‘eye’’) and gryphus (meaning ‘‘grif- ezoidal.PW/PL51.4;EL/EW53.3;EL/PL5
fin’’, originally a mythological creature from 3.4;BW/PW51.5.
AsiaadoptedbytheGreeksandwithamixof Aedeagal sheath (fig. 12) about 0.7 mm
features from a lion and an eagle. The idea of long, 0.4 mm broad; syntergite slightly sur-
the griffin perhaps stemmed from ancient passing apex of S9; T9 about as long as T10;
people’s seeing the exposed remains of T10 triangular; S9 setose in apical one-third,
Protoceratops: Mayor, 2000). The name is rounded at apex, tapering toward base.
a reference to the characteristic eyes and the Genitalia (fig. 13) about 0.5 mm long, 0.3
ambiguous, subfamilial identity of the new mmbroad;medianlobeparallelsidedinbasal
genus. The name is masculine. half and gradually tapering apically in dorsal
DIVERSITY: The type species is presently aspect; paramere weakly sclerotized apically
the only known member of the genus. but strongly so in posterior margin; dorsum
8 AMERICAN MUSEUMNOVITATES NO. 3600
Figs. 6–9. Oculogryphus fulvus, new genus and species, male. 6. Antenna. 7. Head, frontal aspect;
antennaeandmostofthemouthpartsexceptmandiblesandlabrumwerenotshown.8.Hindwing.9.Hind
leg, trochanterto tarsus.
forming a widely V-shaped band connecting ETYMOLOGY: The specific epithet refers to
withmedianlobe;basalpieceaboutaslongas the light brown coloration of the species.
median lobe, bilaterally symmetric, roughly
a V-shaped band, slightly pointed apically.
Female. Unknown. KEYTO LAMPYRID GENERAIN EASTAND
TYPE MATERIAL: Holotype, male; ‘‘N. SOUTHEAST ASIAAND AUSTRALIA
Vietnam 1985, pr. Vinh phu, Tam dao 3.6.- (males only)
11.6. V. Sˇvihis lgt.’’. The holotype will be TheconsideredarearangesfromIndiainthe
preservedintheNationalMuseum ofNatural westandAustraliaintheeast,zoogeograph-
Science, Taichung City, Taiwan (NMNS). ically including the whole Oriental region
2007 JENGET AL.: NEWVIETNAMESE FIREFLY GENUS 9
Figs.10–13. Oculogryphus fulvus, new genusand species, male.10. Abdominal tergite8,dorsal aspect.
11. Abdominal visible sternites, right half, ventral aspect. 12. Aedeagal sheath, dorsal aspect. 13. Male
genitalia,dorsal(A), ventral (B),andlateral (C)aspects.
(OR, including Indian subcontinent, Ryukyu Archipelago), New Guinea, Pacific
Himalayas, Indochina and Malay islands, and Australia of the Australian
Peninsula, Taiwan, Philippines, Borneo, region (AU). The proposed key, following
and Sunda Islands), the Palaearctic region Crowson’s definition of Lampyridae, pro-
in EastAsia(PA,includingEasternEurasia vides an account of the 34 lampyrid genera
continent, Korean Peninsula, and Japan- presently documented in this area (inclusive
10 AMERICAN MUSEUMNOVITATES NO. 3600
of the new genus). It is partly based on or 6. Apices of parameres of genitalia largely or
modified from Ballantyne and Lambkin totally concealedbymedian lobe,notvisible
(2001, 2006, for couplets 3–10), Maulik in ventral aspect .. .. .. . .. .. .. .. .. . 7
— Apices of parameres of genitalia barely or not
(1921)andWittmer(1979)(forcouplets22–
concealed by median lobe, visible in ventral
24), and Wittmer (1937, for couplet 33). It
aspect. . .. .. .. .. .. .. . .. .. .. .. .. 10
should be noted that the key is tentatively
7. Apexof elytradeflexed. .. . .. .. .. .. .. ..
proposed, especially for the nontypical lam-
. .. .. . .. .. . PteroptyxOlivier (OR, AU)
pyrids (couplet 20 and below). With the
— Apexof elytranotdeflexed. .. .. .. .. .. . 8
exception of Oculogryphus, most of the 8. Median carinaof V6present.. .. .. .. .. ..
nontypicallampyridgeneraweretransferred .................ColophotiaDejean(OR)
from Drilidae and placed under Ototretinae — Median carinaof V6absent .. .. .. .. .. . 9
orOtotretadrilinaebyvariousauthors.Most 9. Photogenic organs on V6 bipartite; aedeagal
ofthemarepoorlyknownatpresent,usually sheath bearingparaprocts. .. .. .. .. .. ..
with no more than revisional works on .............PyrophanesOlivier(OR,AU)
— Photogenic organs on V6 one-pieced; aedeagal
restricted zoogeographic faunas (e.g.,
sheath withoutparaprocts .. .. .. .. .. ..
Drilaster of Ryukyu Archipelago by
.......someLuciolaspecies5(PA,OR,AU)
Kawashima et al., 2005). Characters for
10. Elytra broad, with lateral explanate margin of
these genera are provided herein based on
elytra quite broad in elytral base, unambig-
examinationofthetypespecies(*),together uously visible in humeral area dorsally;
with some additional exemplar species (**), usually at least two elytral costae well
onsomenontypespeciesonly(#),orinafew developed; hypomeron with frontal margin
casesonreferencesalone(1). at angle of 30–45 degrees with pronotal
1. Abdomenwithsixvisiblesternites(ventrites) 2 lateral margin laterally.. . .. .. .. .. .. 11
— Abdomenwithmore thansixventrites. . . 12 — Elytra more or less elongate, with lateral
2. Antennae serrate and depressed; coloration of explanate margin of elytra very narrow in
elytrapink tovariousdegrees.. .. .. . .. . humeral basal area, thus partially concealed
...........PristolycusGorham(OR,PA) by humerus dorsally; elytral costae usually
— Antennae filiform or somewhat moniliform; poorly developed, or weakly developed;
colorationof elytranever pink . .. .. . .. 3 hypomeron with frontal margin at angle of
3. Compound eyes with posterior margin clearly 70–90 degrees with pronotal lateral margin
emarginate4.. .. . .. .. Bourgeoisia Olivier laterally .. . Luciola LaportedeCastelnau
(OR) andLuciola cowleyi Blackburn(AU) (OR, PA, AU)andHotaria Yuasa6(PA)
— Compound eyes never or insignificantly emar- 11. Abdominaltergite 8 asymmetricbilaterally..
ginate posteriorly. .. .. .. .. .. .. .. . .. 4 . .. .. . .. .. .. .Luciola ovalisgroup (OR)
4. Elytron with superimposed costa arising from — Abdominaltergite 8 symmetricbilaterally. ..
humeral area and extending to near Atyphella (includings.g.Pygatyphella,AU)
apex. .. .. . CurtosMotschulsky (OR, PA) 12. Antennaebipectinate .. .. . .. .. .. .. .. ..
— Elytronwithoutclearhumeralcosta;ifpresent, . .. . CyphonocerusKiesenwetter (PA, OR)
not particularly more imposed than other — Antennaenot bipectinate . . .. .. .. .. .. 13
costae.. .. .. .. . .. .. .. .. .. .. .. . .. 5
5. Median apical area of abdominal tergite 8 5Including Luciola flavicollis Macleay, L. orapallida
Ballantyne, and L. nigra Olivier from Australia, L.
narrowed and deflexed, closely approaching
kuroiwae Matsumura from Ryukyu, L. trilucida from
the incurved and hooked apex of ventrite 6;
Taiwan, and some undetermined species fromIndochina
some species with acute and conspicuous
Peninsula.
anterior anglesonpronotum .. .. .. . .. . 6Hotaria was treated as a subgenus by McDermott
..............PygoluciolaWittmer(OR) (1966), or a synonym of Luciola by Kawashima et al.
— Median apical area of abdominal tergite 8 not (2003). Indeed, Hotaria and Luciola s.str. (based on L.
narrowed or deflexed; anterior angles of italica)sharedmanymorphologicalcharactersinmalesas
well as flightlessfemales which have complete elytra but
pronotum neveracute . .. .. .. .. .. . .. 6
absenceofhindwings.BallantyneandLambkin’s(2001,
2006)phylogeneticanalysessuggestacloserelationshipof
thesetwogroups.HoweverLuciolacurrentlyrecognizedis
4SomespeciesofLampyroideahavecompoundeyeswith ahighlydiversegroup,containingmorethan300species
an emarginate posterior margin in males, but the genus worldwide, and definitely needs a redefinition.
doesnotoccurintheareaandisthereforeexcludedfrom Accordinglywe have kept Hotariaas a valid name until
thekey. itsstatusisdecidedphylogenetically.