A striking new and species of tiger- (: , , ) from the Caribbean, with molecular and morphological analysis of its systematic placement Benoît Vincent, Mehrdad Hajibabaei, Rodolphe Rougerie

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Benoît Vincent, Mehrdad Hajibabaei, Rodolphe Rougerie. A striking new genus and species of tiger- moth (Lepidoptera: Erebidae, Arctiinae, Arctiini) from the Caribbean, with molecular and morpho- logical analysis of its systematic placement. Zootaxa, Magnolia Press, 2014, 3760 (2), pp.289-300. ￿10.11646/zootaxa.3760.2.8￿. ￿hal-02639378￿

HAL Id: hal-02639378 https://hal.inrae.fr/hal-02639378 Submitted on 28 May 2020

HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript ADN, 28S, phylogénie 28S, ADN, Accepted byL.Gall: 10 Dec. 2013; published: ?? Month 2014 over 3000 meters, which contributes to the high level of endemicity on the island (Huber Caribbean region. The insularity ofthis countryis associ DominicanTheRepublic is located onthe islandofHispaniola, partGreater oftheAntilles archipelago in the Introduction Motsclés: 28S. nucléaire gène du D2 région d’unerésultats analyse phylogénétique du fragment du gène tématiquenouveau dece seinla genreau sous-famille de L’habitus, sys- Laposition Dominicaine. etfigurés. décrits de République décrite sont tiinae, etfemelle mâle lesgenitalia Westindia Résumé NA, phylogeny Key words: the region gene. COI therDNA geneand D2 DNAbarcode mitochondrial of offragment 28S the of derived from phylogeny andamolecular morphology a light of comparative in isdiscussed Arctiinae within genus new from Dominican Republic. Habitus, male and female genitalia aredescribed and figured. The systematic position of the Westindia Abstract ns Cedex 2, France. E-mail: [email protected] 4 3 05 Cedex Paris F-75231 mologie), 2 1 VINCENT BENOIT placement systematic ofits analysis Arctiinae, Arctiini) from theCarib strikingA newgenus andspecies of Copyright © 2014 Magnolia Press systematic placement. ). Here wedescribe this asanew genus andspecies, andpresent morphological and molecular analysis of its discovered. Among moths collected in August 2007 wasa striking and unique member of the Arctiinae (tiger Haxaire &Schmit2001) and newgeneraofmoths (Thiaucourt 2006; Rawlins &Miller 2008)have been variety of newspecies (Rougerie & Herbin 2006; Vincent 2005a, b; 2006, 2009, 2011; Vincent Gelabert2008;Santiago-Valentin & Olmstead 2004). Within thepast decade,result asa intensive of collecting, a 1 rue Roger Rameau, 93110 Rosny sous Bois, France. E-mail: @gmail.com tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and INRA Orléans, UR633, Unité de Recherche en Zoologie Forestière, 2163 Avenue de la Pomme de Pin, CS40001 Ardon, 45075 Orléa 45075 Ardon, CS40001 Pin, de Pomme la Avenue de Forestière, 2163 Zoologie en Recherche de Unité UR633, Orléans, INRA Biodiversity Institute of Ontario, University of Guel 50 (Ento n° postale case 602, USM Evolution, et Systématique Département Naturelle, d’Histoire National Muséum Correspondant, Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in www.mapress.com Zootaxa Vincent Vincent, a new genus, is proposed for proposed is genus, new a Vincent, Arctiinae, Arctiini, République Dominicaine, nouvelle genre, nouveau espèce, Dominicaine, Arctiinae, République Arctiini, Arctiinae, Arctiini, Dominican Republic, new genus, new new new genus, species, Arctiini, Republic, Dominican Arctiinae, 0000 (0): 000 (0): 0000 , nouveau genre est proposé pour http://zoobank.org/urn:lsid:zoobank.org:pub:00000000-0000-0000-0000-00000000000 1 2 / zootaxa , MEHRDAD HAJIBABAEI , MEHRDAD – 000 / Comment citer cedocument: 10.11646/zootaxa.3760.2.8

http://dx.doi.org/10.11646/zootaxa.0000.0.0 http://dx.doi.org/10.11646/zootaxa.3760.2

ph, 579 Gordon Street, N1G 2W1, Guelph, Ontario, Canada. W. haxairei bean, wi W. tiger-moth (Lepidoptera: Erebidae,

3 haxairei Vincent, une nouvelle espèce de lasous-famille des Arc- & RODOLPHE ROUGERIE &RODOLPHE Article ated with amountainwith system peaksreachingslightly Vincent, of anewspecies Neotropical tiger-moth described est discutéesurbase la d’une étude morphologique des et mitochondrial COI utilisé comm e code-barres etde COI utilisé la ADN mitochondrial th molecula Westindia r andmorphological

4 , COI, DNA barcoding, 28S rD- Westindia ISSN ZOOTAXA edition) (print 1175-5326 ISSN

1175-5334 et al ,

COI, code-barres

. 2010; Perez- et al.et

(online edition) (online 2009 ; 2009

1 - - Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript 2 DaCosta & Weller (2005)) too is sparse to substantially assist in the selection of candidates for related genera. phylogenies for Arctiinae (Jacobson andWeller (2002); Zaspel andWeller (2006); Bendib andMinet (1998) and Pericopina and Spilosomina) made this selection challenging. Moreover, the taxon sampling used in available , , subtribes the in listed genera (217 Arctiinae the within genera of diversity great the selected forcomparisons to the newtaxon: series of both sexes in the MW, also from the region of Constanza. species), andonly the other specimens locatedwere asingle male (lacking in thecollection MNHN data) and a extensive surveys of the available literature (including the descriptions and illustrationsby Grote of Cuban Museum,Pittsburgh, Pennsylvania). description A matching thenew species described hereinnotlocated was in (ANSP; it is worth noting that many Grote types could not be located, but might be present at the Carnegie types provided by Jason Weintraub, curator of the Academy ofNatural Sciences, Philadelphia, Pennsylvania, USA mostly represented by taxadescribed from Cuba by Grote, and has also been partly examined via photographs of AntillesGreater speciessubtribesof the within the Arctiina, Phaegopterina Pericopina.remaining and The thirdis decades while conducting taxonomic research onArctiinae. These institutions hold the type material for thirdstwo Humboldt Universität, Museum of Natural History, Smithsonian Institution, Washington D.C., USA); ZMHB (Zoologisches Museum, university, Oxford Museum, (National (University UMO UK);USNM Germany); Oxford, Frankfurt-am-Main, Germany); (Rijksmuseum RNHL vanNatuurlijkeHistories, Leiden, Netherlands); SMF (Senckenberg Museum, London, MNHN(Muséum UK); national d’Histoire Nature ofNaturalHistory, Museum (American AMNH NewYork Zaspel& Weller (2006),DaCosta &Weller(2005), and Miller (1991). Institutional acronyms are asfollows: (2010) with respect to tribal delineations. Genitalic terminology follows Klots (1970), Bendib & Minet (1998), metathorax wasseparated from the mesothorax. specimen wasalso cleared using the sameprotocol. cleansing After ofthe scales and the internal soft tissues, the were madeusing adigital camera attached to aLeica MZ12 stereomicroscope. The whole body ofone male chlorazol-KOH (10%)solution,stainedusing aheated with Republic. Specimenskilled were byinjection of ammonia and stored inglassine envelopes. Genitaliacleared were vapor light (operated continuously from 0630pm to 0630 am) near the town of Constanza, in the Dominican Specimen collectingand morphological study. Materialsmethods and showed novariation in length within our ingroup except for the three included species of barcode sequences, respectively). both genes were available (GenBank accession numbers GU350477 and GU654969 for 28S and DNA COI [Hübner, 1808] wasselectedasanoutgroup, andis the closest relativearctiine to for moths which sequences of thePPCE clade of JacobsonWeller & (2002)within the Arctiinae). Thenoctuid moth The 28SWESTIND1). sequences usedwereavailable only for21species within the last four subtribes (forming available at(www.boldsystems.org) BOLD within the public dataset DS-WESTIND1 (dx.doi.org/10.5883/DS- availableof 32speciesa set including for representatives phylogeneticmarkerin theArctiinae. The taxa usedinthisstudy are listedTable in barcodes(COI) 1.DNA were geometrid moths (Abraham et al available fromanother study and because this gene proved to be informative at the tribe and subfamily level in D2B and D3Ar (Saux also amplified and sequenced the D2 expansion segment of the ribosomal large subunit (28S) using the primer pair followedthe protocols describedin Vaglia Lepidoptera campaign (see www.lepbarcoding.org fordeta Barcoding in Guelph (Ontario, Canada), along with many Neotropical tiger moths developed within the iBOL tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and

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Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in Zootaxa Molecular analyses. carefully were genera five following the Arctiinae, Neotropical of knowledge author’s first the on Drawing The first author has visited the aforementioned institutional collections, among others, over the past two TreatmentArctiinaea subfamily ofthe as ofErebidaefollows Zahiri Phylogeneticanalysis sequences. of DNA 0000 (0) © 2014 Berlin, Germany). Germany). Berlin, et al., Tissue samples were processed via DNAba

Magnolia Press 2004). We selected this fragment because sequences of several taxa werealready Comment citer cedocument: 10.11646/zootaxa.3760.2.8 ., 2001; Young, 2006), although it hasnever previously been used asa

http://dx.doi.org/10.11646/zootaxa.3760.2 , Elysius,Pseudamastus, , et al.

(2008). In addition to the 658 bp fragment of the COI gene, we Males and females of the new taxon were collectedtheMales newtaxonfemales and weremercury of at All sequences All were aligned manually;28S sequences rDNA ils). DNAextraction, amplificationils). PCR andsequencing , USA); BMNH (British of Natural Museum History, (British BMNH , USA); black, andmountedin Euparal.Illustrations onslides of allwithinthe Arctiinae; subtribesof records all are lle, Paris, France); MW (Museum Witt, Munich, rcoding at the Canadian Centre for DNA for Centre the Canadian rcoding at et al. et

(2012) andLafontaine &Schmidt and Opharus Helicoverpa armigera Helicoverpa armigera Gorgonidia , all of which of , all

VINCENT . Note that that . Note ET AL. ET Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript NEW ARCTIINAE GENUS FROM DOMINICAN REPUBLIC ARCTIINAE NEW DOMINICAN GENUS FROM tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in Comment citer cedocument: 10.11646/zootaxa.3760.2.8

http://dx.doi.org/10.11646/zootaxa.3760.2

Zootaxa 0000 (0) 2014 © Magnolia Press

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3 Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript follows: COI nt1-2 / COI nt3 / 28S. / nt3 /COI nt1-2 COI follows: dataset waspartitioned bygene (COI / 28S) or alternatively considering COI codon positions in apartition as separating two branches: one dorsal, with avery pointed tip, the other ventral ending with two digitiform processes uncus. Scaphiumsclerotized, concave and tongue-shaped (Fig. 5). Valvae shortand symmetrical with alargenotch and elongated. Tuba analis twice aswidethe tegumen atits base and extending beyond the apical end of the covered with long scales. closed ventrally. Metepisternal tympanal organ present and well developed. Patagia and tegulae developed and spurs, respectively; the spurs smooth, medium-sized and all equal in length. Tympanal Pocket IVpresent and apically acute, less than half the length of the tibia. Meso- and metathoracic legs with one and two pairs of tibial ventrally.This tuft disappears progressively present. Antennae with 87flagellomeres, each bearing apair of dorsal setae and adense of tuft golden bristles Proboscislongerthan thorax medium-sized with sensillae upward. Second segment thinner, slightly curved and longer than the first. Third segment short and rounded. 4 a wrinkled scaphium and valves characterizedbifid by adeep notch. following combination of characters is diagnostic: hindwing with M2 strongly stalked with M3; male genitalia with streaks, white hindwings andwingspan a greater than 50mmisunique amongthe subfamily. Structurally the pattern. Thecombination anorange of forewing withivory-white two transverse bandsand three longitudinalgrey assessed below. other related species arediscovered in the future. The predict which onesof the unique characters described below maybepotentially diagnostic at the genus level if diagnostic features of this new genus. However, because members of which it would share deriveddiagnostic characters. We describe herewhat weconsider to bethe main Diagnosis. Westindia haxairei species: Type Figs. 1–5 Westindia Systematic Part inferringconfidence values through bootstrapping until the stopping criteria are met (Pattengale www.phylo.org)at theSan Diego Supercomputer Center (Miller al et RAxML v.7.2.7 (Stamatakis, 2006; Stamatakis node wasevaluated through a 1000 bootstrap (BS) re-sampling. ML phylogenetic analyzes were performed using using the Close Neighbor Interchange (CNI) algorithm, with search level 2 and 30 initial trees. The stability of each parsimony (MP) methods.MP analyses were carriedoutin 5(Tamura MEGA missingdata forin 28S 11 taxa. Sequences were analyzedusing maximumlikelihood (ML)and maximum combined; for dataset (2)weanalyzed COIsequences separately, aswell as acombination 28S ofCOI + with Arctiinae subtribes (PPCECallimorphina + and Arctiina). For dataset (1),weanalyzed both genesseparately and by 11species for which only COI sequencesareavailable –this dataset comprises 32taxa intotal,representing all for whichsequence were dataavailable for both genes (COI, 28S; seeTable 1); and the (2) samedataset augmented taxon + 20 species in 14 genera within subtribes Pericopina, Phaegopterina, , and (PPCE) the presenceof a1bpinsertion in position 300of the amplicon.analyses Our consideredtwo datasets: (1) the new sequences straightforwardand unambiguous. The sequence for havea 4bpinsertion (TTTA) position in 184ofthe fragment sequenced, making the alignmentwiththe restof the and prominent without distinguishablemarkings. stalked with M3; Sc+R1 very thin where it reaches the costal margin. contact withM2;are separate. CuA1andM3 tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and

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Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in Zootaxa Description. Vincent,genus new It proved impossible to associatethenew species unequivocallyan to already existinggenus with the 0000 (0) © 2014

posteriorly. Westindia Male (Fig. 1). Head Forewing

n.gen.can bedistinguishedexternally from all otherknown Arctiinae byits wing Eighth sternum trapeziform andas long asseventh sternum. Uncus and tegumen narrow

Magnolia Press Vincent, monotypy by species new Comment citer cedocument: 10.11646/zootaxa.3760.2.8 Male terminalia Male . Elongate, narrow with apex acute. Venation is of the quadrifid type with M3 in in M3 with type Venationacute. quadrifid apex the with isof narrow . Elongate,

. Labial palpi thin and strongly curved upward. First segment strongly curved http://dx.doi.org/10.11646/zootaxa.3760.2

in the apical third. third. apical the in Hindwing et al (Figs. 2-3 ; 5-7). Eighth tergum rectangular, slightly sclerotized ., 2008) implemented onthe CIPRES Portal v. 3.1(http:// higher placement ofthe new genus within Phaegopterina is . Reduced andnarrowed; itsapex acute. VeinM2 strongly we establish a monospecific genus, it is impossible to at its tip. Eyes large without interfacetal setae. Ocellus Helicoverpa armigera only differed inlength by Thorax Abdomen ., 2009), using the GTRCAT model and . Prothoracic legs with epiphysis foliate, foliate, epiphysis with legs . Prothoracic

. Large andtapered, entirely orange et al et ., 2011) including all sitesand et al.,et

VINCENT 2010). The ET AL. ET Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript NEW ARCTIINAE GENUS FROM DOMINICAN REPUBLIC ARCTIINAE NEW DOMINICAN GENUS FROM Westphal leg., in coll W. Speidel (München). vii.1999; R. Marx leg., in Museum Witt (München). 1male Dominican Republic, Constanza,1200m, 21.XI.1995, 2 males and 2 females, Dominican Republic, Monseñor Noue males and2 females, Dominican Republic, Monseñor Nouel, Cordillera Central, Casavito, 1050-1280m, vii.1999 ; 70°41.677’; 05.viii.2007; attracted to UV light, J.Haxaire and O.Paquit Dominican Republic, La Vega department, Road Valle Nuevo to Constanza Km 10, alt. 2015m. N 18°50.123’ W D.C.); 2 males, same data as the holotype but collected on 08.viii.2007, in collection of the first author; 2 males, in the Natural History Museum (NHM, London) and the National Museum of Natural History (USNM, Washington BEVI0555/KF930922). Deposited in collections of the first author except for two specimens deposited respectively holotype,barcode2 males withsequence a available (SampleID/GenBankBEVI0553/KF930928,Acc.: Muséum National d’Histoire Natu National d’Histoire Muséum as the holotype; genital prep. BV 373; Barcode SampleID/GenBank Acc.: BEVI0554/KF930920. Deposited in the KF930933: Deposited in the Muséum National d’ light, J. Haxaire and O. Paquit leg pK 10,Reserva Scientifica Ebano Verde, alt. 1360m.N 19°10.729’ W70°30.988’; 15.viii.2007; attractedto UV Type material Figs. 1–6 Westindia haxairei 1.FIGURES and thin (5.4 mm), with vesica 5mm long and lacking cornuti (Fig 3). (Fig 6). Vinculum thin with two membranous lobes at the articulation with the tegument (Fig 7). Aedeagus long tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in Etymology. Westindia haxairei . Holotype The generic name is derived from the West Indies (=Caribbean) Vincent Vincent ; male: Dominican Republic, Monseñor Nouel department, Road El Blanco to Constanza new species n. sp. 1: Holotype, dorsal view. Comment citer cedocument: 10.11646/zootaxa.3760.2.8 relle (MNHN, Paris). Paratypes Paris). (MNHN, relle

.; genital prep. BV 372; Barcode SampleID/GenBank Acc.: BEVI0552/ http://dx.doi.org/10.11646/zootaxa.3760.2

Histoire Naturelle (MNHN, Paris). Allotype, Paris). Naturelle(MNHN, Histoire l, Cordillera Central, La Palma deConstanza, 600m, : 5malesand: 4females,same dataasthe

Zootaxa leg. 0000 (0) 2014 © Magnolia Press , in collection of the firstauthor. 2

female: same data data same female:

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5 Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript 6 Westindia 2–4. FIGURES haxairei tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and

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Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in Zootaxa 0000 (0) © 2014

Magnolia Press Comment citer cedocument: 10.11646/zootaxa.3760.2.8 n. sp. 2: Holotype, genitalia ; 3: 4: genitalia.Holotype, female, aedeagus; Paratype

http://dx.doi.org/10.11646/zootaxa.3760.2

VINCENT ET AL. ET Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript NEW ARCTIINAE GENUS FROM DOMINICAN REPUBLIC ARCTIINAE NEW DOMINICAN GENUS FROM valvae (va). (tg), andtegumen (vin) vinculum between (ml) lobe Holotype, genitalia, membranous 7 genitalia, valvae; Holotype, ;6 (ta) FIGURES 5–7. tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in Westindia haxairei Comment citer cedocument: 10.11646/zootaxa.3760.2.8 n. sp.

5 Holotype, genitalia, lateral view of the uncus (un), scaphium (sc) and tuba analis tuba and (sc) scaphium (un), uncus the of view lateral Holotype, genitalia, 5 http://dx.doi.org/10.11646/zootaxa.3760.2

Zootaxa 0000 (0) 2014 © Magnolia Press

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7 Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript contrasted, the postmedian interruptedand the costathree at streaksgrey the not visible. by a thin yellowish area.Ventrum—Colors more vivid th bands, with three longitudinal grey streaks, slightly converging basally. The central grey coloration is surrounded apex. The costa is marked by brownish scales. The central area of the forewing, between the two ivory-white itfused, andbecomeswhen wider reaching theis fringe whichanal white toward angle the andblack toward the middle; thissecond section of the bandis strongly narrowed initsmiddle, where the black lines becomealmost starts fromthe costa parallel to the antemedian band, then turns to reach the external margin ofthe wing in its white bands bordered bythick blacklines. Antemedian band narrower in its central area; the postmedian band red hue, the proximal two thirds with opalescent scattered scales. The forewing is crossed by two transversal ivory- claw toothed. ventrally. dorsally, line orange Tarsal tibia longitudinal white a single with tarsi and Fore-tibia white. spurs tibial middle and hind legs black with dorsal surface streaked with two white longitudinal lines; area around the hind- wholethorax. headandVentral andsurface dorsalthe samecolor. orange. Femurofall legs Tibiatarsi andofthe result of the presence of long orange scales tipped with yellow, giving an orange powdered with yellow hue to the palpi orange powdered with yellow. white scale patches on the distal end of each segment. thin whiteline, except forthe apical twenty flagellomeres whitedorsally, and forthe proximal ten segments with pedicel narrow. Antennae monofiliform with 87 flagellomeres; dorsal and ventral surfaces black, delimited by a 8 new genus All the phylogenetic hypotheses generated from the combin Discussion Westindia species was not collected during previous sampling in the same area during April/May 2004, suggesting that cordilleraDominicanin the Republic.the specimens Allcollected to known uswere July August, and inthisand continuous ivory-white coloration. specimens display a1mminterruption of this band whereas in others the contraction is almostabsent allowing a bristles. Dorsal pheromone glands with apair of long unbranched tubes. into alarge wrinkled and rounded bulla seminalis. Papillae anales trapeziform, uniformly covered withlong signum. There is a protuberance close to the insertion of the ductus bursae. The ductus seminalis is short, opening ribbon shaped, curved near the insertion of the corpus bursae which shape is pyriform, wrinkled and without Pseudopapillae entirely joined and covered with short bristles. Ductus bursae uniformly sclerotized, elongated, tergiteA6,slightly sclerotized andrectangular. Sternitelarge A7 witha strong V-shaped posteriorindentation. of golden bristles on the ventral face of the flagellomeres. single lobe lacking cornuti (Fig. 3). Female. excavation on the ventral margin. Phallus long, slightly curved. Caecum penis short and rounded. Vesica with a atend.some bristle the Vinculumshort pointed with a saccus.Juxta slightly sclerotizedaU-shapedwith Ends of the ventral branch with two digitiform processes: one broad and rounded, the other fine and curved with large, symmetrical,bifid with a deep notch (Fig. 6). The dorsal branch is long and curved with a very pointed tip. sclerotized laterally.Scaphium sclerotized, concave and tongue-shaped (Fig. 5).Transtillasubrectangular. Valves Uncus with a pointed apex slightly curved ventrally. It is covered by long bristles laterally. Anal tube slightly Ventral surfaceidentical in color. marked with black scales, forming aregularpunctuation along the pleurites. Sternites uniformly vivid orange. reddish orange scales. The posterior tergites with shorter scales anda slightly stronger coloration. Each spiracle is its base and becoming grey toward its apex. Discoidal lunula grey. Abdomen indentation toward the tip ofthe discal cell. Ventrum -Same coloration than dorsal face, but for the costa orange at and semihyaline.anal slightlygrey areais asimilarlycostaliswith Thegrey;colored the markedin area tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and

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Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in Zootaxa Distribution, phenology and biology. W. haxairei Etymology. Variation. Description is amid-summer flier. The early stages and foodplants are unknown. 0000 (0) © 2014 Westindia within the subtribe Phaegopterina, in agreement with our morphological observations. Forewing The postmedian bandexhibits somevariations inthe extentits ofcentral contraction.Some . Male The new species is dedicated to our colleague Jean Haxaire

1) (Fig. . Dorsum-Ground color orange the with central areamorestrongly colored withavivid

Magnolia Press Comment citer cedocument: 10.11646/zootaxa.3760.2.8 Wingspan 57–65 mm; forewing length 26–30 mm. Male genitalia

Thorax http://dx.doi.org/10.11646/zootaxa.3760.2 The female is identical to the male but forthe absence ofadense tuft

. All thorax, legs excepted, is orange powdered with yellow, with the powdered as orange is excepted, legs thorax, . All (Figs. 2–3; 5–6) Tegumen and Uncus narrow and elongated. seemsrestricted to middle elevation localities of the central

The apical third with fewbristles. Frons, head and labial an on the dorsal face.The antemedian band poorlyis Female genitalia ed datasets consistently support the placement of the . . Tergites A1–4 covered by long vivid (Fig. 4).Tergite A7twice aswide as

Head Hindwing Antennal scape spherical, Dorsum - White - White Dorsum

VINCENT ET AL. ET Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript NEW ARCTIINAE GENUS FROM DOMINICAN REPUBLIC ARCTIINAE NEW DOMINICAN GENUS FROM below branches on the MP tree for shared nodes. treefor onthe below branches MP Phaegopterina clade is highlighted in black and an arrow points at points arrow an and black in ishighlighted clade Phaegopterina the which in tree ML best-score the of topology the represents the box upper-left in tree The branches. above given are 50% both COI and 28S genes for 21 ingroup taxa (outgroup 8. FIGURE resolution ofthelimited phylogenein COI sequences, results not shown). Overall, and as could be expected, the limited taxon and gene sampling results supported asmonophyletic by COI sequences alone of (BS 83 noticeable exception of subtribe Arctiina forwhich no 28S sequences were available and which is consistently gene analyses (results not shown) produced generally unre placed consistently still but 9) (Fig. of theextended dataset (including all tribesin Arctiinae) did not recovera monophyletic Phaegopterina subtribe of 65 and 55, respectively) of the reduced dataset with representatives of the PPCE clade only (Fig. 8). The analysis However, the monophyly of the subtribe wasonly poorly supported after ML and MP analyzes (bootstrap support Species of ornamentation;male genitalia the figured byToulgoët(1985) quite are different, notably inthe valves and uncus. Dominica and Martinique, respectively, but the forewings are uniformly brown or yellowish, without Toulgoët, 1985, has two subspecies transverse bands, and the hind wings have yellowish to pinkish coloration white).(never The genus have ayellowish abdomen with black transverse bands, but the forewings are brown do and not have bright Westindia to pattern wing haxairei prospected byentomologists for several centuries. In fact, no species known to usbears any close similarity in Ctenuchina (Fig.wellas the 9), monophyly as Ctenuchina of + Euchromiina. Thepositiongenus of genesthese s worthnotingthat is attention, it little statisticalmost branches. support for Considering th that by Jacobson &Weller (2002). 1854, asinferred from our analyses, confirms its outlier st tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in The discovery of this new arctiine moth in the Dominican Republic came asasurprise in aregion regularly Consensus tree of the four most parsimonious trees obtained after the parsimony analysis of dataset (1) combining combining (1) dataset of analysis parsimony the after obtained trees parsimonious most four the of tree Consensus Halysidota Hübner, [1819], of which several occurin the Caribbean ( Comment citer cedocument: 10.11646/zootaxa.3760.2.8 . Both . Both P. alsa alsa Westindia amongof haxairei membersthis subtribe inall analyses.Separate

http://dx.doi.org/10.11646/zootaxa.3760.2 tic relationships within thesubfam relationships within tic Ammalo helopsAmmalo

(Druce,1890) and Helicoverpa armigera s Arctiina, Pericopina andupport themonophyly Pericopina ofsubtribes Arctiina, e phylogeny of these moths has to date received relatively (Cramer, and [1776]) atus within the subtribe Phaegopterina, asalready noted solved trees with poorly supported groupings, with the Westindia haxairei after ML analysis with the nt12/nt3partition of the P. alsa lalannei

Zootaxa , not values shown). Bootstrap over confidence 0000 (0) 2014 © Magnolia Press ily Arctiinae, with generally poor generally with ily Arctiinae,

n. sp.. ML bootstrap values are given H. cinctipes Elysius cingulata Toulgoët, 1985 described from Grote, [1866], (Walker, 1856)

Pseudamastus Walker, Walker, H. ata

·

9 Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript 10 50% confidence givenBootstrap over node.values are ateach partitions. distinct as the genes with two (2) dataset extended the theof analysis from resulting tree ML 9.FIGURE Best-score the outer margin of the forewing (see Vincent 2012). (basal, antemedial andpostmedial) onthe forewings andagreypatchextending fromthe apexofthe discal cell to and patterns reminiscentof yellowish grey-white subterminal band separates them. Lastly, species of Gorgonidia Dyar, 1898 have wing colors body and forewings and acomparable wingspan, but the 1895), describedfrom Mexico, is somewhat similar to and these processesareabsent in Watson (1980) have a vinculum with digitate, non-setose processes on each side at the junction with the tegument, Halisidota masoni of those to shape Watson, 1980, H.insularis possesses two of the three character conditions proposed by Jacobson & Weller (2002) to define the subtribe a closest relative, although our analyses highlighted characters useful at higher taxonomic levels. tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and

Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in · The uniqueand distinctivefeatures of Westindia haxairei

Zootaxa 0000 (0) ©0000 2014 (0) Westindia ; ; normale genitalia presence mark) abdominal of areniform color; of (Rothschild, 1909), and H.elota , but none has similar color and ornamentation (e.g., yellow to brown forewings, except W. haxairei

Comment citer cedocument: Magnolia Press 10.11646/zootaxa.3760.2.8 W. haxairei.

http://dx.doi.org/10.11646/zootaxa.3760.2 , but theseall, butdiffer by the presence ofthree incompleteyellow bands

Among the Central American species, W. haxairei presence of a black spot on each tegulae and alarge (Möschler,and 1886)),size haveforewings similarin make it difficult topropose a knownarctiine genus as in having anorange background color of the

Halysidota masoni Halysidota Halysidota

VINCENT figured by figured Westindia (Schaus, (Schaus, ET AL. ET Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript NEW ARCTIINAE GENUS FROM DOMINICAN REPUBLIC ARCTIINAE NEW DOMINICAN GENUS FROM Bendib, A. (1998) D., Abraham, Ryrholm, N., Wittzell, H., Holloway, J.D., Scoble, M.J. & Löfstedt, PhylogenyC. of Molecular (2001) the Literature cited Rica. Janzen andWinnie Hallwachs on the lepidopteran diversity in Areade Conservacion Guanacaste in Costa (ACG) phylogenetic analyses weregenerated as part of a joint research project between Mehrdad Hajibabaei, Daniel H. NSERC, Genome Canada and the Ontario Genomics Institute. Sequences of Costa Rican taxa used for the barcodingDNA hosted by the Biodiversity Institute of Ontario, University of Guelph (Canada), through funding by observation and interpretation of some characters. barcode DNA data were generated at the Canadian Centre for offered the material for study, and to Joël Minet (Muséum The first author is grateful to Jean Haxaire and Odile Paquit who collected the specimens of this new genus and Acknowledgements sampling.and character Becker, 2011). In the current state ofknowledge, amore precise placement of Westindia must await further taxon after morphological comparison with two species occurring in Cuba (C. gortynoides Grote, 1866 and C. alayorum between Westindia haxairei and the genus , asapparent in Fig. 9 (with no statistical support), wasruled out uninformative at this stage with respect to phylogenetic inferences within the tribe. The possible relationship glands with fairly wide opening”), a condition found in many Phaegopterina (Bendib & Minet, 1998) and (1998)to further infer the relationships within this lineage, is of type “1” in Elysius Praeamastus, Halysidota, placement of lower angle of the discal cell) and the pseudo-papillae of female genitalia fused throughout their length support the group or Phaegopterina. In addition, the quadrifid venation in hindwings (vein M2present and branching from the shapedbulla seminaliscomparableis tothat described byBendib (1998)as adiagnostic featureofthe genus prothoracicof glands)not was observed andtheirabsence mayrepresentsecondary a inloss the as phaegopterine Weller (2002) as“vein Sc+R1 [...] present but does not r much attenuated toward the hindwing margin (which weconsider fitting the condition described by Jacobson & Phaegopterina: (1) the presenceon the tympanumofaventrally closed pocket IV; and(2) the hindwing vein Sc+R1 Lafontaine, Lafontaine, J.D. & Schmidt, B.C.Annotated check (2010) listof DaCosta, M.A. & Weller,M.A. & DaCosta, Arctiidae:Arctiinae) (Lepidoptera: Callimorphini classification the of and S.J. Phylogeny (2005) significance. phylogenetic and Evolution (Lepidoptera). Arctiidae in glands pheromone Female (1998) J. Minet, A & Bendib, Huber, B.A., Fischer, N.& Astrin, J.J. (2010) ofHigh level endemism in Haiti’s last forests: arevision remaining of Haxaire,J. &Schmit, P. (2001) Un nouveau sphingide de Republique Dominicaine: Klots, A.B. (1970) Lepidoptera. In &Weller, N.L. (2002) S.J. Jacobson, Hebert, P.D.N., Dewaard, J.R. & Landry, J.F. (2010) DNA barcodes for 1/1000 of the kingdom. kingdom. animal ofthe P.D.N., &Landry, 1/1000 Hebert, for J.R. J.F. Dewaard, barcodes DNA (2010) tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in Zootaxa femelles des Lepidoptera). (Geometroidea: inSubfamilies Geometridae of Mexico. pp. 115–130. Comptes rendus de l'Académie des sciences. Série 3, Sciences de la vie la de Sciences 3, Série sciences. des l'Académie de rendus Comptes (Araneae: Pholcidae) on Hispaniola, using morphology and molecules. molecules. and using on morphology Pholcidae) Hispaniola, (Araneae: http://dx.doi.org/10.1098/rsbl.2009.0848 359–362. Sphingidae). http://dx.doi.org/10.3897/zookeys.40.414 Adults http://dx.doi.org/10.1111/j.1096-3642.2009.00559.x 244–299. . Thomas Say Monograph Series, Entomological Society of America, 98 pp. Say Series,Entomological America, . Thomas Society of Monograph , 1025, 1–94.

Zookeys Westindia . Thèse de doctorat du Muséum National d’Histoire Naturelle, spécialité Entomologie, 300 300 Entomologie, spécialité pp. . National d’Histoire deNaturelle, duMuséum Thèse doctorat Lambillionea Contribution à l’étude des Arctiidae (Lepidoptera : Noctuidea): phylogénie et morphologie de l’abdomen l’abdomen de et morphologie des ArctiidaeContribution l’étude phylogénie à :Noctuidea): (Lepidoptera RegoBarros, 1956(Jacobson& Weller, 2002).The presence a largely of rounded diverticule- , 40, 1–239. in a lineagein defined byBendib(1998: 167) that includes thegenera , 101 (4), 605–608. 605–608. (4), , 101 Comment citer cedocument: : Tuxen, S.L. (Ed.), (Ed.), Tuxen, : S.L. 10.11646/zootaxa.3760.2.8

and A cladistic study of the Arctiidae by of (Lepidoptera) using characters Immatures and

http://dx.doi.org/10.11646/zootaxa.3760.2 Amastus

. The configuration of dorsal saccular glands, proposed by Bendid . Copenhagen, Munksgaard, of Taxonomist’s genitalia of glossary each the hindwing margin”). The third character (presence Molecular Phylogenetics andMolecular Evolution Phylogenetics the (Insecta, Lepidoptera) of North America north north America North of Lepidoptera) (Insecta, Noctuoidea the national d’Histoire Naturelle, Paris) for his help in the helpthe his in for Paris) Naturelle, d’Histoire national , 321 (12), 1107–1114.

Zootaxa Zoological Journal of the Linnean Society Westindia haxairei 0000 (0) 2014 © Magnolia Press

Cauthetia carsusi , 20, 65–77. , 20, (“a pair of elongate Biology Letters Biology

(Lepidoptera (Lepidoptera , Hemihyalea, Halysidota Modisimus , 158, 158, ,

·

, 6, , 6, 11 . Version postprint Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript Manuscrit d’auteur / Author Manuscript 12 Haiti) and Republic (Dominican Hispaniola of (2008) D.E. Perez-Gelabert, bibliography. and checklist : A (Lepidoptera Notodontidae the Miller, of and classification Cladistics (1991) J.S. adult and larval based on : Noctuoidea) Vincent, B. (2005a) Description de deux nouvelles espèces d’Arctiidae de République Dominicaine (Arctiidae, (Arctiidae, Dominicaine République de d’Arctiidae espèces nouvelles deux Vincent, de Description (2005a) B. Vaglia, T., Haxaire, J., Kitching, I.J., Meusnier, I. & Rougerie, R. (2008) Morphology and DNA barcoding reveal three cryptic Toulgoët, H. de (1985) Description d’une nouvelle sous-espèce d’Arctiide de la Martinique avec création d’un genre nouveau. genre nouveau. création d’un avecToulgoët, laMartinique de d’Arctiide sous-espèce nouvelle d’une Description (1985) H.de Stamatakis, A., Hoover,P. &Rougemont,J.(2008) Fast A Bootstrapping Algorithm for the RAxMLWeb-Servers. Pattengale,N.D., Alipour, M., Bininda-Edmonds, O.R.P., Moret, B.M.E. &Stamatakis, A. (2010) How many bootstrap Miller, M.A., Holder,M.T., Vos, R.,Midford,P.E., Liebowitz,T., Chan,Hoover, L., P. &Warnow,T. (2009)The CIPRES Stamatakis, A. (2006) RAxML-VI-HPC:Maximum Likelihood-based Phylogenetic Analyses with Thousands of Taxa of and Thousands with Analyses Phylogenetic Likelihood-based RAxML-VI-HPC:Maximum (2006) A. Stamatakis, Saux, C.,Fisher, B.L.& Spicer, G.S. and sequences 28SrDNA bynuclear asinferred phylogeny ant (2004) Dracula Santiago-Valentin, knowledge current to introduction plants: Caribbean of biogeography Historical R.(2004) Olmstead, E.& Rougerie, R & Herbin, D. (2006) Rawlins, J.E. & Miller, J.S. Dioptine (2008) of region: moths two the Descriptionof with new caribbean notes on genera the Vincent, B.(2005b)Description de trois nouveaux Thiaucourt, P.Thiaucourt, (2006) Tamura, K., Peterson, D., N.,Stecher, Peterson, G., Nei,M.& Kumar, S.(2011) Evolutionary Genetics Molecular MEGA5: Watson, A (1980) A revision of the Vincent, B.(2012)A review of thegenus new two of (2011) Vincent, Description B. Vincent, B.(2009) Description de deux nouvelles espècesappartenant au genre dugenre espèce d’une deuxième Description (2006) B. Vincent, Young, C.J. (2006) Molecular relationships of the Australian Ennominae (Lepidoptera: Geometridae) and implications for the the for implications and Geometridae) (Lepidoptera: Young, Ennominae Australian the of relationships C.J. Molecular (2006) Vincent, B, Laguerre, M & Rougerie, R (2009) Contribution à la connaissance du genre genre connaissance du la à Contribution (2009) RVincent, Rougerie, & M B,Laguerre, Zaspel, J.M. & J.M.Zaspel, & Weller, Review generic S.J. tiger of limits genera (2006) the of moth Zahiri,R., Holloway, J.D., Kitching, I.J., Lafontaine, J.D., Mutanen, M. &Wahlberg, N. (2012) Molecularphylogenetics of tiger-moth (Lepidoptera: Erebidae,Arctiinae, Arctiini) fromtheCaribbean, withmolecular and

Vincent, B.,Hajibabaei ,M.,Rougerie, R.(2014).A strikingnew genus andspecies of morphological analysis of its systematic placement. Zootaxa, 3760 (2), 289-300. DOI : Zootaxa, 2014,3760(2), 289-300 Version définitive dumanuscrit / publié dans Final version of the manuscript published :in · Zootaxa http://dx.doi.org/10.1089/cmb.2009.0179 morphology. Phaegopterinae). Phaegopterinae). species within the the within species (18 4 (1), 1–8. Biology http://dx.doi.org/10.1093/bioinformatics/btl446 are necessary? replicates [Online]. AvailablePortal from: http://www.phylo.org 24August 2011) (accessed Mixed Models. http://dx.doi.org/10.1016/j.ympev.2004.06.017 457–468. Amblyoponinae). Formicidae: ant (Hymenoptera: systematics for implications http://dx.doi.org/10.2307/4135610 and possibilities a perspective. from phylogenetic Hemileucinae). (Lepidoptera:Saturniidae, http://dx.doi.org/10.2992/0097-4463(2008)76[203:dmotcr]2.0.co;2 biology and biogeography (Lepidoptera: Notodontidae: Dioptinae). Phaegopterinae). Phaegopterinae). http://dx.doi.org/10.1093/molbev/msr121 Evolution Analysis using Maximum Likelihood, Evolutionary Distance, and Maximum Parsimony Methods. Bulletin theBulletin of British (Entomology) Museum (Natural History). France Arctiidae). ADN (Lepidoptera codes barres des Apport espèces. nouvelles deux Arctiini). Arctiinae, Erebidae, http://dx.doi.org/10.3897%2Fzookeys.75.811 75, 69–77. Arctiidae). Arctiidae). phylogeny of the Geometridae from molecular and morphological data. data. morphological and molecular from Geometridae the of phylogeny Schäffer (Lepidoptera: Arctiidae:Arctiinae) and their biogeography their and Arctiidae:Arctiinae) Schäffer (Lepidoptera: http://dx.doi.org/10.1111/j.1365-3113.2011.00607.x Systematic Entomology Noctuoidea). (Lepidoptera, Erebidae

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Bulletin de la Société Entomologique de France de Entomologique Société la de Bulletin L’entomologiste Bulletin of Bulletin the Museum American Naturalof History Bioinfomatics Antillisa Lépidoptères Lépidoptères Xylophanes neoptolemus Xylophanes Journal of Computational Biology Computational of Journal gen. n., définition et notes sur les espèces appartenant à ce genre (Notodontidae). (Notodontidae). ce genre à appartenant lesespèces notes sur définition et n., gen. , 65 (1), 47–53.

Comment citer cedocument: Magnolia Press , 22, 2688–2690., 22, 10.11646/zootaxa.3760.2.8 , 3 (1),, 3 1–8. , 3 (2),, 3 9–18. The European Entomologist Halysidota tesselaris Hispaniodirphia lemaireiana Nouvelle Revue d’Entomologie Nouvelle Revue

http://dx.doi.org/10.11646/zootaxa.3760.2 Arctiinae). (Arctiidae, theRepublic Dominican Harrisfrom Gorgonidia Zootaxa and

loelia Taxon , 1204, 53–59. Dyar, 1898, with the description of one new species (Lepidoptera (Lepidoptera species new one Dyar,of description the 1898, with : species-groups (Lepidoptera: Sphingidae). Zootaxa Sphingidae). (Lepidoptera: species-groups species-group ( species-group Harrisde (Arctiidae, République Dominicaine Lophocampa , 53, 299–319. , 111 (2), 223–226. , 4 (1),, 4 33–54. Ferguson de République Dominicaine (Lepidoptera, Caribarctia (Lepidoptera, Dominicaine Ferguson de République 17, 337–354. , 37(1),102–124. Suppl. 40(1),1–65. Antilles n. sp., Greater anew from the saturniid , 204, 1–230. . , (N.S.) 2 (1), 5–9.

Annals Annals Carnegieof Museum Zootaxa

Halysidota Zootaxa

, 1159, 1–68. Molecular Phylogenetics and Evolution Molecular Phylogenetics

, 1264, 1–147. Lophocampa sensu stricto) (Lepidoptera: Arctiidae). Arctiidae). stricto) sensu (Lepidoptera: Bulletin de la Société Entomologique de de Entomologique Société la de Bulletin

Opharus Walker and Harris, 1841(Lepidoptera Walker avec de description , 76 (4), 203–225. Molecular Biology and Holomelina

, 1923, 18–36. 1923, , VINCENT Lépidoptères Systematic Herrich- Zookeys

ET AL. ET 33, , ,