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<mods:title id="9EBABEDA2935516481C985BABBC8ACB3">Systematics of Indo-Pacific fireflies with a redefinition of Australasian Atyphella Olliff, Madagascan Photuroluciola Pic, and description of seven new genera from the Luciolinae (Coleoptera: Lampyridae)</mods:title>
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<mods:namePart id="4B221D4C4693E9ADC6C5576611941261">Ballantyne, Lesley A.</mods:namePart>
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<mods:namePart id="09791FA74C5925A5EEFFD61F7548EF86">Lambkin, Christine</mods:namePart>
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<subSubSection id="C32734F8BE10FF80FF3C56D6234EEF89" box="[151,361,415,441]" pageId="60" pageNumber="61" type="nomenclature">
<paragraph id="8B826773BE10FF80FF3C56D6234EEF89" blockId="60.[151,361,415,476]" box="[151,361,415,441]" pageId="60" pageNumber="61">
<heading id="D0CAD01FBE10FF80FF3C56D6234EEF89" bold="true" box="[151,361,415,441]" fontSize="11" level="1" pageId="60" pageNumber="61" reason="1">
<emphasis id="B949BB61BE10FF80FF3C56D6234EEF89" bold="true" box="[151,361,415,441]" pageId="60" pageNumber="61">
<taxonomicName id="4C3D1CF0BE10FF80FF3C56D62328EF89" ID-CoL="5FVV" authority="Ballantyne &amp; Lambkin, 2009" authorityName="Ballantyne &amp; Lambkin" authorityYear="2009" box="[151,271,415,441]" class="Insecta" family="Lampyridae" genus="Lloydiella" kingdom="Animalia" order="Coleoptera" pageId="60" pageNumber="61" phylum="Arthropoda" rank="genus" status="gen. nov.">
<emphasis id="B949BB61BE10FF80FF3C56D62328EF89" bold="true" box="[151,271,415,441]" italics="true" pageId="60" pageNumber="61">Lloydiella</emphasis>
</taxonomicName>
<taxonomicNameLabel id="A27A061ABE10FF80FEBD56D6234EEF89" box="[278,361,415,441]" pageId="60" pageNumber="61" rank="genus">gen. n.</taxonomicNameLabel>
</emphasis>
</heading>
</paragraph>
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<subSubSection id="C32734F8BE10FF80FF3C568B236CEFEC" box="[151,331,450,476]" pageId="60" pageNumber="61" type="description">
<paragraph id="8B826773BE10FF80FF3C568B236CEFEC" blockId="60.[151,361,415,476]" box="[151,331,450,476]" pageId="60" pageNumber="61">
(
<figureCitation id="13067BF6BE10FF80FF34568B2364EFEC" box="[159,323,450,476]" captionStart-0="FIGURES 204211" captionStart-1="FIGURES 212224" captionStartId-0="162.[151,269,1874,1898]" captionStartId-1="163.[151,269,1698,1722]" captionTargetBox-0="[151,1435,204,1847]" captionTargetBox-1="[151,1436,194,1656]" captionTargetId-0="figure-76@162.[151,1436,194,1850]" captionTargetId-1="figure-75@163.[151,1436,194,1674]" captionTargetPageId-0="162" captionTargetPageId-1="163" captionText-0="FIGURES 204211. Lloydiella spp. paratype males. 204, 208, 210 Ll. uberia; 205, 209 Ll. japenensis; 206, 207, 211 Ll wareo (BPBM). 204206 dorsal whole body (204, 206 have abdomen dissected); 207210 dorsal pronota; 211 four specimens on one card, L to R male ventral, female ventral (no head or pronotum), male dorsal, male ventral." captionText-1="FIGURES 212224. Lloydiella spp. paratype males. Ll. japenensis 212214, 217219, 222. Ll. majuscula holotype 223. Ll. uberia 215, 220, 224. Ll. wareo 216, 221, 224. 212, 215, 216 aedeagi ventral; 213 aedeagus left laterodorsal; 214 aedeagus slightly laterodorsal; 217, 218, 220, 221 aedeagal sheath, ventral; 219 aedeagal sheath dorsal; 222224 abdominal ventrites 6, 7 and tergite 8 ventral." pageId="60" pageNumber="61">Figs 204224</figureCitation>
)
</paragraph>
</subSubSection>
<subSubSection id="C32734F8BE10FF80FF3C555B204CEC1C" box="[151,619,530,556]" pageId="60" pageNumber="61" type="type_taxon">
<paragraph id="8B826773BE10FF80FF3C555B204CEC1C" blockId="60.[151,1437,530,2036]" box="[151,619,530,556]" pageId="60" pageNumber="61">
<emphasis id="B949BB61BE10FF80FF3C555B231EEC1C" bold="true" box="[151,313,530,556]" pageId="60" pageNumber="61">
<typeStatus id="5486D9D1BE10FF80FF3C555B22F3EC1C" box="[151,212,530,556]" pageId="60" pageNumber="61">Type</typeStatus>
species.
</emphasis>
<taxonomicName id="4C3D1CF0BE10FF80FEEB555B204CEC1C" authority="Lea." authorityName="Lea." box="[320,619,530,556]" class="Insecta" family="Lampyridae" genus="Atyphella" kingdom="Animalia" order="Coleoptera" pageId="60" pageNumber="61" phylum="Arthropoda" rank="species" species="majuscula">
<emphasis id="B949BB61BE10FF80FEEB555B2017EC1B" box="[320,560,530,555]" italics="true" pageId="60" pageNumber="61">Atyphella majuscula</emphasis>
Lea.
</taxonomicName>
</paragraph>
</subSubSection>
<subSubSection id="C32734F8BE10FF82FF6D557321B9EFFA" lastPageId="62" lastPageNumber="63" pageId="60" pageNumber="61" type="diagnosis">
<paragraph id="8B826773BE10FF80FF6D557323D6ED5C" blockId="60.[151,1437,530,2036]" pageId="60" pageNumber="61">
<emphasis id="B949BB61BE10FF80FF6D55732319EC64" bold="true" box="[198,318,570,596]" pageId="60" pageNumber="61">Diagnosis</emphasis>
. Distinguished most obviously among New Guinean species by its strong dorsal colouration of orange pronotum and dark elytra; pronotal width subequal to humeral width; anterior hypomeron not flat to neck; 23 interstitial lines; antennae longer than GHW; labial palpi apical segment dentate and laterally flattened, 2 or more teeth; median longitudinal trough may be present on ventral surface of T8; anterior prolongations of T8 longer than posterior entire portion, narrow and expanded vertically; aedeagus with LL/ML wide; LL not diverging along their median dorsal length and narrower at their apex than ML; BP hooded. Known female macropterous and coloured like male. Associated larva with laterally explanate tergal margins concealing laterotergites from above.
</paragraph>
<paragraph id="8B826773BE10FF80FF6C543322E3EACC" blockId="60.[151,1437,530,2036]" pageId="60" pageNumber="61">
<emphasis id="B949BB61BE10FF80FF6C5433232AEDA4" bold="true" box="[199,269,890,916]" pageId="60" pageNumber="61">Male.</emphasis>
Pronotum dorsal surface lacking irregularities in posterolateral areas and longitudinal groove in lateral areas; punctation dense. Anterior margin not explanate. Pronotum wider across posterior area than rest, never subparallel-sided; pronotal width subequal to humeral width. Anterolateral corners rounded obtuse; lateral margins in anterior half divergent posteriorly; lateral margins in posterior half usually diverge then converge with rounded convergence except diverging along length in
<taxonomicName id="4C3D1CF0BE10FF80FC1A5353260AEA03" authority="Ballantyne &amp; Lambkin, 2009" authorityName="Ballantyne &amp; Lambkin" authorityYear="2009" box="[945,1069,1050,1075]" class="Insecta" family="Lampyridae" genus="Pygatyphella" kingdom="Animalia" order="Coleoptera" pageId="60" pageNumber="61" phylum="Arthropoda" rank="species" species="japenensis" status="sp. nov.">
<emphasis id="B949BB61BE10FF80FC1A5353260AEA03" box="[945,1069,1050,1075]" italics="true" pageId="60" pageNumber="61">japenensis</emphasis>
</taxonomicName>
<emphasis id="B949BB61BE10FF80FB9D5353265DEA04" bold="true" box="[1078,1146,1050,1076]" pageId="60" pageNumber="61">
<taxonomicNameLabel id="A27A061ABE10FF80FB9D5353265DEA04" box="[1078,1146,1050,1076]" pageId="60" pageNumber="61" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
and some
<taxonomicName id="4C3D1CF0BE10FF80FB5553532761EA03" authority="Ballantyne &amp; Lambkin, 2009" authorityName="Ballantyne &amp; Lambkin" authorityYear="2009" box="[1278,1350,1050,1075]" class="Insecta" family="Lampyridae" genus="Lloydiella" kingdom="Animalia" order="Coleoptera" pageId="60" pageNumber="61" phylum="Arthropoda" rank="species" species="wareo" status="sp. nov.">
<emphasis id="B949BB61BE10FF80FB5553532761EA03" box="[1278,1350,1050,1075]" italics="true" pageId="60" pageNumber="61">wareo</emphasis>
</taxonomicName>
<emphasis id="B949BB61BE10FF80FAFB535327B3EA04" bold="true" box="[1360,1428,1050,1076]" pageId="60" pageNumber="61">
<taxonomicNameLabel id="A27A061ABE10FF80FAFB535327B3EA04" box="[1360,1428,1050,1076]" pageId="60" pageNumber="61" rank="species">sp. n.</taxonomicNameLabel>
</emphasis>
; lateral margins lacking indentation at mid-point, or sinuousity in horizontal or vertical plane, indentation near posterolateral corner, and irregularities at corner; posterolateral corners rounded or angulate; rounded corners obtuse, or subequal to 90°, angulate corners 90° or less, never very acute, inclined obliquely to median line; posterolateral corners project to or beyond median posterior margin and separated by shallow emargination or not.
</paragraph>
<paragraph id="8B826773BE10FF80FF6C524327BBEB44" blockId="60.[151,1437,530,2036]" pageId="60" pageNumber="61">Hypomeron closed. Median area of hypomeron not elevated in vertical plane, lateral margins not sinuate from above; anterior area of hypomeron not flat to side of head, posterior area flat and strongly adpressed; median area sloping smoothly from anterior raised area to posterior flat area; pronotal width/ GHW index 1.6.</paragraph>
<paragraph id="8B826773BE10FF80FF6C52CB26E3E854" blockId="60.[151,1437,530,2036]" pageId="60" pageNumber="61">Elytral punctation dense, not linear, not as large as pronotum, nor widely and evenly spaced; apices not deflexed; epipleuron and suture extend beyond mid-point of elytron, not extending as a ridge around apex, not thickened in their apical half; 2 or 3 interstitial lines, none exceed suture; elytral carina absent; in horizontal specimen viewed from beneath epipleuron at elytral base wide, covers humerus, viewed from above arises anterior to posterior margin of MS; epipleuron developed as a ridge along most of its length; sutural margins approximate along most of length in closed elytra; lateral margins parallel-sided or convex.</paragraph>
<paragraph id="8B826773BE10FF80FF6C513B2194E9C4" blockId="60.[151,1437,530,2036]" pageId="60" pageNumber="61">Head moderately depressed between eyes; moderately exposed in front of pronotum not capable of complete retraction within prothoracic cavity; eyes close to moderately separated beneath at level of posterior margin of mouthpart complex; eyes above labrum close to moderately separated; frons-vertex junction rounded, no elevation in median area; posterolateral eye excavation not strongly developed, not visible in resting head position; antennal sockets on head between eyes, not contiguous, separated by&lt;ASW; clypeolabral suture present, flexible, not in front of anterior eye margin when viewed with labrum horizontal; outer edges of labrum reach inner edges of closed mandibles. Mouthparts functional; apical segment of labial palpi nonlunate, strongly flattened, like a wide triangle, inner edge dentate, with 2 or more teeth. Antennae 11 segmented; length greater than GHW to twice GHW; no segments shortened, flattened or expanded; pedicel not produced; FS1 not shorter than pedicel; all FS longer than wide.</paragraph>
<paragraph id="8B826773BE11FF81FF6C57D1238CEEEA" blockId="61.[151,1436,152,2018]" pageId="61" pageNumber="62">Legs with inner tarsal claw not split; lacking MFC; no femora or tibiae swollen or curved; no basitarsi expanded or excavated.</paragraph>
<paragraph id="8B826773BE11FF81FF6C57A12704ED9A" blockId="61.[151,1436,152,2018]" pageId="61" pageNumber="62">
Abdomen lacking cuticular remnants in association with aedeagal sheath; no ventrites with curved posterior margins, nor with anterior margins extending into emarginated posterior margin of anterior segment; LO in V7 entire, occupying most of V7; entire LO reaching to sides not posterior margin; neither anterior nor posterior margin of LO medially emarginated; posterior area of V7 behind LO not arched or swollen, muscle impressions not visible in this area; LO in V6 entire, occupying almost all V6. MPP present, symmetrical, apex rounded, entire, not laterally compressed, short or L=W; MPP not strongly inclined dorsally nor engulfed by T8 apex; lacking dorsal ridge, ventral longitudinal trough. V7 lacking median carina, median longitudinal trough, narrow anteromedian depression on anterior face of LO, lacking PLP, incurving lobes or pointed projections, median dimple or reflexed lobes. T7 lacking prolonged posterolateral corners. T8 usually well sclerotised; symmetrical, sometimes slightly longer than wide in visible posterior area where lateral margins either converge gently posteriorly, or W=L, never subparallel-sided; visible posterior area not narrowing abruptly, lacking median posterior emargination, median posterior projections, not inclined ventrally nor engulfing posterior margin of V7 nor MPP, not extending conspicuously beyond median posterior margin of V7; T8 ventral surface lacking flanges, lateral depressed troughs, asymmetrical projections, median posterior ridge; with obvious elongate, margined, median longitudinal trough in
<taxonomicName id="4C3D1CF0BE11FF81FBA2545126A6ED01" box="[1033,1153,792,817]" class="Insecta" family="Lampyridae" genus="Atyphella" kingdom="Animalia" order="Coleoptera" pageId="61" pageNumber="62" phylum="Arthropoda" rank="species" species="majuscula">
<emphasis id="B949BB61BE11FF81FBA2545126A6ED01" box="[1033,1153,792,817]" italics="true" pageId="61" pageNumber="62">majuscula</emphasis>
</taxonomicName>
only; concealed anterolateral arms of T8 shorter than, as long as or longer than visible posterior portion, narrow horizontally, expanded dorsoventrally, not laterally emarginated before their origins, apices lacking bifurcation of inner margin and bases lacking ventrally directed pieces; lateral margins of T8 not enfolding sides of V7.
</paragraph>
<paragraph id="8B826773BE11FF81FF6C54F121E9EAF2" blockId="61.[151,1436,152,2018]" pageId="61" pageNumber="62">Aedeagal sheath never&gt; 4 times as long as wide; lacking paraprocts; asymmetrical in posterior area where sheath sternite emarginated on right side from point of attachment of tergite; sheath not angulate on R or L sides; not subparallel-sided, not emarginated on either side preapically, anterior half of sheath sternite broad, apically rounded; tergite lacking lateral arms that extend anteriorly at sides of sternite; tergite not subdivided, lacking lateral projecting pieces along posterior margin of tergite 9; tergite 9 lacking dark anterior transverse band (a narrow anterior area of sheath tergite often more heavily sclerotised, but not darker, than rest); tergite lacking cuticular projection to left for muscle attachment.</paragraph>
<paragraph id="8B826773BE11FF81FF6C539926F7E862" blockId="61.[151,1436,152,2018]" pageId="61" pageNumber="62">Aedeagus L/W 3/1; LL lack lateral appendages, visible from beneath at sides of ML; LL/ML wide; LL of equal length; slightly shorter than median lobe; LL not diverging along inner margins and separated there by&gt; half their length; LL base width does not=LL apex width which is narrower than ML; LL apices not expanded horizontally, not outturned; preapical inner and outer margins of LL obliquely truncate; dorsal base of LL symmetrical, not excavated, median margin prolonged, pointed, entire; LL lacking hairy appendages along outer ventral margins; apices of LL not outturned nor inturned; lacking projection on left lateral lobe; inner margins lack slender leaflike projection; preapical inner margin of LL obliquely truncate, lacking strongly developed tooth at anterior end of emargination. ML symmetrical, lacking paired lateral teeth, single tooth to left side, not strongly arched, apex not in arrowhead shape, not bulbous, not inclined ventrally. BP not very narrow, not strongly sclerotised, not strongly emarginated along its anterior margin; hooded.</paragraph>
<paragraph id="8B826773BE11FF81FF6C511620C6E9A2" blockId="61.[151,1436,152,2018]" pageId="61" pageNumber="62">
<emphasis id="B949BB61BE11FF81FF6C5116230EE849" bold="true" box="[199,297,1631,1657]" pageId="61" pageNumber="62">Female.</emphasis>
Macropterous and assumed capable of flight. Pronotum lacking irregularities in posterolateral areas of dorsal surface; punctation moderate to dense, not as large as elytron; always wider across posterior margin; indentation of lateral margin and irregularities at posterolateral corner absent; pronotal width subequal to humeral width; outline as for male. Elytral punctation not as large as that of pronotum, nor evenly spaced; interstitial lines 23; elytral carina absent. Head of winged female form, with eyes smaller than male, can be retracted within prothoracic cavity, lacking any anterior prolongation; antennae on head between eyes; clypeolabral suture present and flexible. No legs or parts thereof swollen and /or curved. LO in V6 only, lacking any elevations or depressions or ridges on V7.
</paragraph>
<paragraph id="8B826773BE11FF82FF6C50D621B9EFFA" blockId="61.[151,1436,152,2018]" lastBlockId="62.[151,1436,152,658]" lastPageId="62" lastPageNumber="63" pageId="61" pageNumber="62">
<emphasis id="B949BB61BE11FF81FF6C50D6233DE989" bold="true" box="[199,282,1951,1977]" pageId="61" pageNumber="62">Larva.</emphasis>
Known only in
<taxonomicName id="4C3D1CF0BE11FF81FE7550E921DEE98A" authority="(Ballantyne &amp; Lambkin, 2000)" baseAuthorityName="Ballantyne &amp; Lambkin" baseAuthorityYear="2000" box="[478,1017,1952,1978]" class="Insecta" family="Lampyridae" genus="Atyphella" kingdom="Animalia" order="Coleoptera" pageId="61" pageNumber="62" phylum="Arthropoda" rank="species" species="majuscula">
<emphasis id="B949BB61BE11FF81FE7550E92059E989" box="[478,638,1952,1977]" italics="true" pageId="61" pageNumber="62">Ll. majuscula</emphasis>
(
<bibRefCitation id="EFAC1A82BE11FF81FD2450E921D6E98A" author="Ballantyne, L. A. &amp; Lambkin, C." box="[655,1009,1952,1978]" pageId="61" pageNumber="62" pagination="15 - 93" refId="ref103088" refString="Ballantyne, L. A. &amp; Lambkin, C. (2000) Lampyridae of Australia (Coleoptera: Lampyridae: Luciolinae: Luciolini). Memoirs of the Queensland Museum 46 (1), 15 - 93." type="journal article" year="2000">Ballantyne &amp; Lambkin, 2000</bibRefCitation>
)
</taxonomicName>
; terrestrial; tergal plates sclerotised to margins, lateral tergal margins explanate, slightly ridged, hairy and spinose, covering laterotergites which are not visible from above; arrangement of plates in ventral aspect of thorax and abdomen agrees with that described (
<bibRefCitation id="EFAC1A82BE12FF82FEBC57892051EEEA" author="Ballantyne, L. A. &amp; Lambkin, C." box="[279,630,192,218]" pageId="62" pageNumber="63" pagination="15 - 93" refId="ref103088" refString="Ballantyne, L. A. &amp; Lambkin, C. (2000) Lampyridae of Australia (Coleoptera: Lampyridae: Luciolinae: Luciolini). Memoirs of the Queensland Museum 46 (1), 15 - 93." type="journal article" year="2000">Ballantyne &amp; Lambkin, 2000</bibRefCitation>
;
<bibRefCitation id="EFAC1A82BE12FF82FD28578921C2EEEA" author="Ballantyne, L. A. &amp; Menayah, R." box="[643,997,192,218]" pageId="62" pageNumber="63" pagination="101 - 109" refId="ref103282" refString="Ballantyne, L. A. &amp; Menayah, R. (2002) A description of larvae and redescription of adults of the firefly Pteroptyx valida Olivier in Selangor, Malaysia (Coleoptera: Lampyridae: Luciolinae), with notes on Luciolinae larvae. Raffles Bulletin of Zoology 50 (1), 101 - 109." type="journal article" year="2002">Ballantyne &amp; Menayah, 2002</bibRefCitation>
). Protergum wider than long, lacking tubercles along its anterior margin; posterolateral corners acute; median posterior margins of terga 111 lacking either rounded or pointed projections, posterolateral corners slightly acute; posterolateral corners of terminal tergum slightly produced; punctures in anterior area of terga 211 slightly larger than rest of area; median line extends from anterior to posterior margins in most segments, lacks raised margins; lacking brush of hairs around tarsal claws; mandibles lacking inner teeth; antennal segment 3 short, sense cone adjacent to segment 3 short and wide; with laterosternites on abdominal segments 18.
</paragraph>
</subSubSection>
<subSubSection id="C32734F8BE12FF82FF6C569E2656EC2A" pageId="62" pageNumber="63" type="etymology">
<paragraph id="8B826773BE12FF82FF6C569E2656EC2A" blockId="62.[151,1436,152,658]" pageId="62" pageNumber="63">
<emphasis id="B949BB61BE12FF82FF6C569E2374EFC1" bold="true" box="[199,339,471,497]" pageId="62" pageNumber="63">Etymology.</emphasis>
Although named in honour of a male the generic name is a feminine noun latinised from the English name Lloyd (the most appropriate masculine derivative was already in use).
</paragraph>
</subSubSection>
<subSubSection id="C32734F8BE12FF82FF6C556E2196ECA2" pageId="62" pageNumber="63" type="discussion">
<paragraph id="8B826773BE12FF82FF6C556E2196ECA2" blockId="62.[151,1436,152,658]" pageId="62" pageNumber="63">
<emphasis id="B949BB61BE12FF82FF6C556E2367EC71" bold="true" box="[199,320,551,577]" pageId="62" pageNumber="63">Remarks.</emphasis>
I have much pleasure in naming this genus for James Lloyd in appreciation not only of a lifetime dedicated to the pursuit of the elusive firefly and its problems, but also in recognition of his and Dorothys considerable support and kindnesses to me in my own pursuit.
</paragraph>
</subSubSection>
</treatment>
</document>