PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3519, 17 pp., 5 figures July 31, 2006 New Data on Miocene Butterflies in Dominican Amber (Lepidoptera: Riodinidae and Nymphalidae) with the Description of a New Nymphalid ENRIQUE PEN˜ALVER1 AND DAVID A. GRIMALDI2 ABSTRACT Anew,virtuallycompleteandwell-preservedfemalespecimenofVoltiniadrambaHall,Robbins, and Harvey, 2004 (Lepidoptera: Riodinidae) provides new data on this fossil species, and a new fossil species of the Recent genus of Nymphalidae Dynamine Hu¨bner, 1819 (Lepidoptera: Nymphalidae) is described as Dynamine alexae n.sp., on the basis of a male specimen. The two species are preserved in Miocene amber from the Dominican Republic. Dynamine alexae n.sp. represents the first adult nymphalid butterfly found as a fossil in amber. The four taxa of butterflies found up to the present in Dominican amber indicate post-Miocene extinctions in Hispaniola,probablycausedbyinsularization.ThebutterfliesfoundinDominicanamberdonot support a hypothesis of a Gondwanan origin for many butterfly tribes and subfamilies as previously proposed; we conclude that this hypothesis is implausible based on the age of the butterflies as inferred from the fossilrecord. Some palaeoecologic andtaphonomic questionsare discussed. INTRODUCTION amberpreservessoftinternaltissues,including cells, organelles, and even endosymbiotic Amberpreservesdelicatearthropods,main- spirochetes and protists (Grimaldi, 1996; ly insects, with high fidelity, including micro- Grimaldi et al., 1994; Henwood 1992a, scopic features like setae and sensilla on the 1992b; Wier et al., 2002). externalcuticleaswellaswingandothercolor The most diverse organisms in amber are patterns (Grimaldi, 1996). In some cases insects, particularly of the orders Diptera, 1DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory([email protected]). 2DivisionofInvertebrateZoology,AmericanMuseumofNaturalHistory([email protected]). CopyrightEAmericanMuseumofNaturalHistory2006 ISSN0003-0082 2 AMERICAN MUSEUMNOVITATES NO. 3519 Hymenoptera, and Coleoptera. Lepidoptera tribal groupings; two of them—Mesosemiini specimens tend to be less common and even andEurybiini—possessingfiveforewingradial rareinamber,theprincipalrecordbeingadult veins; the other six possess four forewing moths.Thefossilrecordofadultbutterfliesin radialveins(Hall,2003).TheRiodinidaefossil the world, from the Paleocene to Miocene, record is comprised only of a caterpillar contains approximately 50 specimens known, specimen (DeVries and Poinar, 1997; see the comprising about 33 named species in all discussionaboutotherputativeriodinidfossils livingfamilies,butonlysevenofthespecimens inHalletal.,2004) andtherecentlydescribed have been found in amber, all of them in species Voltinia dramba based on five adult Dominican amber (Grimaldi and Engel, females (Hall et al., 2004), both preserved in 2005). For Miocene Dominican amber, the Miocene Dominican amber. A new female list of fossilized Lepidoptera includes other specimen of Voltinia dramba in the collection families: Blastobasidae, Cosmopterygidae, of the American Museum of Natural History Gelechiidae, Noctuidae, Tineidae and Tortri- (AMNH) provides an opportunity to com- cidae (Poinar [1992]), Tortricidae (Poinar and plete some details in the description of this Brown [1993]), and Oecophoridae (Kristensen interesting species. The most recent investiga- and Skalski [1999]). Grimaldi and Engel tions on the phylogeny of Riodinidae are (2005) figured a caterpillar and two adults of Harvey (1987) in Hall (2003), plus the sub- thefamilyGeometridae,anadultofthegenus sequent contributions by Hall (1998, 1999) Acrolophus (Acrolophidae), and other adults and Hall and Harvey (2002). The most recent of Tortricidae and Gelechioidea; an interest- study on the phylogeny of the subfamily ing tineoid moth figured by these authors is Riodininae, by Hall (2003), is a cladistic a case with the caterpillar inside. The scarce analysis of the 16 genera that have five butterflies found in Dominican amber have forewing radial veins; this study coded data been studied in detail, and are represented as on the adult ecology, wing venation and caterpillars of the families Nymphalidae and pattern, the adult head and body, male and Riodinidae (DeVries and Poinar, 1997; female genitalia, and early-stage ecology and Hammond and Poinar, 1998) and adults of morphology. That analysis confirmed the Riodinidae (Poinar, 1992; Grimaldi, 1996; monophyly of the tribes Mesosemiini and DeVries, 1997; Hall et al., 2004). Here, we Eurybiini, and established two subtribes for report two additional adult butterfly speci- Mesosemiini (Mesosemiina and the new sub- mens of the families Riodinidae and tribe Napaeina). The fossil species Voltinia Nymphalidae as inclusions in Miocene drambabelongstothesubtribeNapaeina.For Dominican amber; the nymphalid specimen Hall (2003), the best hypothesis about the is the first adult known of this family pre- relationships among the four- and five-radial- served in amber. vein groups of Riodininae is Mesosemiini + The family Riodinidae contains about (Eurybiini + all four-radial-vein tribes). a thousand species, most of which occur in The Nymphalidae is the most diverse family the American tropics and are partly charac- of butterflies, with more than 6,000 described terized by the greatly reduced male forelegs species worldwide. The nymphalid butterflies and the silver or leaden spots in the wings, are found in virtually every terrestrial habitat which is why these butterflies are commonly except Antarctica, and have their greatest named metalmarks (see DeVries, 1997). diversity in the Neotropics (DeVries, 1987). Riodinid caterpillars and pupae resemble Thenymphalidsarecharacterizedbyveryshort thoseoflycaenids,whichiswhysomeauthors forelegs in males and females, on which the classify them into a single family (Kristensen, males of many species carry a ‘‘brush’’ of long 1976; De Jong et al., 1996), though the scales.Thefamilycontainsmanycolorful,large families are usually considered sister groups species, and most of the mimicry complexes in (Ackery et al., 1999). The family Riodinidae butterflies. Nymphalid larvae feed on dicotyle- contains three subfamilies (Nemeobiinae, donous plants, very often toxic ones, and they Euselasiinae and Riodininae). Riodininae, are also generally adorned with a variety of the largest subfamily, is divided into eight spines and tubercles (DeVries, 1987; Smith et 2006 PEN˜ALVER AND GRIMALDI: MIOCENE AMBER BUTTERFLIES 3 al., 1994). The species of Nymphalidae have provenance within the outcrops of Dominican beenplacedintomorethan500generaclassified amberminesisunknown,butitsauthenticityis into12subfamilies.Theoldestnymphalidfossil certain, based on physical characteristics and is an undescribed form from the Early Eocene typicalpreservationoftheinclusions. ofGreenRiver(Colorado),butitsclassification The pieces were polished and some parts of within the family is unclear, and other Eocene the specimens were studied using sugar gel records are several species described from appliedbetweentheambersurfaceandaslide, Florissant (reviewed in Grimaldi and Engel, whichreducestheopticaldistortioncausedby 2005). The systematic relationships among the the curvature and imperfections of the amber subfamilies and tribesofNymphalidaeare still surface. Photomicrography used the InfinityE poorly known and thus the phylogeny of the K-2 long distance microscope and the family has been frequently discussed. There is MicrOpticsE fiber optic flash unit (www.mi- some evidence of a sister-group relationship croptics.com). The specimens were drawn between Lycaenidae and Nymphalidae, and using a drawing tube attached to a Zeiss Pieridae is possibly the sister group of the Stemi SV8 stereoscope. Specimens are housed in the amber fossil collection, Department of Lycaenidae + Nymphalidae (De Jong et al., Invertebrate Zoology, AMNH. 1996). Freitas and Brown (2004) presented a historical overview of Nymphalidae phylog- eny.Tworecentstudies,usingthewinglessgene SYSTEMATIC PALEONTOLOGY (Brower, 2000) and mitochondrial and nuclear genes (Wahlberg et al., 2003), concluded that CLASSINSECTALINNAEUS,1758 many of the traditional subgroups are mono- ORDERLEPIDOPTERALINNAEUS,1758 phyletic.Thelatestandmostcompletecladistic analysis to date, using 234 characters from all SUPERFAMILYPAPILIONOIDEA life stages of 95 nymphalid species, has been LATREILLE,1802 published by Freitas and Brown (2004), con- FAMILYRIODINIDAEGROTE,1895 cluding that the taxa can be grouped into six main lineages and it supports the monophyly SUBFAMILYRIODININAEGROTE,1895 and relationships of most presently recognized subgroups. That study also supported the TRIBEMESOSEMIINIBATES,1859 position of the subfamily Libytheinae as the SUBTRIBENAPAEINAHALL,2003 basalgroupofNymphalidae. Due to the poor fossil record of butterflies, GENUSVOLTINIASTICHEL,1910 they have rarely been used in phylogenetic Voltinia dramba Hall, Robbins, studies.Forthisreason itisveryimportantto and Harvey, 2004 study all remains found in the fossil record, Figures 1, 2a–c like the two specimens embedded in Dominican amber described here. MATERIAL: AMNH DR-18-1 (female) in alargeclearpieceofambermeasuring83 mm MATERIAL AND METHODS length 3 44 mm width 3 24 mm thickness. The piece also contains small specimens Dominican amber is fossil resin that was (syninclusions) of three dipterans, two formed by an extinctspecies of Hymenaea tree beetles, one moth (of the superfamily (Leguminosae: Caesalpinioidea), and is mid- Gelechioidea), one hymenopteran of undeter- Mioceneinage(Iturralde-VinentandMacPhee, minedfamilyandoneChalcidoidea,onemite, 1996; see the discussion in Grimaldi, 1995, and one seed. about the erroneous dating of this amber as Thisbutterfly is virtually complete and well Eocene-Oligocene by some authors). This preserved, lacking only the antennae, the amber derives mainly from outcrops in the proboscis and small portions of the posterior mountains in the north and northeast of area of the hind wings. The wings are over- Santiago. The material studied here was ac- lapping.Thevenationandcolorpatternofthe quired through purchase, and as such its exact left wings are very well preserved. The 4 AMERICAN MUSEUMNOVITATES NO. 3519 Fig.1. CameralucidadrawingsofthenewspecimenofVoltiniadramba(AMNHDR-18-1)inMiocene amber from the Dominican Republic. a. Complete dorsal view of body with the color forewing pattern. b. Hind wing with the color pattern preserved. c. Ventral view of body. d. Hind leg. e. Female genitalia. a–cto same scale. specimen occurs in a piece that, prior to vein (straight line), and in cells R to CuA 3 2 purchase, was broken into several portions, (submarginal line comprised of eight spots, which were then fused with adhesive. includingaverysmall spotatbaseofcellR ). 4 DESCRIPTION: Head with eyes bare, having TwoadditionalwhitespotspresentincellM3, entirelybrownmargins.Thoraxbrown,length one closest to the base and another in the 5.08 mm. Forewing shape triangular; length distal part. Faint spot in cell CuA present 2 23.6 mm, greatest width 14.9 mm. Five radial betweenthetwolinesthatcrossthiscell.Hind veins (fig. 1a). Underside with three narrow, winglength19.9 mm,greatestwidth13.9 mm. straight white bars at base of CuA cell, the Underside with three narrow, straight white 2 distal one is faint, and other three in discal barsatbaseofCuA ;distaloneiscurved,and 2 cell, all separated by darker brown areas. A other three in discal cell, all separated by faint bar present at the base of M and M darker brown surfaces (fig. 1b). Narrow, 1 2 cells,separatedfromthediscalcellbyadarker straight white bars in middle of CuA , CuA , 1 2 brown area. Three lines of white spots in cells andA +A cells(onlythedistalcolorpattern 1 2 R toM (straightdiagonalline),incellsCuA of cell M is visible); the bars of CuA and 2 2 1 3 1 to A + A at level of middle disco-cellular CuA have a darker brown basal margin and 1 2 2 2006 PEN˜ALVER AND GRIMALDI: MIOCENE AMBERBUTTERFLIES 5 Fig. 2. Photomicrographs of Miocene butterflies in Dominican amber. a. Female of Voltinia dramba (AMNHDR-18-1)inventralview,andcompleteviewoftheamberpiece.b.Apicaltibialspurinthehindleg ofthesamespecimen.c.DorsalviewofthenewVoltiniadrambaspecimen;thisbutterflyisfossilizedwiththe wings at an angle and so this image was made with six consecutive pictures of the specimen taken at successive focal planes. d. Male of Dynamine alexae n.sp. holotype (AMNH DR-18-2) in dorsal (left) and ventral view(right). Scalebars5 1 cminpanels a,c, d; 50.1 mm in panelb. 6 AMERICAN MUSEUMNOVITATES NO. 3519 acloser small white spoton thedistal margin. the hind wings has not been observable due to Submarginal, darker brown spots at least in theoverlappingofthewings. cellsM ,CuA ,andCuA (thelasttwoarenot Hall et al. (2004) placed V. dramba in the 3 1 2 complete). These spots large, semicircular in tribeMesosemiinibecauseitapparentlylacked shape, withnarrow paleborder. Wing pattern a hind tibial spur, in addition to other incellsSc+R toM notvisible.Legsbrown; characters. However, in the new specimen an 1 2 hind leg with distal tibial spur (length apical spur on the hind tibia is clearly present 0.28 mm) and white scales on coxa (fig. 1d). (figs. 1d, 2b). It is not easy to observe this Lengthoffemora,tibiae,basitarsi,andtarsiII structure, and surely it is present in the fossil + III + IV + V in mm: foreleg 2.15, 1.31, 0.69 specimens studied by Hall et al., though and 0.92; midleg 3.54, 2.31, 1.38 and 1.15; probably obscured. The presence of a spur hind leg 2.69, 3.15, 1.46 and 1.62. Claws does not preclude placement of the fossil unforked. Abdomen brown dorsally, ventral riodinids in the tribe Mesosemiini and in the surfacelaterallybrownandwithalongitudinal genus Voltinia, since this character is present band of pale brown scales (fig. 1c). Length in the extant species V. radiata and V. theata 8.15 mm, greatest width 2.31 mm. Female and in some other genera of Mesosemiini genitalia preserves the ostium of the bursa in (Hall,2003;Halletal.,2004).However,thisis segment VIII and the ovipositor lobes, which an important character for understanding are of moderate size (fig. 1e). relationships in the genus. NEWCHARACTERISTICSFORV.DRAMBA: The The differences observed in the color new specimen has practically the same wing patternofthenewspecimenarenotindicative color pattern as previously described speci- of a distinct species. These differences may be mens of the species, but shows some slight aconsequenceofintraspecificvariationand/or differences.Thenewspecimenhasaverysmall distinct preservation. It is well documented white spot at the base of cell R and a faint that butterfly wing patterns frequently show 4 spot in cell CuA between the two lines that spectacular differences among individuals of 2 cross this cell, both in the forewing. Also, the the same species (e.g., Brakefield and French, newspecimenhastwospotsnotpresentinthe 1999). Some additional spots observed in the original description of the species, one distal new specimen are faint, and possibly not faint spot in cell M and a white spot, in line, evident in all the fossils. Hall et al. (2004) 3 oncellA +A .Inaddition,thenewspecimen even indicated that the wing pattern recon- 1 2 lacks the extra faint spot at the base of cell struction was adapted from the sister species CuA in the hind wing. V.danforthi,andpresumablyisacombination 1 The most important difference observed in ofthewingpatternsofthefivespecimens.We our specimen is the presence of a distal spur describe the differences observed because in the hind tibia. In addition, the ventral variationforthisspeciesispresumablysignifi- surface of the abdomen is laterally brown cant. and has a longitudinal band of pale brown The riodinid caterpillar found previously in scales (vs. the ventral surface of the abdomen Dominican amber was identified as a member being completely brown in the original de- of the genus Theope (DeVries and Poinar, scription). 1997),whichbelongstothetribeNymphidiini. COMMENTS: Thisspecimenwaserroneously referred to as a nymphalid by Grimaldi and FAMILYNYMPHALIDAESWAINSON,1827 Engel (2005: fig. 13.68). Ironically, this identi- ficationwasbasedonstudyofthespecimenby SUBFAMILYBIBLIDINAEBOISDUVAL,1833 severalexperiencedlepidopterists.Themalefor TRIBEDYNAMININIBURMEISTER,1878 this fossil species is as yet unknown. The fracture surfaces of the amber piece hinder Genus Dynamine Hu¨bner, 1819 observation of the right wings, but their venation and color pattern have been drawn TYPE SPECIES: Papilio mylitta Cramer, and compare well with those of the left wings. 1780: 107. By subsequent designation Thecolorpatternoftheanteriormiddleareaof (Scudder, 1875: 160). 2006 PEN˜ALVER AND GRIMALDI: MIOCENE AMBERBUTTERFLIES 7 Dynamine alexae, new species and brown bands: underside with paired Figures 2d, 3, 4, 5 submarginal ocelli in M –M and CuA – 1 2 1 CuA , identical and very circular in shape; 2 MATERIAL: AMNH DR-18-2 (male), ho- ocelli connected by broad, brown, continuous lotype, in an amber piece 32 mm length 3 postmedial band (this band has a broad pale 22 mm width 3 8 mm thickness. brown proximal margin); one wide medial The holotype is incomplete, having lost the brown band preserved has a pale area in the anterior part of the body at the surface of the anal region; distal marginal brown band has amber (figs. 2d, 3), specifically the head, the a longitudinal, pale band for entire length. anterior part of the thorax, most of the right Proximal part of wing is not preserved. wings and proximal portions of the left fore- Margin without expansions or tails. Legs: and hind wings. Preserved are only the distal Tibiae with white scales ventrally and two partofthemid-andhindlegs,mostoftheleft large apical spurs. Mid and hind tarsomeres wings,andabdomenwithgenitalia.Thewings with four rows of strong setae (two ventral partly preserve the color pattern as tones of and two ventrolateral), except for tarsomere brownandareoverlapping.Thewingsurfaces V, which has two rows of ventral setae have small to medium gas bubbles trapped (fig. 3d). Length of tarsomeres of midleg: I,? during immersion in the resin. mm; II, 0.73 mm; III, 0.43 mm; IV, 0.29 mm; ETYMOLOGY: Patronym in honor of Mrs. andV,0.52 mm.Lengthoftarsomeresofhind Alex Goelet, wife of Mr. Robert G. Goelet, leg:I,2.29 mm;II,0.51 mm;III.0.23 mm;IV, Chairman Emeritus and trustee of the 0.26 mm; and V, 0.46 mm. White scales AMNH; for their generous sponsorship of present on tibiae. Claws unforked. Abdomen research at the AMNH. dorsally and laterally dark brown and white TYPELOCALITY: Mid-Mioceneamberfrom ventrally. Male genitalia with valves exposed, mines in the Cordillera Septentrional, north which are long, flattened, and narrow, and and northeast of Santiago, Dominican a narrow tubular structure that possibly Republic (Hispaniola). correspondstotheuncus(figs. 3c,5);theapex DIAGNOSIS: According to wing size this ofeachvalvehastwostrong,shortspines,one new Dynamine species was large. Greatest dorsal and one ventral. width of the hind wing larger than greatest width of the fore wing (HW/FW 5 1.19). COMMENTS: The female for this new fossil species is as yet unknown. The reconstruction White median band under both fore and hind of the wing color pattern of the preserved wings; costal white subband on fore wing areaswaspartial(fig. 4)becausethewingsare coversM –CuA cellandendsinCuA –CuA . 3 1 1 2 incomplete, overlapping, and touching. Two identical ocelli under hind wing. This specimen belongs to the genus Continuous, broad, brown postmedial band Dynamine due to the combination of the under hind wing connects the two ocelli, and following characters: presence of a white has a broad, pale brown proximal margin. Twostrong,shortapicalspinesatapexofeach medianbandonfore-andhindwings,absence male valve, one dorsal and one ventral. of ocelli on the forewing, two ocelli under the DESCRIPTION: Large body size. Forewing hind wing restricted to M1–M2 and CuA1– (figs. 3a,4):thepreservedportionofforewing CuA2 and connected by a broad brown band, indicates a triangular shape (fig. 4), inferred hind wingmargin without expansions or tails, length is 25.80 mm, and the greatest width is and a dark brown abdomen that is white 16.11 mm; L/W inferred 5 1.60. White medi- ventrally. Some species of Dynamine do not anbandonundersidebrokenatlevelofCuA have ocelli, but the two extant species present 1 in two subbands, the apical part of the costal in the Antilles have ocelli. We include the subband covers M –CuA cell and ends in genus Dynamine in the subfamily Biblidinae 3 1 CuA –CuA ; apical part of anal subband and tribe Dynaminini, sensu Oppler and 1 2 ending in vein CuA . Hind wing (figs. 3b, 4): Warren (2003) and Freitas and Brown 1 equal in length and width (inferred length is (2004). The subfamily Biblidinae is one of six 18.75 mmandgreatestwidth19.17 mm);L/W groups that have been established by the inferred 5 0.98. White underside with ocelli cladisticanalysisofFreitasandBrown(2004). 8 AMERICAN MUSEUMNOVITATES NO. 3519 Fig.3. CameralucidadrawingsofDynaminealexaen.sp.(AMNHDR-18-2)inMioceneamberfromthe DominicanRepublic,holotype.a.Bodyventralviewwiththehind-wingcolorpatternpreserved.b.Viewof the upper side of the forewing, showing the preserved color pattern. c. Male genitalia. d. Hind leg. a–b to same scale. 2006 PEN˜ALVER AND GRIMALDI: MIOCENE AMBERBUTTERFLIES 9 The family Nymphalidae has been found wingcolorpatternofD.alexaeissimilartothat previously in Dominican amber, represented of D. egaea on the continent and the West by a caterpillar placed near the genus Smyrna Indies. of the subfamily Nymphalinae (Hammond The species of Dynamine possessing two and Poinar, 1998), and thus without any ocelli that are not present in the West Indies relationship to the new fossil nymphalid are: D. ate, D. artemisia, D. onias, D. glauce, specimen. D. meridionalis, D. agatha, D. pebana, D. COMPARISONWITH EXTANT DYNAMINE: The aerata,D.paulina,D.perpetua,D.gisella,and genus Dynamine has very characteristic wing D. zenobia (for wing pattern and other colorpatterns,especiallythatoftheunderside, characteristics of these species, see D’Abrera, and for this reason it can be confused only 1987). Dynamine alexae differs from all of with the genus Lucinia. However, Lucinia has these species in some characters, most obvi- two larger wing ocelli, with each covering ously in a size significantly larger than that of more of one cell and not connected by any the first seven species listed. Butterflies in the continuous or broken brown band; and the genusDynamineareallsmall-sized(FWlength hind wing margin is not rounded. Ro¨ber from13to28 mmapprox.)and,forthisrange (1916) superficially segregated the species of D. alexae (FW length 5 25.80 mm) equals in Dynamine into three groups: 1) those with size the largest specimens of the largest little difference in wing pattern between males continental species (e.g., D. hoppi gillotti with and females; 2) those with appreciable differ- arangeofFWlengthfrom26to28 mm,orD. enceinthepatterningofthesexesandwithout hecuba from 22 to 24 mm). Furthermore, D. wingocelliontheunderside;and3)thosethat alexae n.sp. has a very broad hind wing differ in the wing pattern of the sexes and comparedtothegreatestwidthofitsforewing. which have wing ocelli. In extant Dynamine species the greatest width The genus Dynamine is represented in the is identical in both wings or even larger in the West Indies by only two species (D. egaea forewing. In addition, Dynamine aerata has Fabricius, 1775 and D. mylitta Cramer, 1780), the posterior wing ocellus larger that the both belonging to group 3. Dynamine alexae anterior one. Dynamine paulina has a very new species has wing ocelli, but it is unknown large anterior ocellus that extends over three whetherthe sexesweredimorphicand whether cells, and the white median band on the the wing uppersides had white spots or bands. undersideoftheforewingisdividedintothree Dynamine mylitta has a range from Mexico to small parts (fig. 4). Dynamine perpetua, D. Argentina including Cuba, and has a wing gisella, and D. zenobia have a very small, pattern very different from that of D. alexae costal, white subband on the fore wing in (theformerhasawhitemedianbandunderthe comparison with D. alexae. forewing divided in three small parts and the The male valves in extant Dynamine vary anterior ocellus on the hind wing is present in greatlyinform.Wehavecompared thevalves CuA –CuA ). Dynamine egaea has a range of D. alexae n.sp. with four species (fig. 5), 1 3 from Mexico to Colombia, including Cuba, which differ most in the structure of the apex. Hispaniola, and Jamaica, where it purportedly The two ‘‘subspecies’’ of D. egaea that were occursasthreesubspecies(D.e.calais,D.e.zetes studied from the West Indies have a broad, andD.e.egaea,respectively).Dynaminealexaeis granulate apex without spines. The continen- much larger than all the West Indies forms tal ‘‘subspecies’’ D. paulina thalassina has (25.80 mm estimated forewing length vs. 21– three strong, short spines (one dorsal and 22 mm;and18.75 mmgreatesthindwingwidth two ventral) in the apex. Lastly, the continen- vs.12.46 mm;seefig. 4).Thebroadcontinuous tal D. glauce and D. artemisia have two small band that connects the two hind wing ocelli is apical spines. similar to the band in the extant subspecies D. egaea calais from Cuba and D. e. egaea from TAPHONOMY Jamaica, and different from the subspecies on Hispaniola that has a narrow, occasionally Amber in general preserves insects and broken, brown band (fig. 4). In general, the other organisms that are uncommon in the 10 AMERICAN MUSEUMNOVITATES NO. 3519 Fig.4. FossilandRecentwingvenationsandundersidecolorpatternsinmalesofthegenusDynamine. ThereconstructionoftheundersidecolorpatternofDynaminealexaen.sp.isadaptedfromseveralRecent species of Dynamine (fig. 3 shows the partly preserved color pattern), and the reconstruction of the wing venation (depicted as gray lines) is adapted from the venation of D. egaea zetes from Hispaniola. All specimens are fromthe AMNH collection. Allto same scale.