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Accepted by Q. Wang: 21 Mar. 2012; published: 1 Aug. 2012
ZOOTAXA
ISSN 1175-5326 (print edition)
ISSN 1175-5334 (online edition)
Copyright © 2012 · Magnolia Press
Zootaxa 3405: 1–34 (2012)
www.mapress.com/zootaxa/Article
1
The external larval morphology of aquatic and terrestrial Luciolinae fireflies
(Coleoptera: Lampyridae)
XINHUA FU1, LESLEY BALLANTYNE2,4 & CHRISTINE LAMBKIN3
1Hubei Insect Resources Utilization and Sustainable Pest Management Key Laboratory, College of Plant Science and Technology,
Huazhong Agricultural University, Wuhan 430070, Hubei, China. E-mail: fireflyfxh@mail.hzau.edu.cn
2School of Agricultural and Wine Sciences, Charles Sturt University, PO Box 588, Wagga Wagga 2678, Australia.
E-mail: lballantyne@csu.edu.au
3Queensland Museum, PO Box 3300 South Brisbane, 4101, Australia. E-mail: christine.lambkin@qm.qld.gov.au
4Corresponding author
Table of contents
Abstract . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
Material and methods . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
Results . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
Key to lucioline larvae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7
Aquatica ficta (Olivier) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
Aquatica hydrophila (Jeng et al.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Aquatica lateralis (Motsch.) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
Aquatica leii (Fu et Ballantyne) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
Aquatica wuhana Fu et Ballantyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14
Luciola cruciata Motsch. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
Luciola owadai Satô et Kimura . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .16
Luciola substriata Gorham . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Luciola aquatilis Thancharoen . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16
Semiaquatic larvae. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Pygoluciola qingyu Fu et Ballantyne . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Terrestrial larvae . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20
Asymmetricata circumdata (Motsch.). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22
Pteroptyx valida Olivier . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .24
Discussion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
Acknowledgements . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .31
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .31
Abstract
The external morphology of aquatic, semiaquatic and terrestrial lucioline larvae was investigated in order to provide an
overview of what traits constitute the extremely ecologically diverse Luciolinae (Coleoptera: Lampyridae). The aquatic
species, Aquatica ficta (Olivier), A. leii (Fu et Ballantyne), A. hydrophila (Jeng et al.), A. lateralis (Motschulsky), A. wu-
hana Fu et Ballantyne, Luciola cruciata Motschulsky and L. owadai Satô et Kimura cannot swim, but instead crawl on
the substrate. They have soft bodies, lateral abdominal tracheal gills and glands on eversible structures that secrete repel-
lent substances. The back-swimming species, Luciola substriata Gorham and L. aquatilis Thancharoen, which inhabit the
surface of ponds, have hardened exoskeletons, and lack gills and eversible glands. Unlike the crawling species, the back-
swimmers have sense organs along the ventral surface of the apical maxillary and labial palpomeres, and are metapneustic
in their later instars. The larval morphology of the aquatic species is contrasted with Pygoluciola qingyu Fu et Ballantyne,
whose larvae are semiaquatic and lack gills, and with the terrestrial larvae of Asymmetricata circumdata (Motsch.) (newly
FU ET AL.
2 · Zootaxa 3405 © 2012 Magnolia Press
associated here) and Pteroptyx valida Olivier. Keys to genera of the larval Luciolinae, to species in the genus Aquatica,
and to larval types based on habitat are provided. An overview of current knowledge of the lucioline larval biology and
taxonomy is given. Larval morphology provides a means to better understand the specialization of trophic behaviour in
aquatic and terrestrial habitats. Discussion covers use of larval characters in phylogenetic analyses, and attempts to deter-
mine habitat and mode of life preferences from morphological considerations.
Key words: Luciolinae larvae, Aquatica, Asymmetricata, Pteroptyx, Luciola, Pygoluciola, eversible glands, gills
Introduction
There is little knowledge about the biology and morphology of the immature stages of the Lampyridae in SE Asia
and it is often impossible to identify firefly larvae. Problems of identification are exacerbated by several factors not
peculiar to the SE Asian region. For example, correct identifications can only be established with reliable associa-
tions between adults and larvae. Adult-larval associations are established by rearing larvae until they metamor-
phose into adults, which is often a complicated undertaking (Archangelsky & Branham, 1998). Moreover, where
successful rearing confirms a larval-adult association, there is the further problem of verifying the identity of the
adult in the association. Lloyd and Walker (1967) stated that some biologists attempting to place a name on their
experimental material failed to properly identify the adult. These authors advocated the conventional use of
vouchered specimens to reduce the problem of erroneous or questionable firefly species identification. DNA
barcoding could serve to validate larval species, but until barcoding becomes cost effective, morphological charac-
ters of immature stages still remain convenient sources of species specific information.
Another problem relates to the evaluation of existing literature concerning larvae. Many publications address-
ing larvae provide such poor descriptions of external morphology, if any, that subsequent identification is virtually
impossible (Table 1). Few of these publications deal with larvae associated by rearing.
Lack of knowledge of larval morphology hinders phylogenetic analysis (Ballantyne & Lambkin 2000, 2001,
2006, 2009), and often prevents correct larval identification. The situation in the Luciolinae is however improving.
Ballantyne (1968) briefly described the larva of Bourgeoisia hypocrita Olivier but could not determine its mode of
life. Larvae of the “surf firefly” (Atyphella aphrogeneia Ballantyne & Buck) are apparently terrestrial while living
in the surf spray zone on coral reefs (Ballantyne & Buck 1979). A possible Atyphella guerini larva was associated
by location as only one firefly species has been recorded from New Ireland (Ballantyne 2000). Ballantyne and
Lambkin (2000) described larvae of five Atyphella and three Luciola species from Australia. In addressing the
larva of Pteroptyx valida Olivier, Ballantyne and Menayah (2002) overviewed literature regarding Luciolinae lar-
vae and developed a new and consistent terminology for the ventral plates of the larva, thus overcoming the confus-
ing and inconsistent previous terminology. Ho (2002, unpublished PhD) described aspects of the morphology of 21
species of Taiwanese firefly larvae, of which eight were Luciolinae. Jeng et al. (2003) overviewed aquatic species
and confirmed larval associations in five species from Taiwan. Chen (2003) depicted adults and larvae of several
species of Luciola and Curtos Motschulsky from Taiwan. Fu and Ballantyne (2006a, 2008) continued the review of
literature regarding lucioline larvae. Ho et al. (2009) addressed life history studies and some morphology of Luci-
ola ficta Olivier. Fu et al. (2010) erected a new genus of aquatic fireflies, in which all species have larvae reliably
associated. Ballantyne and Lambkin (2009) scored 27 species from larvae (Table 2), with only three species viz.
Magnalata limbata (Blanchard), Luciola cowleyi Blackburn and Bourgeoisia hypocrita with larvae associated on
the basis of label data; the remainder are reliable associations. However, few aquatic larvae were addressed in these
publications and none of these treatments included the use of a scanning electron microscope.
Here, we evaluate the external morphology of nine lucioline species with verified aquatic larval stages using
scanning electron microscopy for six of them. Aquatic larval morphology was compared with that of the terrestrial
larvae of Pteroptyx valida Olivier and Asymmetricata circumdata Motsch. and the semiaquatic larvae of Pygoluci-
ola qingyu Fu et Ballantyne using scanning electron microscopy. Present knowledge of the lucioline larvae was
reassessed in light of these findings.
Zootaxa 3405 © 2012 Magnolia Press · 3
FIREFLY LARVAE
TABLE 1. Records of lucioline larval associations in SE Asia and the Australopacific region. Extracted from Ballantyne and
Menayah (2002) and reassessed.
Footnotes: 1. Description of behaviour suggests back swimming larvae like Luciola substriata Gorham; no gills. 2. Aquatic
larva with single branch to gills. 3. Bertrand (1973:108 Figs 8–10) depicts unidentified aquatic larva with unbranched gills,
toothed mandibles and an elongate structure between the mandibles which probably represents hairs on maxillae and labium. 4.
Raj (1947) considered this larva might be Luciolinae of which only Luciola and Pyrophanes Olivier were known to him from
India; he identified it as Pyrophanes indica (Motsch.), the only Pyrophanes recorded from India. Olivier (1902a) incorrectly
assigned indica to Pyrophanes (Ballantyne 1993). 5. Morphology suggests Luciola praeusta Kiesenwetter. 6. The short
description indicates non-explanate tergal margins but further identification is impossible.
Material and methods
The following definitions of mode of life for larval Luciolinae are used:
Aquatic: living under water and with obvious adaptations for aquatic life (Jeng et al. 2003); either with lateral
abdominal gills or metapneustic, lacking gills.
Semi-aquatic: usually living in riparian environments and regularly entering the water in search of prey; lacking
obvious adaptations to aquatic life, such as gills (Fu & Ballantyne 2008).
Terrestrial: lacking gills and any specialized respiratory and behavioural modifications for an aquatic life.
The terminology of the ventral plates follows Ballantyne and Menayah (2002).
Pygopodial structure was interpreted using Ho et al. (2000).
Rearing: First instar larvae of Luciola substriata were bred from eggs which females laid on the back of giant
duckweed Spirodela polyrhiza (Fu et. al. 2005a). Other aquatic larvae, such as Aquatica ficta, Aq. lateralis, Aq.
wuhana and Luciola cruciata were bred in tap water following the method outlined in Fu et. al. (2006b, c). Larvae
Unsubstantiated records/
Doubtful identification Locality Mode of life References
? Luciola vespertina F. India aquatic Annandale 1906
Unidentified 1Siam
Malaysia aquatic
aquatic Annandale 1900
Annandale 1900
? Pyrophanes similis Olivier 2 Celebes aquatic Blair 1927
Bertrand 1972, 73
Luciola species3 Ceylon aquatic Bertrand 1972, 73
? Luciola picea Gorham India ? Olivier 1900
Pyrophanes indica Motsch.4India terrestrial Raj 1947
Luciola gorhami Ritsema/L. dubia Olivier5India terrestrial Fletcher 1919
Mehta 1932
Gardner 1946
Identification reassessed Locality Mode of life References
Described as Luciola australis F.; probably
Atyphella guerini Ballantyne New Ireland terrestrial Olivier 1883
Identification reassessed in Ballantyne 2000
Described as Luciola flavicollis Macleay,
probably Luciola nigra Olivier Australia terrestrial Armitage 1908
Identification reassessed in Ballantyne &
Lambkin 2000
Bred to adult
(inadequate morphology) Locality Mode of life References
Colophotia brevis Olivier6Indonesia terrestrial Blair 1927
No adults
Luciola trivandrensis Raj India Raj 1941
Described from larva only; adults not associ-
ated
FU ET AL.
4 · Zootaxa 3405 © 2012 Magnolia Press
of Asymmetricata circumdata were bred from eggs. Five males and three females were collected in Xishuangbanna
Tropical Botanical Garden (Jinghong City, Yunnan Province, China) on April 28, 2011. Males and females were
put in a round transparent plastic box (20 cm diameter, 6 cm high) and allowed to mate. Moist filter papers were put
in the box. After females laid eggs on the papers, males and females were killed fixed and preserved by immersion
in 80% ethanol. Eggs were transferred back to the same box and hatched at 25 °C and natural photoperiod. Tap
water was sprayed on the eggs every day. Neonate larvae were transferred to and reared in the same boxes at 25 °C
and natural photoperiod with crushed land snails Bradybaena ravida sieboldiana Pfeiffer 1850 as food. Tap water
was sprayed every two days. Thancharoen provided specimens of Luciola aquatilis Thancharoen she had bred.
SEM: The following larvae were used: Aquatica ficta, five 6th instars; Aq. lateralis, three 5th instars; Aq. leii,
three 5th instars; Aq. wuhana three 4th instars; Luciola cruciata, three 5th instars; L. substriata, two 1st instars, three
6th instars; Pygoluciola qingyu, three 5th instars; Asymmetricata circumdata, two 6th instars; Pteroptyx valida, two
4th instars. The collection locality and details, and deposition of specimens are included under the material
examined for each species.
Larvae were killed in liquid nitrogen. The heads, tracheal gills of six species and pygopodia of three species
were dissected and fixed in a solution of 2% glutaraldehyde and 2.5% paraformaldehyde in a 0.1 M sodium phos-
phate buffer, pH 7.4. The samples were washed with 0.1 sodium phosphate, pH 7.4, and fixed in 2% osmium
tetraoxide in a 0.1 M pH 7.4 phosphate buffer for 2 h at room temperature. They were then dehydrated in a
30–100% ethanol gradient for 15 min at each concentration. This was followed by three changes in terbutilic alco-
hol and the samples were dried by sublimation in a VFD-20 desiccator (Vacuum freezing drier) for 1 h. Once dried,
samples were placed on aluminium bases with double-sided carbon adhesive tape and sputter coated with gold (Fu
et. al. 2007; Fu & Ballantyne 2008; Fu et. al. 2009; Fu et. al. 2010). The specimens were observed under a JSM-
6390 LV scanning electron microscope at an accelerating voltage of 25 kV.
Images taken using the SEM include Figs 1 to 60.
Unless indicated otherwise, specimens are deposited in the Natural History Museum, Huazhong Agricultural
University (NHMHAU).
Abbreviations for figures: A1, 1st antennal segment; A2, 2nd antennal segment; A3, 3rd antennal segment; EO,
eversible organ; G, galea; LP, labial palpus; M, mandible; MXP, maxillary palpus; S, spiracle; SC, sensory cone;
TG, tracheal gill.
Drawing: Fu traced the outline of Figs 91–93 from SEM images.
Photography: A Dino-lite digital microscope pro was used for Figs 67–72. A Nikon D700, 105mm macro
lens, macro flash plus an extension tube were used for Figs 73–78, 79–84, 85–90.
Results
General features of lucioline larvae (modified and expanded from Ballantyne & Menayah 2002)
Body: (Figs 73, 75, 77, 79, 81, 83, 85, 87, 89). Elongate, slender, usually tapering somewhat in front and
behind; with 3 thoracic and 9 abdominal segments; terga usually more heavily sclerotized than underside of body,
appearing little sclerotised in Curtos spp.; in aquatic species with gills, dorsal and ventral plates may be separated
by wide areas of membrane (Figs 73–82). In Pygoluciola qingyu, and probably larvae of Luciola chinensis (L.), L.
praeusta Kiesenwetter, L. terminalis Olivier, and L. anceyi Olivier the external dorsal plates are well sclerotised
with no areas of exposed membrane; the ventral surface retains flexibility because of intersegmental membranes.
Luciola substriata Gorham and L. aquatilis Thancharoen (Figs 83–86) have dorsal and ventral surfaces bearing
heavily sclerotised plates with no membranous areas visible. All body segments except the last with a dorsal
medial longitudinal line. On the ventral surface an elongate pleural suture runs from the anterior margin of segment
2 to the posterior margin of abdominal segment 9, delimiting median sternal elements ventrally, and laterotergites
dorsally; laterotergites bear spiracles in the thorax, and either spiracles or gills in the abdomen. In L. substriata and
L. aquatilis laterosternites are lacking in the abdomen.
Head: (Figs 1–6, 12–30, 34–60) invisible beneath prothoracic tergum if retracted, concealed at rest within pro-
thorax; extensible neck membrane forming two layered envelope around retracted head; prognathous, well sclero-
tized, small, flattened, parallel sided; ocellus on each side; head capsule divisible into median dorsal frontoclypeus,
bounded laterally and posteriorly by frontal arms of epicranial suture; lateral epicranial halves reflexed ventrally
Zootaxa 3405 © 2012 Magnolia Press · 5
FIREFLY LARVAE
but not meeting. Antennae: (Figs 4, 17, 23, 27, 34, 35, 41, 47, 48, 52, 59) 3 segmented, with elongated scape and
pedicel, and apical very short (3rd) segment (the flagellum), lying beside small sense cone; elongate "articulating
membrane" forms two layered envelope around retracted antenna. Mouthparts: (Figs 1–3, 5, 6, 13–16, 18, 19–30,
34–36, 46, 49–51, 53–58, 60) well developed; mandibles falcate, strongly sclerotized, densely covered in fine hair
along outer margins, perforated along length by a fine canal that opens on the outer margin just behind the apex;
with or without a basal retinaculum; retinaculum having 1 or 2 teeth; maxillae and labium fused forming compound
plate covering most of ventral head area. Maxillae with short, squat, four segmented palp; basal segment large and
well defined, segments 2 and 3 very short and diminishing in width towards apex, apical segment longer and nar-
rower than palpomere 3, shape differs between genera and species; palp may obscure galea, which is long, thin,
two segmented, bearing long and short setae at apex, and on its inner margin an elongate, flattened, dense profusion
of anteromedially directed hairs (lacinia); cardo well sclerotised, articulating with broad elongate stipes. Labial
palpi small, two segmented, arising at anterolateral corners of small prementum which lacks a ligula; postmentum
elongate, not well sclerotised and colourless, and joined along sides by membrane to the cardines.
Thorax: prothorax usually longer than wide, anterior margin often bluntly rounded, and containing retracted
head within (e.g. Fig. 34); lateral margins slightly divergent posteriorly; ventral surface little differentiated, with
very narrow strips of cuticle dorsal and lateral to coxae 1 representing prothoracic episterna and epimera; meso and
metathoracic segments shorter than prothorax and often rectangular in outline; ventral surface of meso and
metathorax (Figs 74, 76, 78, 80, 82, 84, 86, 88, 90) with median sternal elements delimited by an elongate pleural
suture from the laterotergites; ventral surface composed of two areas in each segment: 1. an anterior presternum
with paired laterotergites bearing well developed (biforous) spiracles in the mesothorax; spiracles may be rudimen-
tary in the metathorax (Thancharoen et al. (2007) indicated variability of spiracle number and functionality in the
thorax of certain Lampyridae larvae); 2. a median subrectangular sternal area bearing the legs, above the coxae of
which the meso- and metathoracic episterna and epimera are visible as thin sclerotised strands, and margined later-
ally by paired laterotergites. Dorsal and ventral plates often widely separated in aquatic species by expanded mem-
branous areas. In predacious species (e.g. Pygoluciola qingyu) dorsal plates hardened, darkened, ventral surface
retaining flexibility because of pale intersegmental membranes. In Luciola substriata (Fig. 84) a set of hardened
plates anterior to coxae of segments 2 and 3 (Jeng et al. 2003; Fig. 40) may be a secondary development to provide
flexibility. Legs: (Figs 74, 76, 78, 80, 82, 84, 86, 88, 90) five segmented; short, cylindrical coxae widely separated
at their bases, with apices inclining medially; trochanters elongate, joining femora obliquely; tibiae covered with
short strong setae, terminating in a single apical claw - the tarsungulus; legs 1 are shorter than legs 2 and 3.
Abdomen: terga 3–9 often subequal in length, usually tapering in width from tergum 6 backwards; in terrestrial
species (Figs 88, 90) segments 1–8 have single laterotergites at each side with sclerotised plates bearing spiracles;
in some aquatics (Figs 73–82) laterotergites bear forked gills and eversible organs (Figs 7–12) arising from the
membranous area above the gill bearing plates; ventral area of segments 1–8 with median subrectangular, often
pigmented, sternal area (ventral plates very lightly coloured in gilled aquatics Figs 74, 76, 78, 80, 82); median ster-
nal area margined by elongate, narrow, paired, sometimes pigmented laterosternites, delimited by folds from lat-
erotergites above and median sternal plate below (e.g. Figs 88, 90); Luciola substriata and L. aquatilis lacking
laterosternites in abdomen and ventral body area well sclerotised and darkened (Figs 84, 86); ventral area of seg-
ment 9 simple plate with no differentiation into areas; light organ present beneath antepenultimate abdominal seg-
ment (8); Ohba and Sim (1994) described it as paired; abdomen terminated by a series of eversible filaments
(pygopods) bearing recurved hooks (Figs 61–66, 91–93). Segment 10 may be the thin ring of cuticle at the poste-
rior end of segment 9.
Generic characteristics of lucioline larvae
While ten of the 19 lucioline genera recognised here have larvae and adults associated by rearing, in several
instances the record is for one or two species only (Table 2; e.g. Pteroptyx, Colophotia Dejean, Asymmetricata Bal-
lantyne and Lloydiella Ballantyne; Ballantyne & Lambkin 2009). The following larval associations are tentative:
Aquilonia costata, Magnalata limbata, Bourgeoisia hypocrita (label data, similarity of larval data to adults, and the
only firefly species known from Fiji, respectively).
FU ET AL.
6 · Zootaxa 3405 © 2012 Magnolia Press
TABLE 2. Characteristics of lucioline larvae. Arrangement of genera follows Ballantyne & Lambkin (2009) and Fu et al.
(2012).
Genus/Species
Habitat1
Ter ga l
margins2
Gills
Meta-pneus-
tic
Glands on
eversible
structures
References
# *Aquatica Fu Ballantyne &
Lambkin
- 5 species
ANE1 +-+ Fu et al. 2010
Aquilonia Ballantyne T E1 - - - Fu et al. 2012
*Asymmetricata Ballantyne
- circumdata (Motsch.) T E1 - - - Ballantyne & Lambkin 2009
# *Atyphella Olliff
-7 Australian species T E1 - - - Ballantyne & Lambkin, 2000, 2009
#* Bourgeoisia Olivier
- hypocrita (Olivier) ? E2 - - - Ballantyne 1968; Ballantyne & Lamb-
kin, 2000, 2009; Deheyn & Ballantyne
2009 ; Fu et al. 2012
*Colophotia Dejean
- praeusta Kiesenwetter
- brevis Olivier
TNE1 --- Ballantyne & Lambkin 2009
Blair 1927 (see Table 1)
Convexa Ballantyne ? ? ? ? ? Larva unknown
Ballantyne & Lambkin 2009
*Curtos Motsch.
-costipennis
(Gorham)
-okinawanus Matsumura
- sauteri Olivier
- fulvocapitalis Jeng et Satô
T
T
T
T
NE1
NE1
NE1
NE1
-
-
-
-
-
-
-
-
-
-
?
?
Ballantyne & Lambkin 2009; Jeng et
al. 1998; Chen 2003; Ho 2002.
Emeia Fu et al.
-pseudosauteri (Geisthardt). T NE1 in thorax,
NE2 in abdomen - - - Geisthardt (2004) ; Fu et al. 2012
Gilvainsula Ballantyne ? ? ? ? ? Larva unknown
Ballantyne & Lambkin 2009
Lampyroidea Costa ? ? ? ? ? Larva unknown
Ballantyne & Lambkin 2009
# *Lloydiella Ballantyne
- majuscula (Lea) T E1 - - - Ballantyne & Lambkin 2000, 2009
# *Luciola Laporte
-anceyi Olivier3
-aquatilis Thancharoen
-australis (F.)
-cerata Olivier3
-cruciata Motsch.
-filiformis Olivier
-flavicollis MacLeay
-gorhami Ritsema
-kagiana Matsumura
-nigra Olivier
T
A
T
T
A
T
T
T
T
T
NE2
NE2
NE1
NE2
NE1
E1
NE1
NE2
E1
NE1
-
-
-
-
+
-
-
-
-
-
-
+
-
-
-
-
-
-
-
-
?
-
-
-
+
-
-
-
-
-
Ho 2002; Chen 2003
Ballantyne & Lambkin 2009;
Thancharoen et al. 2007
Ballantyne 1988; Ballantyne & Lamb-
kin 2009
Ho 2002; Chen 2003
Ballantyne & Lambkin 2009; McDer-
mott, 1966
Okada 1928; Minami 1966; Ohba et
al. 1994
Chen 2003
Ballantyne & Lambkin 2000, 2009
Ho 2002; McDermott 1966
Chen 2003
Ballantyne & Lambkin 2000, 2009
continued next page
Zootaxa 3405 © 2012 Magnolia Press · 7
FIREFLY LARVAE
Footnotes: 1. A aquatic; T terrestrial; SA semiaquatic. 2. E Explanate (E1 widely explanate, margins thin; E2 explanate poster-
olateral corners; margins thick); NE not explanate (NE1 laterotergites usually visible at sides of body; NE2 laterotergites not
visible at sides of body). 3. Luciola cerata, praeusta, terminalis and anceyi have similar larval types to P. q ingy u and L. cow-
leyi. 4. # Larval specimens described in literature listed. *Scored in Ballantyne & Lambkin 2009.
Key to lucioline larvae
1. Aquatic (metapneustic or with gills along the sides of the abdomen) (Figs 73–86). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .2
- Terrestrial/semiaquatic (lacking gills) (Figs 87–90) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .9
2. Metapneustic in later instars; hard bodied with no obvious membranous areas; back swimmers just below water surface . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Luciola substriata Gorham /L. aquatilis Thancharoen (Figs 83, 84)
- With gills arising from the sides of abdominal segments 1–8 (Figs 73–86). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .3
3. Protergum lacking marginal pale markings. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Aq. hydrophila (Jeng et al.)
-Protergum always with some pale lateral margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .4
4. Protergum with 2 pale markings only (at anterolateral corners) (Fig. 77) . . . . . . . . . . . . . . . . . . . . . .Aq. leii (Fu et Ballantyne)
TABLE 2. (continued)
Genus/Species
Habitat1
Ter gal
margins2
Gills
Meta-pneus-
tic
Glands on
eversible
structures
References
-orapallida Ballantyne
*- owadai Matsumura
-*parvula Kiesenw. 4
-substriata Gorham
T
A
T
A
NE1
NE1
E1
NE
-
+
-
-
-
-
-
+
-
?
?
?
Ballantyne & Lambkin 2000, 2009
Ballantyne & Lambkin 2009; McDer-
mott, 1966
Okada 1928; Minami 1966; Ohba et
al. 1994
Chen 2003
Ballantyne & Lambkin 2009; Than-
charoen et al. 2007; Fu et al. 2005a, b
*Magnalata
-limbata (Blanchard) T E1 - - - Ballantyne & Lambkin 2009
Missimia Ballantyne ? ? ? ? ? Larva unknown Ballantyne & Lamb-
kin 2009
Photuroluciola Pic ? ? ? ? ? Larva unknown Ballantyne & Lamb-
kin 2009
# Pteroptyx
-maipo Ballantyne
malaccae Gorham
-valida Olivier
T
?SA
T
NE1
NE1
NE1
-
-
-
-
-
-
-
-
-
Ballantyne et al. 2011
Ballantyne & McLean 1970;
Ballantyne, 2002; Lomboot et al.
2007
Ballantyne & Menayah 2002;
Ohba & Sim 1994; Ballantyne &
McLean 1970;
Lomboot et al. 2007; Ballantyne et
al. 2011.
Pygatyphella Ballantyne ? ? ? ? ? Larva unknown
Ballantyne & Lambkin 2009
#* Pygoluciola Wittmer
-qingyu Fu et Ballantyne
- cowleyi 3 (Blackburn) SA
?T
NE2
NE2
-
-
-
-
-
-
Fu & Ballantyne 2008
Ballantyne & Lambkin 2009
Pyrophanes Olivier ? ? ? ? ? Larva not reliably associated;
Ballantyne & Lambkin 2009;
(Table 1)
FU ET AL.
8 · Zootaxa 3405 © 2012 Magnolia Press
- Protergum with more than 2 pale markings around margins . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .5
5. Protergum sub-parallelsided; with four separate pale marks at anterolateral corners and along lateral margin anterior to the pos-
terolateral corners (Fig. 75). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Aq. lateralis (Motsch.)
- Protergum with lateral margins not sub-parallelsided; pale markings not as above . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .6
6. Median dark marking on protergum prolonged narrowly anteriorly and posteriorly to meet anterior and posterior margins (Fig.
81) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Luciola cruciata Motsch.
- Median dark marking on protergum not prolonged narrowly anteriorly, median dark area may assume the shape of a cross with
wide anterior and lateral arms, thus protergum having four discrete pale markings at anterolateral and posterolateral corners. 7
7. Protergum mainly dark marked, with four discrete pale areas at anterolateral . and posterolateral corners; no pale markings on
remaining terga; median line narrow (Fig. 73) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . ..Aq. ficta (Olivier)
- Protergum either marked as above or with pale areas extending obliquely across posterolateral corners to almost meet in
median line; always with pale markings on remaining terga; median line narrow or wide . . . . . . . . . . . . . . . . . . . . . . . . . . . . .8
8. Protergum mainly dark, with four discrete pale areas at anterolateral and posterolateral corners; terga of body segments 2–11
very small and separated by wide median line; small pale markings at posterolateral corners of terga of body segments 2–11;
terga of body segments 10, 11 lacking pale lateral margins (Ohba et al. 1994: 17, Fig.1). . . . . . . . . ..L. owadai Satô et Kimura
- Protergum mainly dark with extensive pale markings obliquely across posterolateral corners; terga of body segments 2–11
large, not widely separated by narrow median line; extensive pale markings across most of posterior margins of body segments
2–8 or 9; lateral margins of terga of body segments 10, 11 pale (Fig. 79) . . . . . . . . . . . . . . . . . .Aq. wuhana (Fu et Ballantyne)
9. Semiaquatic – living near shallow bodies of water and able to enter them and submerge for long periods; lacking gills; dorsal
body plates very heavily sclerotised and lacking membranous areas; mandibles with 2 inner teeth (Fu & Ballantyne 2008; Figs
2–4, 23–25) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .Pygoluciola qingyu Fu et Ballantyne
- Dorsal body plates usually dark coloured, may not be well sclerotised; mandibles with single tooth, or not toothed; probably
terrestrial . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .10
10. Mode of life not determined, probably terrestrial; lacking gills; dorsal body plates very heavily sclerotised and lacking mem-
branous areas; mandibles with a single tooth . . . . . . . . . . . . . . . . . . . . . . . .Luciola cerata, praeusta, terminalis, Pygo. cowleyi
- Terrestrial; lacking gills; dorsal body plates not usually very heavily sclerotised; mandibles not toothed . . . . . . . . . . . . . . . 11
11. Tergal margins not laterally explanate; laterotergites often visible at sides of body when specimen viewed from above.(e.g.
Figs 89, 90) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .12
- Tergal margins laterally explanate, usually along lateral margins, sometimes only at posterolateral corners (e.g. Figs 87, 88)13
12. Dorsal body plates usually dark coloured and reaching to sides of terga . . . . . . . . .Pteroptyx and Colophotia spp. (e.g. Fig. 89)
- Dorsal body entirely pale, tergal plates pale coloured and not reaching to sides of terga . . . . . . . . . . . . . . . . . . . . . .Curtos spp.
13. Thoracic and/or abdominal terga prolonged at posterolateral corners . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .14
- All tergal lateral margins widely explanate, not prolonged, except for terminal tergum . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
.Magnalata sp., Atyphella spp., Luciola parvula, L. filiformis, L. kagiana, Lloydiella sp., Asymmetricata circumdata (Figs 87,
88)
14. Lateral margins of thoracic terga very broad, not prolonged except for very short pointed posterolateral corners on terga 2, 3;
lateral margins of abdominal terga 1–8 narrowly prolonged (Fu et al. 2012 Figs 5, 6) . . . . . . . . . . . . . . . . Emeia pseudosauteri
- Lateral margins of both thoracic and abdominal terga narrowly prolonged (Fu et al. 2012 Figs 50, 51) . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bourgeoisia hypocrita
Aquatic larvae
Aquatic larvae are of two types:
1. Mode of life: having tracheal gills and spiracles; usually living in shallow well oxygenated water, cannot swim
(Figs 73–82).
Body: (Figs 73–82) protergum lacking median line; body soft with extensive membranous areas, tergal plates
not explanate, and often widely separated in median line; laterotergites visible in thorax and abdomen; plates on
ventral body surface usually not coloured, very flexible; laterotergites and laterosternites discernable in abdomen.
Fork shaped tracheal gills (Fig. 7) with a non-functional spiracle occurring at the tip of the shorter branch at sides
of first 8 abdominal segments. Several species have 10 pairs of eversible gland bearing sacs (Figs 7–12) arising at
the sides of the meso and metathorax and abdominal segments 1–8 (Fu et al. 2007, 2009; Fig. 7); glands bearing
from 2−7 spines (Fu et al. 2007, 2009; Figs 9–12). Pygopodia of Aq. leii consisting of 6 main stalks branching at
their apices into two; concentric rings of recurved hooks surrounding the entire expanded stalks (Figs 61, 62, 91).
Head: Antennal segment 3 elongate, surmounted by closely adpressed short finger like projections, subequal in
length and width to adjacent sense cone; apex of segment 2 not strongly oblique (Figs 3, 4, 15, 17, 21, 23, 27).
Mouthparts: Mandibles: (Figs 2, 15, 21, 28, 67–69) lacking retinaculum, not much expanded at base; inner
edge of apical half with row of very short setae (edge appears serrate); most of dorsal surface irregular and resem-
Zootaxa 3405 © 2012 Magnolia Press · 9
FIREFLY LARVAE
bling overlapping scales; single line of thick stout setae bordering lateral margin of scaled area on dorsal surface,
setae often split at apices; outer edge of mandibles bearing hairs in preapical area only. Maxillae (Figs 1, 5, 6, 13,
14, 18, 19, 20, 24, 25, 26, 29, 30): apical palpomere with terminal sense organs. Labium (Figs 1, 3, 13, 16, 19, 22,
25, 29): palpi close at their bases, apical palpomere with terminal sense organs; ventral surface of prementum with
hairless “keel” at least 3 times as long as wide.
2. Mode of life: Metapneustic (Figs 83–86), lacking gills in later instars; lateral bristles in first and possibly second
instars may function as gills (Fu & Ballantyne 2009; Figs 31–33); back swimmers common at or just beneath water
surface with their terminal spiracles piercing the surface (Fu et al. 2005a).
Body (Figs 83, 85): protergum with fine median line; dorsally flattened; well sclerotised plates covering dorsal
and ventral body surfaces (no exposed membranous areas), tergal plates not laterally explanate, laterotergites not
visible from above at sides of abdomen; ventral plates of abdomen heavily sclerotised, dark, consisting of median
sternal plate and lateral paired laterotergites carrying spiracles; abdomen lacking laterosternites; spiracles arising
from posterolateral areas of the dorsal surface of abdominal segment 8; apparently lacking defensive glands (Fu et
al. 2010; Figs 84, 86).
Head: antennal segment 3 elongate, surmounted by closely adpressed short finger like projections, subequal in
length and width to adjacent sense cone; apex of segment 2 not strongly oblique (Figs 35, 41, 83–86).
Mouthparts: Mandibles: (Figs 39, 70) with narrow retinaculum, bearing a dense profusion of curved setae;
inner edge of apical half lacking row of very short setae; most of dorsal surface irregular and resembling overlap-
ping scales; single line of thick stout setae bordering lateral margin of scaled area on dorsal surface, setae often
split at apices; outer edge of mandibles bearing hairs along most of length. Maxillae: apical palpomere with sense
organs along ventral surface (Fig. 42). Labium: palpi moderately separated at their bases, apical palpomere with
sense organs along ventral surface; ventral surface of heart shaped prementum with hairless “keel” about as long as
wide (Fig. 40).
Aquatica ficta (Olivier)
Figs 1–12, 67, 73, 74
Luciola ficta Olivier, 1909c:249. Pic, 1911:188. McDermott, 1966:104. Ho & Jiang, 1997:42. Yeh, 1999:1. Jeng et al.
2003:545. Ho, 2002:48. Chen, 2003:163. Ho et al, 2009:100.
Luciola ovalis Hope. Matsumura 1918:84 misidentification. Chang 1994:1 misidentification.
Aquatica ficta (Olivier). Fu et al. 2010:6.
Material examined. CHINA: 20 sixth instars bred from an original population of five males and three females
collected from Fuqing County, Fuzhou City, Fujian Province, May 15, 2008 (NHMHAU).
Diagnosis. Protergum having rounded sides; with four pale spots at anterolateral and posterolateral corners;
remaining terga lacking pale markings; thoracic and abdominal terga not widely separated, median line narrow;
abdominal terga 1–8 with plates obliquely emarginate in anterolateral corners (Figs 73–74); abdominal glands with
3–7 spines (Figs 7–12); antennal segment 3 with closely adpressed finger like projections subequal in length to
adjacent sense cone (Figs 1–4); mandibles lacking retinaculum (Fig. 67); palpi with terminal sense organs (Figs 5,
6).
Aquatica hydrophila (Jeng et al.)
Luciola hydrophila Jeng et al., 2003:544 (Fig. 39).
Aquatica hydrophila (Jeng et al.).Fu et al., 2010:6.
Diagnosis. Protergum having rounded sides, lacking pale markings (Jeng et al. 2003; Fig. 39); thoracic terga 2, 3
and abdominal terga 1–8 with wide pale median line, otherwise lacking any pale markings, plates obliquely emar-
ginate in anterolateral corners. Pygopodia with basal row having two retractable filaments.
FU ET AL.
10 · Zootaxa 3405 © 2012 Magnolia Press
Remarks. Larvae not examined by the authors herein. Jeng et al. (2003) briefly described the larva which they
considered approached that of L. owadai; differentiated on the basis of dorsal colour patterns and pygopod struc-
ture.
FIGURES 1–6. Aquatica ficta larval head, 6th instar, SEM. 1 anterior, dorsal surface uppermost; 2 dorsal anterior half of head;
3 ventral; 4 right antenna; 5–6 maxillary palpus; 6 detail apical sensory area.
Zootaxa 3405 © 2012 Magnolia Press · 11
FIREFLY LARVAE
FIGURES 7–12. Aq. ficta larva, SEM. 7 tracheal gills and eversible organs, first and second abdominal segments from above;
8 non-functional spiracle at base of tracheal gill; 9–12 various protuberances on surface of eversible organs.
Aquatica lateralis (Motsch.)
Figs 13–18, 68, 75, 76
Luciola lateralis Motschulsky, 1860:144. Olivier 1902a:82; 1907:53. Okada, 1928:101. Minami, 1966:73; McDermott,
1966:108. Bertrand, 1972:601; 1973:107. Ohba, 1986:156; 1988:12. Ohba et al. 2001:91. Jeng et al. 2003:546. Suzuki et
al., 2004:287. Fu et al. 2009:155.
Luciola viticollis Kiesenwetter 1874:261. Gorham, 1883:409. Olivier, 1902a:77; 1902b:189. Okada, 1931:146. McDermott, 1966:102.
Aquatica lateralis (Motsch.) Fu et al., 2010:1.
FU ET AL.
12 · Zootaxa 3405 © 2012 Magnolia Press
FIGURES 13–18. Aq. lateralis larval head, 5th instar, SEM. 13 anterior, dorsal surface uppermost; 14 apex of galea; 15 dorsal;
16 ventral; 17 right maxillary palpus and antenna; 18 tip of sensory area maxillary palpus.
Material examined. JAPAN: Five fifth instars, bred in lab, population collected from Tokyo Metropolitan
University, Minami–Ohsawa, Hachioji, Tokyo, May 28, 2008 (NHMHAU).
Diagnosis. Protergum with parallel sides; four pale areas, two at anterolateral corners, and two along lateral
margin just anterior to the posterolateral corners; median line wide, reaching maximum width at anterior margin of
thoracic tergum 3; thoracic terga 2, 3 with midlateral paler markings Figs 75, 76); abdominal glands with 4–7
spines including a median spine (Fu et al. 2009; Fig. 3); antennal segment 3 with closely adpressed finger like
Zootaxa 3405 © 2012 Magnolia Press · 13
FIREFLY LARVAE
projections subequal in length to adjacent sense cone (Figs 13–17); mandibles lacking retinaculum (Fig. 68); palpi
with terminal sense organs (Figs 13, 18).
Remarks. Okada (1928) first reported 10 pairs of bifid defensive organs along the sides of the body, and indi-
cated an odour of resin and peppermint in both this species and Luciola cruciata. His figure 1b (page 102) of Luci-
ola lateralis shows gills protruding from the sides with defensive organs on the right side of thoracic segment 3 and
on either side of abdominal segment 3. Figure 75 herein indicates only the dorsal branch of the gills, while Fig. 76
shows both the dorsal branch and a short ventral branch on abdominal segments 3-5.
FIGURES 19–24. Aq. leii larval head, 5th instar, SEM. 19–20 anterior dorsal surface uppermost; 21 dorsal; 22 ventral; 23 right
antenna; 24 tip of maxillary palpus.
FU ET AL.
14 · Zootaxa 3405 © 2012 Magnolia Press
Aquatica leii (Fu et Ballantyne)
Figs 19–24, 77, 78, 91
Luciola leii Fu & Ballantyne, 2006a:339. Fu et al. 2006b:860; 2007:117; 2009:155.
Material examined. CHINA: Sixteen fifth instars bred from an original population of nine males and four females
collected from Huazhong Agricultural University, Wuhan City, Hubei Province, May 14, 2010 (NHMHAU).
Diagnosis. Protergum with two small pale areas at anterolateral corners; remaining terga lacking pale mar-
kings (paler or darker markings in this species are in the membranous areas not the dorsal tergal plates); lateral
margins of protergum rounded; median line narrow (Figs 77, 78); glands with 2–6 marginal spines, lacking median
spine (Fu et al. 2009; Fig 1A); pygopodia arising from 6 basal stalks; rings of recurved hooks completely surroun-
ding each stalk (Figs 61, 62, 91); antennal segment 3 with closely adpressed finger like projections subequal in
length to adjacent sense cone (Figs 19, 21–23); mandibles lacking retinaculum; palpi with terminal sense organs
(Figs 20, 22, 24).
Remarks. The dorsal colouration of the larva depicted in Fu and Ballantyne (2006a; Fig. 8) differs from that
depicted here in having much larger anterolateral pale areas on the protergum, and pale margins on the terminal ter-
gum (Fig. 77).
Aquatica wuhana Fu et Ballantyne
Figs 79, 80
Aquatica wuhana Fu et Ballantyne 2010:1.
Material examined. CHINA: Three fourth instars bred from an original population of five males and two females
collected from E Jia Bian Village, Jiang Xia District, Wuhan City, Hubei Province, May 28, 2008 (NHMHAU).
Diagnosis. Protergum with extensive pale areas at anterolateral, and obliquely across posterolateral corners,
extending medially almost meeting across posterior margin; each set of dorsal plates of thoracic terga 2, 3 and
abdominal terga 1–3 with pale posterolateral corners, pale area extending narrowly across posterior margin to
expand slightly at inner area; abdominal terga 4–6 with pale markings restricted to posterolateral corners and to one
side of median line; abdominal terga 8, 9 pale along lateral margins; narrow median line extending from anterior
margin of thoracic tergum 2 to posterior margin of abdominal tergum 8 (Figs 79–80); abdominal glands with 3–7
spines (Fu et al. 2010; Figs 48, 49); antennal segment 3 with closely adpressed finger like projections subequal in
length to adjacent sense cone; mandibles lacking retinaculum; palpi with terminal sense organs (Fu et al. 2010;
Figs 44–47).
Luciola cruciata Motsch.
Figs 25–30, 69, 81, 82
Luciola cruciata Motschulsky, 1854:53. For economy of scale the references in McDermott, 1966:102 are not repeated here.
Bertrand, 1972:599, 601; 1973 Fig. 7. Ohba, 2001:45. Fu et al. 2009:155.
Material examined. JAPAN: Five fifth instars bred from an original population of three males and a female
collected from Tokyo Metropolitan University, Minami-Ohsawa, Hachioji, Tokyo, June 5, 2008 (NHMHAU).
Diagnosis. Similar to larvae of Aquatica spp., distinguished by the dorsal colour patterns, especially that of the
protergum; protergum with extensive median dark area narrowly prolonged in the middle along anterior and poste-
rior margins; dark area may extend to sides; most of posterior half of thoracic terga obliquely pale across corners
and narrowly across posterior margin; pale areas across posterior margins of most abdominal terga restricted to 2
separate areas with the outer area larger; median line wide (maximum width across abdominal terga 3, 4), and terga
widely separated (Figs 81, 82; Okada 1928, Fig 1a, Plate VIII E); eversible organs when everted with anterior and
posterior branches longer and thinner than those of Aq. leii or Aq. lateralis (Fu et al. 2009 Table 1); glands with 6
or more spines including 2 median spines; antennal segment 3 with closely adpressed finger like projections
Zootaxa 3405 © 2012 Magnolia Press · 15
FIREFLY LARVAE
subequal in length to adjacent sense cone (Figs 25, 27, 28); mandibles lacking retinaculum (Fig. 69); palpi with
terminal sense organs (Figs 25, 26, 29, 30).
Remarks. Okada (1928: Fig. 1a, page 102) depicts the everted defensive organs above the lateral gills. Figures
81 and 82 herein do not show everted abdominal glands.
FIGURES 25–30. Luciola cruciata larval head, 5th instar, SEM. 25–26 anterior, dorsal surface uppermost; 27 right antenna; 28
dorsal; 29 ventral; 30 apical maxillary palpomere.
FU ET AL.
16 · Zootaxa 3405 © 2012 Magnolia Press
Luciola owadai Satô et Kimura
Luciola owadai Satô & Kimura, 1994: 159. Ohba et al., 1994: 13.
Diagnosis. Similar to larvae of Aquatica spp., distinguished only by the dorsal colour patterns, especially that of
the protergum; protergum with extensive median dark area prolonged in middle along anterior and posterior mar-
gins; dark area may extend to sides; [Satô & Kimura (1994) and Ohba et al. (1994) described the median dark area
on the protergum as cross shaped with each arm relatively wide and subequal in width, unlike cruciata where the
anterior and posterior margins narrowly prolonged]; most of posterior half of thoracic terga obliquely pale across
corners and narrowly across posterior margin; pale areas across posterior margins of most abdominal terga
restricted to 2 separate areas with the outer area larger; median line wide (maximum width across abdominal terga
3, 4), and terga widely separated (as in Figs 81, 82); abdominal glands not investigated; antennal segment 3 with
closely adpressed finger like projections subequal in length to adjacent sense cone; mandibles lacking retinaculum.
Remarks. The authors herein did not examine specimens of this species which is now protected in Japan
(Ohba pers. com.). Ohba et al. (1994:17 Fig. 1) depicted the gills as single branched with a very short posterior
swelling which could be the retracted second branch. Jeng et al. (2003:544) differentiated owadai and hydrophila
on pronotal colouration, length of gills (those of hydrophila are shorter with more obtuse apices), and pygopod
structure (in owadai the basal row of pygopods has four retractable filaments).
Luciola substriata Gorham
Figs 31–42, 70, 83, 84
Luciola substriata Gorham 1880:100; 1895:305. Olivier 1902a:86; 1913a:271; 1913b:59. Ho et al. 1997: 47. Ho, 2002:50. Jeng
et al. 2003:539. Fu et al. 2005a: 83; 2005b:501; 2009: 243.
Material examined. CHINA: Eleven first and six sixth instars bred from an original population of 12 males and
four females collected from Xianjian village, Hongshan District, Wuhan City, Hubei Province. July 23, 2008
(NHMHAU).
Diagnosis. Dark brown well sclerotised metapneustic larvae with all body plates closely adpressed; mandibles
lacking teeth; apical labial and maxillary palpomeres with sense organs along ventral surfaces; not possible to dis-
tinguish from larvae of L. aquatilis.
Remarks. Ballantyne and Lambkin (2009) scored two different populations of males of this species from both
mainland China and offshore China (Taiwan) with results indicating morphological distinctiveness and two distinct
species in an unresolved species complex within Luciola substriata.
Luciola aquatilis Thancharoen
Figs 85, 86
Luciola aquatilis Thancharoen et al., 2007:55.
Material examined. THAILAND: 13º 40’20.01” N, 100º33’13.50” E, Six sixth instars, population from Samut
Prakarn Province Bangyor district, Phrapradaeng, 4.88 m, November 13, 2003, collected and bred in laboratory by
A. Thancharoen (NHMHAU).
Diagnosis. Dark brown well sclerotised metapneustic larvae with all body plates closely adpressed; mandibles
lacking teeth; apical labial and maxillary palpomeres with sense organs along ventral surfaces; not possible to dis-
tinguish from larvae of L. substriata.
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FIREFLY LARVAE
FIGURES 31–36. Luciola substriata first instar larva, SEM. 31, 32 body, dorsal (31) and ventral; 33 ventral abdominal seg-
ments 3–7; 34 anterior portion of prothorax and retracted head, ventral; 35 left antenna, ventral; 36 apical maxillary palpomere,
ventral.
Semiaquatic larvae
Mode of life. Lacking gills; The semiaquatic designation arises from the ability of these larvae to enter water,
apparently searching for prey, and to remain submerged for a considerable period.
Body: (Fu & Ballantyne 2008, Figs 2–4, 23, 33, 34) protergum and remaining terga except terminal tergum
with raised uneven median line; tergal plates not explanate, margins thickened and laterotergites not visible at sides
FU ET AL.
18 · Zootaxa 3405 © 2012 Magnolia Press
of body when viewed from above; all tergal plates except the last with four short rounded projections along the
posterior margins; dorsal body with heavily sclerotised plates and no obvious exposed membranous areas; ventral
body not heavily sclerotised, flexible; laterotergites and laterosternites visible in abdomen (Fu & Ballantyne 2008);
no eversible abdominal structures bearing glands. Fourteen pygopodia arise from six basal stalks (the two dorsolat-
eral stalks branch into three); densely packed recurved hooks occur only on the ventral surface of each exerted
pygopod (Fig. 92).
FIGURES 37–42. L. substriata larval head, 6th instar, SEM. 37–38 anterior dorsal surface uppermost, 39 anterior half of head,
dorsal; 40 ventral; 41 left antenna, dorsal; 42 apical maxillary palpomere, ventral.
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FIREFLY LARVAE
Head: Antennal segment 3 elongate, surmounted by several hairs, subequal in length and width to adjacent
sense cone; apex of segment 2 not strongly oblique (Figs 43– 48; Fu & Ballantyne 2008; Figs 28, 29).
Mouthparts: Mandibles: with 2 toothed retinaculum, most of dorsal surface with very faint closely parallel
ridges; single line of thick stout setae just inside lateral margin on dorsal surface, setae not appearing split at apices;
outer edge of mandibles bearing hairs along most of margin (Figs 43, 46, 71). Maxillae: (Figs 44, 45, 46) apical
palpomere with terminal sense organs (Fu & Ballantyne 2008 Fig. 31 LA = galea not lacinia). Labium: (Figs 44,
45) palpi widely separated at their bases; ventral surface of heart shaped prementum with hairless “keel”.
Remarks. Lomboot et al. (2007) also recorded Pteroptyx malaccae as semiaquatic as it lives near brackish
bodies of water in damp soil, but no evidence of these larvae submerging to find prey or of an ability to survive
immersion was presented, and this larval type is dealt with under terrestrial larvae.
FIGURES 43–48. Pygoluciola qingyu larval head, 5th instar, SEM. 43 dorsal, anterior half; 44–45 ventral; 46 mandibles; 47
right antenna; 48 detail apex right antenna.
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Pygoluciola qingyu Fu et Ballantyne
Figs 43–48, 63–4, 71, 92
Pygoluciola qingyu Fu et Ballantyne, 2008:1.
Material examined. CHINA: Eight fifth instar larvae, E Mei Mountain, Sichuan Province, July 8, 2006
(NHMHAU).
Diagnosis. Very similar in morphology to larvae of L. gorhami, L. praeusta. L. terminalis, L. cerata, L. anceyi
from which it differs most obviously in having 2 inner teeth on the mandibles (Figs 46, 71). Found near shallow
bodies of water along road sides. Known to be actively predacious on living ants.
Remarks. It is very possible that some of the other species with similar dorsal morphology, but a single
toothed retinaculum, (e.g. L. gorhami, L. praeusta. L. terminalis, L. cerata, L. anceyi) could be found to be semi-
aquatic (Chen 2003; Fu & Ballantyne in prep.). There are however no records to suggest these species are actively
predacious like P. qingyu. Chen (2003) recorded L. terminalis as terrestrial, the adults occurring close to rice fields,
while L. praeusta was found alongside mountain roads in an environment that approaches that of Pygoluciola
qingyu with larvae recorded as terrestrial. Fletcher (1919:28) found L. gorhami in Pusa India under fallen and rot-
ting leaves, indicated it fed only on dead Lepismidae (Thysanura), and would not touch living food. Mehta (1932)
considered the spinose tergal plates equated with a burrowing mode of life.
Terrestrial larvae
Terrestrial larvae are of two types.
1. Having laterally explanate tergal margins. Body: (Figs 87, 88) median line extending from anterior margin of
protergum to posterior margin of penultimate abdominal segment; tergal margins laterally explanate on all but ter-
minal tergum and covering ventral body structures including laterotergites; thoracic tergal margins widely explan-
ate in Emeia pseudosauteri, narrowly prolonged in abdominal segments 1–8; tergal plates well sclerotised,
intersegmental membranes visible if larva is extended; plates on ventral body surface usually coloured and sepa-
rated by membrane; laterosternites and laterotergites (bearing spiracles) visible in abdomen; lacking lateral eversi-
ble structures and glands. As. circumdata having at least 58 pygopodia arising from at least 22 basal stalks;
recurved hooks surround the entire surface of each expanded pygopod (Fig. 93).
Head: antennal segment 3 in As. circumdata (Figs 51, 52) , some Atyphella spp., and Emeia pseudosauteri (Fu
et al. 2012; Fig. 28), surmounted by hairs, slightly longer than adjacent short, squat, broad sense cone.
Mouthparts: Mandibles (Figs 49, 72) lacking retinaculum; dorsal surface smooth, not resembling overlapping
scales. Maxillae: (Figs 50, 51, 53) apical palpomere with terminal sense organs. Labium: (Figs 50, 51, 53) palpi
occurring at anterolateral corners of prementum; apical labial palpomere with terminal sense organs; ventral sur-
face of prementum longer than wide, largely hairless. e.g. Asymmetricata circumdata, Emeia pseudosauteri.
Remarks. Bourgeoisia hypocrita larvae (Fu et al 2012; Figs 50, 51) have abdominal tergal plates with 2 thin
posterolateral prolongations and a brush of hairs at the apex of the tarsungulus on all legs. Larvae of L. parvula, L.
filiformis and L. kagiana have a similar tergal plate arrangement and remain to be investigated further (Chen 2003).
Atyphella spp., Magnalata limbata, Lloydiella majuscula larvae are not investigated here beyond information con-
tained in Ballantyne & Lambkin (2009).
2. Lacking laterally explanate tergal margins. Body: (Figs 89, 90) all terga except the last with median
line; tergal plates not sclerotised to margins, membranous areas on dorsal surface usually visible, laterotergites vis-
ible from above at sides of thorax and abdomen; plates on ventral body surface usually coloured, flexible; venter of
abdomen with median sterna, margined by elongate laterosternites and at sides laterotergites bearing spiracles on
all but terminal segment (Fig. 90).
Head: (Fig. 59) antennal segment 3 elongate, surmounted by hairs, elongate, slender, about as long as elongate
sense cone; apex of antennal segment 2 strongly oblique and segment 3 may appear to be longer than adjacent
sense cone.
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FIREFLY LARVAE
Mouthparts: (Fig. 55) mandibles lacking retinaculum, but with dense brush of hairs along inner basal half;
most of dorsal surface slightly irregular; single line of thick stout setae bordering lateral margin on dorsal surface;
outer edge of mandibles bearing hairs all along margin. Maxillae: (Figs 56, 58) apical palpomere with terminal
sense organs. Labium: (Figs 56, 58) with palpi separated at their bases; ventral surface of prementum with largely
hairless “keel” at least 3 times as long as wide.
Remarks. Some Pteroptyx, Colophotia praeusta (Ballantyne & Lambkin 2009), probably also Pyrophanes and
some Australian Luciola have dorsal plates not projecting at the sides, and a relatively soft body, with laterotergites
usually visible at the sides. They lack lateral eversible structures and glands and mandibles are not toothed.
FIGURES 49–54. Asymmetricata circumdata larval head, 5th instar, SEM. 49 dorsal; 50–51 ventral (51 anterior half of head
only); 52 right antenna, ventral; 53 apical labial and maxillary palpomeres and galea; 54 apex of apical maxillary palpomere.
FU ET AL.
22 · Zootaxa 3405 © 2012 Magnolia Press
FIGURES 55–60. Pteroptyx valida larval head 4th instar. SEM. 55 dorsal; 56 ventral (56, 57 anterior end to right of page); 57
anterior dorsal surface uppermost; 58 apical labial and maxillary palpomeres and galea from below; 59 right antenna, from
below; 60 tip of apical maxillary palpomere.
Asymmetricata circumdata (Motsch.)
Figs 49–54, 65–6, 72, 87, 88, 93
Luciola circumdata Motschulsky, 1854:50 (female). Lacordaire, 1857:338. Olivier, 1885:364 (Male, female); 1902a:84. Bour-
geois, 1890:184. McDermott, 1964:44; 1966: 101. Ballantyne, 1987b:181, Fig. 1.
Asymmetricata circumdata (Motsch.) Ballantyne & Lambkin 2009:32.
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FIREFLY LARVAE
Material examined. CHINA: Eight sixth instars bred from an original population of 4 males and 1 female
collected from Xishuangbanna Tropical Botanical Garden, Chinese Academy of Science, Jinghong City,
Xishuangbanna Province, May 16, 2010 (NHMHAU).
Diagnosis. The only species of Asymmetricata with reliably associated larvae; distinguished by its laterally
explanate tergal margins, short antennal sense cone and the largely yellow dorsal colouration, broken by a continu-
ous wide median band of black running from the anterior area of thoracic tergum 2 to the posterior margin of termi-
nal tergum (Figs 49–54, 87, 88). At least 58 expanded pygopodia arise from basal stalks which may branch more
than once; recurved hooks surround the entire surface of the expanded pygopod (Fig. 93).
FIGURES 61–66. Larval pygopodia. 61–62 Aquatica leii (5th instar); 63–64 Pygoluciola qingyu (6th instar); 65–66
Asymmetricata circumdata (6th instar).
FU ET AL.
24 · Zootaxa 3405 © 2012 Magnolia Press
Pteroptyx valida Olivier
Figs 55–60, 89, 90
Pteroptyx validum Olivier, 1909a: 397; 1910: 48.
Pteroptyx valida Olivier. McDermott, 1966: 117. Ballantyne & McLean, 1970: 256. Wing et al., 1983:86. Lloyd & Wing, 1981:
459. Lloyd et al.,1989:373. Lloyd, 1977:63; 1979: 300; 1981:97; 1983:141. Ballantyne, 2001:79.
Material examined. MALAYSIA: Selangor, two 4th instar larvae (of a total of 33) bred from eggs collected from a
gravid female by Rasainthiran Menayah; used for SEM.
Diagnosis. Without laterally explanate tergal margins and laterotergites usually visible at sides of body when
viewed from above (Figs 55–60, 89–90).
Remarks. Redescribed by Ballantyne and Menayah (2002) from specimens bred by Menayah.
Discussion
As in many studies on firefly morphology in SE Asia this study has been constrained by the number of reliably
associated larvae available for dissection and examination.
Aquatic larvae are clearly of two types. Bottom dwellers do not swim, have lateral abdominal gills, sensory
areas at the tips of the palps, and lack hairs at the tip of antennal segment 3. Eversible glands along the sides of the
body produce repellent protective materials. All Aquatica, and two Luciola species are bottom dwellers. It is not
yet possible to distinguish between the gilled aquatic larvae of Aquatica, and those of Luciola cruciata and L. owa-
dai, where larval morphology may be primarily an adaptation to mode of life (Fu et al. 2012; Fig. 7, Nodes 1 and
3). Back swimming larvae are metapneustic, can swim easily and live just below the water surface. Not only is
their entire external body covered in hard plates lacking obvious membranous areas, but the palpi have sensory
areas along the ventral (and hence uppermost) surface when the larva is back swimming. Antennal segment 3 lacks
hairs at its apex. They lack lateral abdominal glands and their main protection in the aquatic environment may be
their hard exoskeleton.
Terrestrial larvae are recognised by the absence of gills or terminal spiracles on dorsal protrusions, and those
investigated have antennal segment 3 surmounted by hairs. Laterally explanate tergal margins are characteristic of
the Atyphella “complex” of Ballantyne and Lambkin (2009) (viz. Magnalata spp., Atyphella spp., Lloydiella spp.)
as well as Asymmetricata. It is not yet possible to distinguish these genera using larval morphology and where spe-
cific distinctions exist, they rely largely on colour patterns (Ballantyne & Lambkin 2000). Laterally explanate ter-
gal margins also occur in several Luciola species (L. parvula, L. filiformis, L. kagiana). The larval type of Luciola
italica (type species of Luciola s. str.) is unknown.
Terrestrial larvae lacking laterally explanate tergal margins are very similar and it is not possible to distinguish
generic differences between larvae of Pteroptyx, Colophotia and Australian Luciola partly because the reliable
associations needed to identify species are lacking. Four species of Australian Luciola were distinguished at spe-
cific level in Ballantyne and Lambkin (2000) by the density and arrangement of small rounded projections on the
dorsum of all body segments except the last.
The larvae of Bourgeoisia hypocrita are probably terrestrial (Ballantyne 1968; Deheyn & Ballantyne 2009) but
nothing is known of their life cycle. They have a brush of hairs arising at the apex of the tarsungulus in all legs but
the significance of this in relation to possible habitat is unexplained. Adult (flightless) females were found in a ter-
restrial environment among rotting leaves and it is reasonable to suppose the larvae inhabit a similar area. The tri-
lobite-like larva of Emeia pseudosauteri is terrestrial.
In Australia laterally explanate plates occur in all known Atyphella, which have been found in leaf litter in rain
forest areas along the Queensland and northern New South Wales east coast. Luciola australis and L. orapallida,
which lack explanate tergal margins, occur in tidal mangrove flats (L. australis around Gladstone and Townsville),
with L. orapallida from further north in eastern Cape York Peninsula. It might be conjectured that the slim body
shape of the mud flat dweller facilitates its passage through the wet soil, while the dish like tergal plates of an
Atyphella sp. makes progress under leaf litter easier, were it not for the fact that larvae of L. nigra (lacking laterally
explanate tergal margins) and A. scintillans (having laterally explanate tergal margins) both occur in leaf litter situ-
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FIREFLY LARVAE
ations in relict rain forest in suburban back yards in parts of Brisbane (Ballantyne 1993). Additionally L. flavicollis
and L. nigra larvae occur in open forest locations along the Queensland coast.
FIGURES 67–72. Left mandibles, dorsal. 67 Aq. ficta; 68 Aq. lateralis; 69 L. cruciata; 70 L. substriata; 71 P. qingyu ; 72 A.
circumdata.
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26 · Zootaxa 3405 © 2012 Magnolia Press
FIGURES 73–78. Habitus, larvae (scale bar = 5 mm). 73–74 Aq. ficta dorsal and ventral (6th instar); 75–76 Aq. lateralis dorsal
and ventral (5th instar); 77–78 Aq. leii dorsal and ventral (5th instar).
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FIREFLY LARVAE
FIGURES 79–84. Habitus, larvae (scale bar =5 mm). 79–80 Aq. wuhana dorsal and ventral (4th instar); 81–82 L. cruciata dor-
sal and ventral (5th instar); 83–84 L. substriata dorsal and ventral (6th instar).
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FIGURES 85–90. Habitus, larvae (scale bar =5 mm). 85–86 L. aquatlis dorsal and ventral (6th instar); 87–88 A. circumdata
dorsal and ventral (6th instar); 89–90 P. valida dorsal and ventral (4th instar).
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FIREFLY LARVAE
FIGURE 91. Aq. leii semidiagrammatic representation of everted pygopodia from below (5th instar) (scale bar= 0.2 mm).
FIGURE 92. P. qingyu semidiagrammatic representation of everted pygopodia from below (5th instar) (scale bar= 0.2 mm).
Semiaquatic larvae like Pygoluciola qingyu may simply be opportunistic feeders capable of remaining
immersed for some time and the semiaquatic designation inappropriate, as there are no obvious morphological
modifications. Pteroptyx malaccae larvae living in brackish environments beside rivers were recorded as semi-
aquatic but this designation may also be inappropriate as no evidence of the larvae actually submerging to find
prey, or of their ability to remain submerged for a period, was presented (Lomboot et al. 2007). These larvae
inhabit an environment very similar to that occupied by L. australis and L. orapallida, which are treated here as ter-
restrial.
The north American Pyractomena lucifera (Melsh.) (Lampyrinae) larvae occur in fresh water marshes, can
capture prey both above and below the water surface, and may remain submerged for many days (Buschman 1984).
Similarly, the larvae of species of terrestrial tiger beetles (Cicindelinae: Carabidae) can survive periodic flooding
and immersion in water with very low oxygen levels for several days (Brust et al. 2005, Brust & Hoback 2009;
Hoback 1998). There is no evidence yet available to suggest that other, softer bodied firefly larvae could not also
survive immersion for a prolonged period.
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FIGURE 93. A. circumdata semidiagrammatic representation of everted pygopodia from below (6th instar) (scale bar= 0.2
mm).
In P. qingyu the mandibles have 2 teeth on their inner margins and a very hard dorsal exoskeleton and the spe-
cies is known to aggressively attack live prey like ants. It is tempting to interpret the similar morphology of various
other Luciola species viz. Luciola gorhami, L. praeusta and L. terminalis (where the mandibles have a single tooth
and the tergal plates are similarly configured) as indicating a comparable mode of attack. However it appears that
these latter larvae are terrestrial and probably all feed on dead or decaying animal matter.
Interpretations of the various morphologies to determine mode of life has proved difficult with the exception of
those that are clearly aquatic, and larvae lacking gills are assumed to be terrestrial when no other information is
available. The semiaquatic designation can only arise from observation of the larvae in their natural habitat.
Larval types may be of taxonomic significance. Fu et al. (2012) recognised various clades (some as yet unde-
scribed) where similar larval types occur:
1. Luciola s. str. clade (Fu et al. 2012; Fig. 8, Node 38) including the type species L. italica. While unable to
obtain italica larvae, Fu et al. included kagiana and parvula, both with laterally explanate tergal margins in the
larvae (Table 2). Larvae of B. hypocrita (Fu et al. 2012; Fig. 7, Node 38) fell into the wider Luciola s. str.
clade; in hypocrita the tergal margins are prolonged at their posterolateral corners and cover laterotergites from
above.
2. Fu et al. 2012; Figure 8, Node 46 includes two species (L. cerata and L. praeusta) having larvae with well scle-
rotised tergal plates projecting laterally sufficiently to cover the laterotergites, and mandibles with a single
inner tooth (Table 2).
3. Pygoluciola qingyu (Fu et al. 2012; Fig. 8, Node 49) has larvae similar to those described in 2 above, while
mandibles have 2 inner teeth and the semiaquatic larva is aggressive towards living insects especially ants.
This is thus far the only species of Pygoluciola (Fu & Ballantyne 2008; Fu et al. 2012; Fig. 8, Node 48) where
larvae have been reliably associated.
4. The two species of back swimming metapneustic larvae (Table 2) were grouped in a distinctive clade ( Fu et al.
2012; Fig. 8, Node 45) .
5. The new genus proposed for Curtos pseudosauteri (Fu et al. 2012) is known from distinctive trilobite-like lar-
vae (Fu et al. 2012).
6. Certain species of Australian and New Guinean firefly presently assigned to Luciola, are being investigated
further (Table 2). Fu et al. (2012) grouped them in a clade separate from certain other New Guinean Luciola
and Pteroptyx species (Fig. 7, Node 28). Their larvae are elongate, slim, with laterotergites usually visible at
the sides of the abdomen and it is difficult to distinguish them.
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FIREFLY LARVAE
TABLE 3. List of aquatic larvae.
Acknowledgements
We are grateful to Anchana Thancharoen for sending specimens for this study. Fu is supported by Hubei Provincial
Foundation of Natural Science # 4006-086064, Chinese National Science Foundation # 30900147, # 31172137 and
Fundamental Research Funds for the Central Universities (Program No. 2011PY054). Ballantyne thanks Charles
Sturt University for continued support. Lambkin thanks Queensland Museum for continuing to support her
taxonomic and systematic work.
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