MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY NUMBER 24 THE HIGHER CLASSIFICATION, PHYLOGENY AND ZOOGEOGRAPHY OF THE SATYRIDAE (LEPIDOPTERA) BY LEE D, MILLER PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY AT THE ACADEMY OF NATURAL SCIENCES PHILADELPHIA 1968 oo) MEMOIRS OF THE Vir CAaN EE NLOMOLMOCGICAL SOCIETY NUMBER 24 THE HIGHER CLASSIFICATION, PHYLOGENY AND ZOOGEOGRAPHY OF THE SATYRIDAE (LEPIDOPTERA) BY LEE D. MILLER o> > SS MTT, yy My > & oN Q My /, QD Nyy > R MM, PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY AT THE ACADEMY OF NATURAL SCIENCES PHILADELPHIA 1968 Breen esac Eahvl HSONTag> LIBR ARIES - y, Serr: £- SELWYN S. ROBACK EDITOR (Issued October 22, 1968) PRINTED IN THE UNITED STATES OF AMERICA To the memory of Dr. Richard M. Fox — Teacher, inspiration, colleague, friend. He is and will be missed. PREFACE No work is ever truly done alone: this paper is ample proof of the statement. I wish to take this opportunity to thank the many people and organizations who have helped to make it a reality. The research was supported in part by National Science Founda- tion grant number GB-2928 (R. M. Fox, Principal Investigator). The National Science Foundation, through Office of Science Infor- mation Services, made possible the publication of this work by grant number GN-652. Thanks are due the following individuals and institutions for their aid, loan of materials, use of facilities and general encouragement: Carnegie Museum, particularly Dr. M. Graham Netting, Director, Drs. Richard M. Fox and George E. Wallace and Mr. Harry K. Clench of the Section of Insects and Spiders; the Department of En- tomology, British Museum (Natural History), especially Messrs. J. P. Doncaster, Keeper of Entomology, and N. D. Riley, P. E. S. Whal- ley, T. G. Howarth and M. Clifton of the Lepidoptera section; and the Departments of Biology of the University of Pittsburgh and The Catholic University of America. Without their cooperation this pro- ject could not have been completed. The following individuals read the original manuscript which was submitted in quite different form as a dissertation at the University of Pittsburgh: Drs. Fox, Peter Gray, Eliot B. Spiess, Malcolm T. Jollie and Kenneth W. Cummins, Department of Biology, University of Pittsburgh, C. C. Li, Department of Biostatistics, Graduate School of Public Health, University of Pittsburgh, Craig C. Black and Mary Dawson, Section of Vertebrate Paleontology, Carnegie Museum. All were available for consultation and deserve great thanks. Mr. Clifton read the part of the manuscript dealing with the Haeterinae and made the necessary revisions on it based upon his forthcoming revision. Dr. Norman Tindale of the South Australian Museum read and commented upon the zoogeographic analyses, espe- cially those dealing with Australia. Dr. F. Martin Brown and Mr. Harry K. Clench read all or part of the manuscript and commented upon it. Particular thanks, however, are due Dr. Fox, who read and carefully critiqued the entire manuscript. He offered countless valu- able suggestions, most of which have been incorporated into the manu- script. Preparation of the manuscript and the figures has been aided greatly through the activities of two of my graduate students, Misses Jacqueline Y. Sessi and Dorothy A. Shea. Finally, thanks are due the many unnamed friends and colleagues who, knowingly or not, have contributed ideas and moral support for this undertaking. TABLE OF CONTENTS Page HINT RO IDIOKETBIOINT: aecccsaassaddesesodcobodacasobaocc Tod ease ach hase Gases BER CBOE EEE RAT Re aa eee I AW oXS) 12410010) (270.85 aoeceanncnbcecccdqpnodcocadHna aeosuosceosceeeor aoc neetooe are BOERS EECeCCe eerrnee | EVIStOMICAIMS KEtC Maes: eecesre sre ce se tec cee econ cscs earas curate cvavec eoeoneselastésecsvewceccces I Material Studied and Methods Employed ..................ccccccccccceeeeeeeeee 4 COMPARADIVE MM ORRETOIE@ GiyYaierec rrr eenee 7 PEN DIIGIIIDENS)_ gatpsostenoncuc cece octiacaqdcocasucocacccoo acc onCCC SUS OS CRC DCL CSO TOGe COAG REPRE EEE 1 15 IGEN ea sdasnucceccopdecaceoe nage lonoodcca Cac SS GHEE CEC Hc eae GO Cuan oe COE EAS cC ERED ET eee ieee eee 7 ab alla Pier rrccctecconscsessceeeer neces hence cede reve csaeuonberseetowanstensadsoesusueaeeseus 7 MINOR ART: oescccenstacccecns esas ecco dseiek Coa seats dunce uve iaeeGduesoends ROS OC EDC DOSS cece EAE EECEER 8 ROTCLE SS) iv sesessti evs sceucecossotdeesceuecesssessseucevdusesvecetouse 9990096006060000000000060008686 8 Mal eR LOTCLE Sis eitrecccsss sscreccreseusunncerk serve soad crue tue et tee densa lnc oesseiawntess 8 Remaleshonrele gsi rese-setees sesruteccccsuassecaes coos seceveout ccesten eos seee shessivvives 9 NAVE SY7) EEE © 5 SeccanonauadhoacHosqoono sos sSonccaoSbEs cacao sacaa Janae oO Dono pea aRocoe ERE nee 10 WNT S Shapes cect hac cec cs cacvueirasosebosen ses aassooticcccccevieeseuasestesetusussiasdat iss stesnsneanetacts 10 ROTC WANDS) coe cec caste reeeee oa ceeen ac chee eee e se ois nwa nene accel younetentne dee rucceontesass 11 PML LWA GS iresecoseoseaes Soca see ccc ce cee se castes oscewus suaety sacs se tlocicecsceneasuasieesews 12 Androcomiallpatchesieccsasscnscceveesoe snes sostetacscece crest eaec eco eee ee 13 IN fale G eniitialliia aye setea secre eae c eee eee co ee ee eae oa sore aan NE Sa 14 SYSTEMATIC REVISION OF THE GENERA OF THE SATYRIDAE 14 Family as abynidaemB O1sdinvall eisncsccesesessosercceetsstotonrceenorteccaceseerer scenes IIS) Key to the Subfamilies of the Satyridae «00.0.0... cece eeeeees 18 Subfamily Haeterinae Herrich-Schaffer .......c.cccceccccccccccccccecceeeeeeeeees 19 iinibesEacterinirennich=Schiatteiessseesscetterecntcesssesectecechcerecoeecce see 20 General IncludediiniitheMHaecteninie-ceescetsscteceececcecacee see teeeee: 22 Subfamily Brassolinae Boisduval .............cecceceeeeeeeeesseeeeeeeessseeeeeeeneees 23 MnriberBrassolimieB oisduvalteececcssscessscessecececteccereccenoscsesesseeee-ceeetececeenees 23 Generainchided inthe sBrassolimipesrsssesceeevececceeeeesssese se eee cee ceee 27 Subfamily Bimae) Herrich-Schatten --.-.c.s...cctc.ecesvecscceressssecctecsesssssessee 28 KeyatontheminibesrotsthesBimacrecncer see tecee crete ere reer 29 AmibewAmtinghinis new stli bey scessrssssectcoctssssoseessesserecceterecee oss suecenoeuesewent 29 Genera Includedtinithe vAmmtinghimit ties esr eestecee cceeeeeceeceree- 31 iribemBinielennich=Schateneccmece tees creceeccere te ececcre eee eee eeee 32 GenuspinchidedsinsthesBiiniltercecese-ccseeescceeseseseeeecee cree eee 34 MribesVMclanitinisemewathib@eeceeccsssccseeecerecececeeteeecesceceetenes cscs ecee eects 34 Genera Included in the Melanitini ...............cccccceeeeseseeeeeeeeeeeees 36 GenuseiVianaraniamKRanDy eens eee eee eens 36 Subfamily Elymniinae Herrich-Schaffer ....... cee cccccccccccccceceeeeeeeeeeees 38 Neyatonthe mnibessofatheElymmitmaeyeessescsseseescreecscectececeeseeee 40 nibe meth min lanky isc. csterccsettere set soso seve eee cce ccescc eal ice siorisuesstestesve’ 40 Genera Inchidedsiniithemeethimitecctsscestteseccencecsese-eeseeeceecee scenes 47 MEM. AMER. ENT. SOC., 24 TriberZethenmnignewatni eye ess sce ee cee eee 52 Genera IncludedtinitherZethenmiupee eee ee 33} Tribe Elymniunisernrich-Schatherier cece eee ee ee 54 Genera Included in the Elymniini .................ccececcccccsseesssseeeeeeeees 56 ibribe My calesinizmewetnibe ees te cetera oe 37/ Genera Included in the Mycalesini ........................ccccccceeeeeeesssees 60 Subfamily Eritinae, new subfamily ...............ceeeessscceesssceeessscccesseeees 64 Tribe Enitini; new tribes ecccssee eee eee 64 Genera Included in the Eritimi «0.0.0.0... eeeessneeeeeeeessneeeees 66 Subfamily Ragadiinae Herrich-Schaffer ...............cceeeeeeeeeceecceeeeeeeesenes 67 Tribe Ragadiini Herrich-Schaffer ...........:..ccccccccccscscccssccscccsececeeseseneees 67 Genera Included in the Ragadiini ..........0. cc ecceesssececeeeesenseeeee 69 Subfamily Satyrinae Boisduval ..............cccccccceeessseeeceeeecesesssceseceeesnes 69 Key to the Tribes of the Satyrinae .............eeeeeeeeeeeessssseeceeeeenenes 73 TribesHy pocystintmewtribeleesseccree cece seerceecaseees ee se eee erences 74 Genera Included in the Hypocystini ...............ccccecceeeeeeeeeeeees 80 Mribesy.pthiminisnewstribe cecscee eee oe 81 Genera Included in the Ypthimini 2.0.0.0... eeeceeeceeecceceeeeeeeees 85 Genus Ralaconympha Butler wecsssecccee tere ee 87 MribesEupty chime mew. tet Welc es eeesncveeccassees sens sercessorces cece seeesaeeeee 89 Genera Included in the Euptychiini ...............0.:eeeeeeeeeeeeees 92 Tribe Coenonymphini, new tribe ........... ccc eeeeeesseeeeeeeeeeeessseeceeeeeessssees 95 Genera Included in the Coenonymphini ........... ccc ececcececeeeeeees 98 ibribesManioliniMlamipsomeeceessesce cece eases cceeeeeseseeensccee eae eee eee 98 Genera Included in the Maniolini 0.0.0.0... ...ccccccsceessscssscceceeeeeeees 101 Tribe-Erebitini Mutt rasscc2 soe coseces er accesso rae casein tees oe Re 101 Genera Included in the Erebiint ..............0:ceeceestsceceeeeeessseeeeee 104 MribeWOinimis me WHS esses essa sees ose ote eae oes oeaeee eon 105 Genera Iincludedsin' the iD iim poses recone ceeteeeeeteee comers 108 MribePronophilimi}@larks ees se eeeeceeee sees eee ee 108 Genera Included in the Pronophilint «0.0.0.0... ccc ccccececcecceeeeeeeees 114 Mribe Satyrini Boisduval eirse eee ce ete ecreee acc ececcee cece ereaeneeeeeeeee eee 119 Genera Included in the Satyrint 2.0.0.0... cceeseeeseeereceeeceeeeeeeeeeees 123 Alors: IM lenNeIrETTIOT, \WSTETIN/” ce5s90cdececcescc0000900000000000050000000300000805090000000000% 124 Genera Included in the Melanargiini ....................ccc00eeesseereeeeees 126 Genera of WncentaimiRositiomiecs cc crececcceeetencecessteneceee te er ceceeeseeeettoe 127 GenusPampernis Heimlichy sescssrccereecccctccceseescetmeenteeeaseaee eee 127 GenusySerodocisIBillb er cee eerste ee aerate ee 128 li THE EVOLUTION AND ZOOGEOGRAPHY OF THE SATYRIDAE i28 OniginvofathesSatynidacmerecre teers eee ce renee ae eee sese ceo es sesereesecaes 131 Tee ILeter ISURIORY OE ToS SAARI sccocccosoccoccocoocane6oo0s05Nbn9040050805500000 134 Derivation of the Satyrid Faunae of the World ...............cceeeeeeeeees 142 HLT RIN COLO PICS oeeee se eeee eer ene se vem estes eee ences Eee wsetesy evs vansehtuces 145 TTLEYSEIN CRY CLI(C® “cecaneaadodosdo00coc eae oe Socceaeca dodo conn he DEE oC EE PEE EEEEE EEE 145 ASR alCATChi CMe ttesrcters ste cecs ec srsetre eee ccetecce cece ee cee ce ces Sees sssesesesiesiaeess 147 Mhemindo-Mialayanm Ne oiomireccesccrsccese tee ece eee tececctecestieceecsners sess 148 The Australian Region, including the Pacific Islands .............. 149 ‘Tove: Jeni abtoyoyeyny | Reka (Oy 0)) coseecccoccecascsedaadiocodec pea soser DOS uCoBE SO ECO CREE 150 IMac a aSCais seescnsttec seen ccoce cer atc ee everceecteseccetees oie ceucsusescausccnsten seeded 152 TWO SY EN OLGS PUAN TS INE 0 oa esnecrnc ecboceoa cose sexe reo A nee SCE Eat cbacEeECESE CECE Scere -DDEEE PEER EET EEE REEEAE 153 DLINITD EE ores ee a ance eee ean ves raced un ee aahuae ees vasnausecd ces sbasedsecbeoeasui viceneaeeesevs 164 MEM. AMER. ENT. SOC., 24 ili MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY NUMBER 24 THE HIGHER CLASSIFICATION, PHYLOGENY AND ZOOGEOGRAPHY OF THE SATYRIDAE (LEPIDOPTERA) By LEE D. MILLER INTRODUCTION The Problem. — The objective of this study, briefly, is the anal- ysis, both in space and, where possible, in time, of the evolution, phy- logeny and zoogeography of the higher taxa of the Satyridae. The existing studies on the family have been generally regional, and little progress has been made on interrelating the faunae of the various re- gions of the world. Furthermore, these studies were not based on modern, multi-variant taxonomic principles; hence, as will be shown, some fundamental errors in relationships have been perpetrated. Since no adequate taxonomic revision of the higher categories of the Saty- ridae is available, it has been necessary to complete one as a basis for the evolutionary and zoogeographic discussions. Using morphologi- cal and distributional data as bases, a scheme for the evolution and zoogeography of the Satyridae is proposed, and finally a phylogeny of the family is projected. Historical Sketch. — Linné considered all butterflies congen- eric. In the Tenth Edition of his Systema Naturae (1758), the start- ing point of zoological nomenclature, he published descriptions of a MEM. AMER. ENT. SOC., 24 2D THE SATYRIDAE number of satyrids, placing them with all other butterflies in the genus Papilio. The first nomenclatorially valid satyrid genus, Maniola, was erected by Schrank (1801) in his Fauna Boica. Fabricius (1807) described several more genera in Illiger’s rare Magazin fiir Inseckt- enkunde. Latreille (1810) proposed Satyrus in his Considerations générales sur l'ordre natural des animaux . . . crustacés . . . arach- nides_ .. . et insectes . . . Hiibner, between the years 1806 and 1819 (the latter date is bibliographically indeterminate and is brack- eted in all succeeding references), published many satyrid generic names in the Sammlung Exotischer Schmetterlinge and the Verzeich- niss bekannter Schmettlinge [sic]. Several other authors, notably Billberg (1820) and Meigen (1829), proposed early generic names for satyrids, but it was not until 1836 that the family was described, as “Satyrides”, by Boisduval. That author also first proposed the Brassolinae, as “Brassolides”’. The first comprehensive work devoted to the butterflies of the world in which the treatment was anything like a modern one was The genera of diurnal Lepidoptera by E. Doubleday, J. O. Westwood and W. C. Hewitson which appeared in two volumes between 1846 and 1852. In this work the Satyridae were defined, the brassolines delimited (as a subfamily of the Morphidae) and many generic names were added to the Satyridae. The practice of separating the brasso- lines from the rest of the satyrids became firmly established with this publication. The majority of the satyrid generic names were de- scribed in this work between 1849 and 1851, after Doubleday’s death, and are usually considered to have been authored by Westwood. The authors always referred to this work as “Doubleday, Westwood and Hewitson”, even though Hewitson’s chief role was as illustrator; but Brown (1941) holds that text descriptions must be credited to either Doubleday or Westwood alone, while names first proposed in plates (the text and the plates were often separately published) should be credited to Doubleday and Hewitson or to Westwood and Hewit- son. Accordingly, in those instances where the text appeared simul- taneously with, or prior to, the plates, I am crediting authorship to either Doubleday or Westwood, but if the plates appeared before the text descriptions, credit must be given to Doubleday and Hewitson or to Westwood and Hewitson. The first attempt at a higher classification of the satyrids (actually, LEE D. MILLER 3 of all the butterflies) was that of Herrich-Schaffer (1864). For the first time the satyrids were split beyond the Satyrinae and Brassolinae, as “Satyrina” and “Brassolina”. The Biinae (“Biina”), Hacterinae (“Hetaerina” [sic]), Ragadiinae (“Ragadiina”) and Elymniinae (“Elymniina”) were established. This work has been too long over- looked by systematists — all but one of the presently recognized saty- rid subfamilies were defined in it. Rober (1892), in the second vol- ume of Staudinger and Schatz’ Exotische Schmetterlinge, divided the satyrid genera into six groups with a residue of indeterminate genera. Rober’s groups generally correspond to major divisions of the Saty- ridae, but they were not nomenclatorially designated. Moore in the first and second volumes of his Lepidoptera Indica, published between 1890 and 1893, proposed a great many generic names but considered these genera to be apportioned into only the Satyrinae and Elym- niinae. These same divisions are to be found in Gaede (1931). The Rober scheme was followed with little modification by Fruhstorfer, Weymer and Aurivillius in their respective sections of Seitz’ monu- mental Die Grossschmetterlinge der Erde. More recently Clark (1947) subdivided the satyrids, exclusive of the brassolines, into the Satyrinae, Enodiinae, Pronophilinae, Elym- niinae and Pierellinae (a synonym of Haeterinae). In 1948 he emended Enodiinae to Lethinae and divided the brassolines, as a sep- arate family, into the Brassolinae, Caliginae and Biinae (this last not properly a brassoline). In neither paper did Clark give definitive reasons for his classification, hence he has been criticized by such authors as Ehrlich (1958) who proposed a higher classification of the “true butterflies” (Papilionoidea), retaining the separation of the brassolines and the satyrids and placing the former with the Mor- phidae. Table 1 attempts to relate the classifications of Herrich-Schaffer (1864) and Clark (1947, 1948) with that proposed in the present paper. In many instances particular genera were not apportioned by either Herrich-Schaffer or Clark, and those authors’ intentions can only be guessed. With the exception of Clark (1947, 1948) and Ehrlich (1958), no modern author has attempted to place the satyrids in their proper evolutionary positions, and there has been no zoogeographic analysis of this family. Schwanwitsch (1924), however, relied heavily on MEM. AMER. ENT. SOC., 24 4 THE SATYRIDAE TABLE | A comparison of the present subfamilial classification of the Satyridae with those proposed by Herrich-Schaffer (1864) and Clark (1947, 1948). Present Herrich-Schaffer’s Clark’s classification classification classification Hacteninae wees ater ee Hetaerina (sic) ................ Pierellinae Brassolinacieresscsiccceeseseeeeee BLEASSOlinarers tee eeeee eee Brassolinae Caliginae Bina eee ie ee Bilaeeiiccs eee eee eee Biinae (of Brassolidae) Satyrina (part) Satyrinae (part) El ymimiimacieeresccccececeeceoneee Elymmiimay eee eee ecee ee ceeeeeees Elymniinae Satyrina (part) Enodiinae (=Lethinae) Eritinae eee eee SEA D U0 e) sonscnccsasonansceoocacee Satyrinae Ragadinae ete ee Ralgadiinayee.ceee tee Satyrinae SatyriMaeieees ee eee Satynin ayes eee Satyrinae (part) Pronophilinae satyrids in his study of the evolutionary trends toward the develop- ment of the “basic nymphalid pattern”. This work was followed by a number of other papers concerning pattern modifications, two of which are of interest here: in 1925 he published an analysis of the Pierella-type pattern, a paper which has since become a classic in the explanation of the morphological migration of pattern elements; and in 1931 he analyzed the highly modified Melanargia-type pattern. Both papers were significant, but the one explaining “pierellization” has had the most far-reaching implications. Material Studied and Methods Employed. — Since there are between 2500 and 3000 species in the Satyridae, it would be im- practical to attempt a complete revision of all the species in a single work. The number of described genera, on the other hand, is man- ageable — fewer than 400. A fair axiom in systematics may be stated more or less as fol- lows: systematic decisions at one taxonomic level should be reached by examination of most (preferably all) of the entities at the next lower major taxonomic level. In other words, if one is revising a genus, one should see representatives of the species and subspecies included in it to make meaningful taxonomic judgments at the spe- cific and generic levels. In the present study examples of the already- named genera were examined, hence, taxonomic judgments are made at the tribal level, and no synonymization of nomenclatorially valid LEE D. MILLER 5 genera (only pure objective synonyms) has been attempted, nor have the several new genera which certainly exist been named. This is work for careful species-by-species revisions. It is certain that many of the nomenclatorially valid genera are not biological entities; for example, I am unable to ascertain any significant differences at the generic level between the seven nomenclatorially valid genera included in the satyrine tribe Melanargiini. Nevertheless, all of the nomencla- torially valid names are carried in this revision. For the initial work on this project the collection of Carnegie Museum was examined for representatives of the known genera. It soon was apparent that the material at hand was not sufficient, so two months were spent at that Mecca of butterfly systematists, the British Museum (Natural History), examining those genera which were either unrepresented at Carnegie Museum or only poorly rep- resented. Material was found and examined in these collections rep- resenting all but four of the genera that had come to my attention by August, 1964. Those genera that have been described since that time are listed in what I believe to be their proper systematic posi- tions, but none of them has been examined as critically as have the others. Sokal and Sneath (1963: 161-162) explain the “exemplar meth- od” of taxonomy as follows: a taxon may be characterized by a small number of entities within it (even a single specimen) because the variation within taxa is considered to be less than that between taxa. This is somewhat the method that has been employed in the present study. The type-species of a genus is considered to be typical of it, and the various genera are defined chiefly by their type-species in the following pages. Occasionally the type-species is aberrant, but it is the type-species which bears nomenclatorial “responsibility” for the generic name, in any event. Where possible, other species of the genera were examined, though not in as great detail as the type- species. The systematic revision is based entirely upon external morpho- logical characteristics with many attributes being considered. Wing venation drawings were made from specimens either with an ocular grid under a dissecting microscope or with a projection device for larger specimens. Antennae, palpi, fore-, mid- and hindlegs were re- moved from the specimen, bleached in clorox and mounted on perma- MEM. AMER. ENT. SOC., 24 6 THE SATYRIDAE nent slides. Drawings of these structures were made with a camera lucida. The taxonomic and phylogenetic analysis of the family has been generally based on the premises that (1) entities which more nearly resemble one another in many characteristics are most closely re- lated, (2) structures which are reduced and fused represent advanced conditions (Fox, 1956: 24; Fox and Fox, 1964: 71-72: Brown, 1965) and (3) those organisms which bear advanced characteristics (not just a single advanced character, but several of them) are prob- ably themselves advanced (for example, see Mayr, 1965). This is in accord with the current paleontological evidence drawn from ex- isting fossil records. The only taxonomically valuable structures not utilized in this study were the male and female genitalia. Examination of several regional works (for example, Hayward, 1953; Forster, 1964) shows little reliance can be placed on genitalic structures for classification above the generic level, although these structures are certainly of great importance in specific, and sometimes generic, determination. Authorship of higher taxa has caused some problems. According to Art. 36 of the International Code of Zoological Nomenclature (1961), the proposition of the family Satyridae by Boisduval (1836, as “Satyrides”) implicitly creates also the subfamily Satyrinae and the tribe Satyrini. It is this rule which has been followed through- out the present paper for the assignment of authorship to the higher taxa. The zoogeographic analysis has been carried on in light of the principles of vertebrate zoogeography, which is firmly based on and supported by fossil evidence. For a summary of such evidence see Darlington (1957). There are too few butterfly fossils to support or refute any zoogeographic statement. Heavy reliance has been placed on the presence in one area or another of relict, annectant forms: such species may occur nearer the place of origin of the taxon than do others. Although fossil evidence shows that such groups as horses and rhinoceri evolved more extensively in the Nearctic than in the Palearctic, yet are now absent from the former, many other groups have evolved more or less in the places of their greatest present-day density and diversity. In the absence of fossil evidence to the con-— trary, the latter situation is considered to be likely in the satyrid but- terflies. LEE D. MILLER 7 COMPARATIVE MORPHOLOGY The characters utilized in this study are all external and mor- phological. Their distribution and evolutionary patterns are given in the following pages. Antennae. — Four measurements were made on the antennae: the total length; the length of the club, where the club is a discreet entity; the width of the shaft and the width of the club at its thickest point. The antennae of Satyridae demonstrate the tricarinate con- dition typical of the Nymphaloidea (Fig. 249), although the ridges may not be apparent throughout the entire length of the antennae. The antennae are scaled, but only on the basal few segments in the Haeterinae. In general, the antennal club is not greatly thickened nor flattened at the tip in the primitive satyrids (Figs. 114, 122, 128 which are some of the best developed ones), thereby following the general primitive condition in butterflies. Many of the more advanced Satyrinae, conversely, have the antennal club strongly developed, flat- tened at the tips and occasionally exhibiting bizarre configurations in some Hypocystini and Pronophilini (Figs. 163, 166, 169, 291, for example). The antennal clubs of other Satyrinae are shown in Figs. (elie ISIE 2S 2205235, 2495 2622297. Head. — Three measurements were made on the head region: least intraocular distance; height of the eye and width of the eye. The eyes of all satyrids are entire. Little significance could be gleaned from these measurements, but the eyes of the Satyrinae appear to be more ovoid than those of other subfamilies. Labial Palpi. — In common with other butterflies the labial palpi of the Satyridae are three-segmented. The basal segment is so firmly attached to the head that it was often broken in dissection, hence is little used in the present study. Five measurements were made on the palpi: the length of the first segment, when it was not broken; the length of the second segment; the length of the third segment; the maximum width of the second segment and the maximum length of the hairs of the second segment. The relationship between the lengths of the second and third segments of the palpi varies, often without regard to phyletic groups. For example, most of the Satyrini display an extremely short third palpal segment, about one-fifth as long as the second, whereas the Pronophilini, which are closely related to the Satyrini and may have been derived from a fairly recent common an- MEM. AMER. ENT. SOC., 24 8 THE SATYRIDAE cestor, show the longest third palpal segment of any satyrid tribe — in some instances over half as long as the second segment. The sec- ond segment is generally between five and eight times as long as it is wide, again with no apparent phylogenetic correlation. The relative lengths of the hairs of the second segment of the palpi show some in- teresting trends. In general the hairs are longer in the Satyrinae than in the other subfamilies, but this tendency is modified in those groups which have both tropical and temperate or alpine members. In such instances the relative length of the hairs increases as the altitude or latitude increases. Examples of the labial palpi of the Satyridae are shown in many figures in the text, such as Figs. 3, 10, 22, 30, 35, 50, 82, 93, 1007 129, 135, 1437 W298. 214.2255 236) 25052 Ose PDN 5 BMI, Thorax. — Only one thoracic measurement was made: the length of the dorsal thoracic elements. This measurement, unfortunately, is generally dependent only upon the absolute size of the insect. Forelegs. — The forelegs of both sexes are more or less reduced in all satyrids, as in other members of the Nymphaloidea. The fore- coxae are moveable, as determined by examination of many (though not all) genera. The forecoxae are not reduced to the extent that the more distal elements of the forelegs are. The reduction of the pro- thoracic femur, tibia and tarsus has long been the criterion for re- garding the Nymphaloidea as the most advanced of the lepidopterous insects, a concept recently challenged by Ehrlich (1958). Male forelegs. — Four measurements were made on the male forelegs: the length of the tarsus; the length of the first tarsal sub- segment; the length of the tibia and the length of the femur. The miniaturization of the male forelegs generaliy follows phylogenetic lines, being least miniaturized (and reduced; for a discussion of the terms “miniaturized” and “reduced” see Fox, 1967) in the Haeter- inae, Brassolinae, Biinae, Elymniinae and Eritinae and most in the Ragadiinae and Satyrinae (compare the graphs in Figs. 1, 20, 170, 309, for example). In the first five subfamilies the femur, tibia and tarsus usually are all relatively well developed (an exception is Cae- rois, Fig. 27, where the tibia and the tarsus are much miniaturized). The foretarsus in these primitive subfamilies is typically monomerous, but some genera may have up to four freely articulating subsegments. Examples of the forelegs of typical primitive satyrids are Figs. 4, 11, LEE D. MILLER 9 23, 51, etc. In the Ragadiinae and Satyrinae the forelegs are much more miniaturized and reduced than in the other five subfamilies, and occasionally there is an apparent total loss of the tarsus (Ypthimini, Fig. 177). The tibia and femur are also greatly miniaturized, but the coxa does not seem affected by this general trend. An anomalous situation exists in the Satyrini and Erebiini, however, where a few genera display great miniaturization of the total foreleg but still re- tain up to four distinct, though tiny, tarsal subsegments. The male foreleg of Ragadia is shown in Fig. 136, and those of representative Satyrinae are shown in Figs. 144, 173, 193, 199, 215, 226, 237, 251, ANG. AQ, BUA, Cee Female forelegs. — Five measurements were made on the female forelegs: the length of the tarsus; the length of the first tarsal sub- segment; the length of the fifth tarsal subsegment, if any was present; the length of the tibia and the length of the femur. The female fore- legs potentially show more precise inter- and intrasubfamilial rela- tionships than do those of the males. The forelegs of the Haeterinae, Brassolinae, Biinae and Elymniinae are quite well developed (Figs. 1, 20, 41, 48, 88, 89, 98), always have pentamerous tarsi and usually have posttarsal pulvilli present, except a few of the more advanced Elymniinae. In these families the tarsus is not, or only weakly, clubbed (for example, Figs. 5, 15, 24,52, 112). In only two genera, Paralethe and Antirrhea (Fig. 24) are there posttarsal claws, and always there is but a single claw. This represents a more advanced condition generally than that shown in the Ithomiidae (Fox, 1956: 13). The female forelegs of the Eritinae (Figs. 125, 131) and the Ragadiinae (Fig. 137) are miniaturized beyond the point shown in the preceding subfamilies, but they retain the full complement of tarsal subsegments; the tarsus itself is clubbed and not so freely ar- ticulated as in the primitive subfamilies. In the Satyrinae the fore- legs range from miniaturized, but with five tarsal subsegments, to highly reduced, as much so or more than in the males. Hypocystini, Ypthimini (interestingly where the greatest reduction takes place in the male foreleg), Euptychiini, most Pronophilini and some members of several other tribes have pentamerous tarsi (Figs. 145, 174, 194, 200, 267, for example), but the tarsus is always clubbed. The fore- legs of female Satyrini, Melanargiini, Dirini, Coenonymphini and Maniolini are always reduced and apparently lack one or more tarsal MEM. AMER. ENT. SOC., 24 10 THE SATYRIDAE subsegments (Figs) 26,227) 2380252, 3005 Sil 3esion example)r The miniaturization of the forelegs of Satyrinae through the phyletic lines may be seen by examination of Figs. 140, 170, 211, 295, 309, etc. Walking Legs. — Six measurements were made on both the mesothoracic and metathoracic legs: the length of the tarsal claws; the length of the tarsus, exclusive of the claws; the length of the first tarsal subsegment; the length of the tibia; the length of the tibial spurs, if any, and the length of the femur. The more important taxonomic characters are on the mesothoracic legs. The lack of tibial spurs characterizes the Haeterinae, Biini and Elymniini (Figs. 6, 33, 96), and these spurs have been lost sporadically in such other groups as the Ypthimini (Fig. 179) and Coenonymphini (Fig. 221). The posttarsal claws are bifid in the Melanitini (Fig. 39) and in a few other genera (Samanta and Manataria, Fig. 47). In all other Saty- ridae the claws are simple. The relative lengths of the midtibia and the first midtarsal subsegment are important taxonomically, particu- larly within the Satyrinae. Usually the midtibia is between one and a half and two times as long as the first midtarsal subsegment, but the midtibia is barely longer than the first tarsal subsegment in the Satyrini (Fig. 301) and in some Dirini (Fig. 259), whereas the mid- tibia is always more than twice as long as the basal midtarsal sub- segment in the Coenonymphini (Fig. 217). All members of the Satyrus series of the Satyrini display a heavy spine at the dorsal distal end of the midtibia (Fig. 301), a characteristic weakly shown by some Dirini (as Torynesis, Fig. 259), thereby indicating a close re- lationship between these two tribes. Dorsal spinosity of the midtibia is shown by some members of most groups and is characteristic of others: Haeterinae, Zetherini, Pronophilini, Dirini, Satyrini and Me- lanargiini. Examination of the charts of relative lengths of legs shows that the hindlegs are occasionally significantly longer (Satyrini, Fig. 295, for example) or shorter (the Pronophila series of the Pronophilini, Fig. 260, for example) than the midlegs, but generally these legs are subequal. Wings. — The wings have long been the most-used basis for the classification of butterflies, both from the aspect of venation and the distribution of androconial patches. Undoubtedly too much empha- LEE D. MILLER 11 sis has been placed in the past on wing venation and androconial dis- tribution alone, but it is illogical to ignore the structure of the wings because of previous preoccupation. Forewings: The following four measurements were made to de- limit the shape of the forewing: the length from the base to the end of vein Rs; the length from the base to the end of Me; the length from the base to the end of 2A and the distance between the end of Rs and the end of 2A. The cell was delimited by two measurements: from the base to the origins of Mi and Ms. Five measurements were made to determine the relative positions of the origins of the veins: the dis- tance from the origin of Rs to the origin of M:; from the origin of Mi to the origin of My; from the origin of Me to the origin of Ms; from the origin of Ms; to the origin of Cu; and from the origin of Cun to the origin of Cus. Inasmuch as the inflation of the forewing veins has long been considered a key characteristic of the Satyridae, the following three measurements were made: the thickest part of Sc; the thickest part of the cubital stem and the thickest part of 2A. In gen- eral the longest forewing cell is displayed by the most primitive sub- families, the Haeterinae (Figs. 2, 7, 8), Brassolinae (Figs. 9, 13, 14, 17, 19) and Biinae (Figs. 21, 26, 28, 29, 34, 40), where the cell is frequently more than three-fifths the length of the wing. In most other groups the cell is about half as long as the wing, but the Elym- niini are characterized by an extremely short forewing cell (Figs. 90- 92), often less than a third the length of the wing. There appear to be no broad phylogenetic implications shown by the relative place- ment of the veins along the cell, though several relationships are shown within subfamilies, particularly the proximity of the origins of Ms and Cu: in the Elymniini (Figs. 90-92). The classic definition of the Satyridae includes a statement about the inflation of the stems of the three basic forewing veins. This is simply not diagnostic: the inflation of the veins of Melanargia (Fig. 310), for example, is con- siderably less than that of the true nymphalid genus Mestra. Within the Satyridae particular inflation patterns are characteristic: the My- calesini are characterized by the inflation of all three stems, that of 2A being more or less quadrate (Figs. 99, 104-106, 109, 113, 115, 119); all stems are inflated in the Hypocystini (Figs. 141, 147-150, 156, 161, 162, 165, 167, 168) and Coenonymphini (Figs. 212, 219), but the inflation of 2A is rounded, not quadrate. In contrast, most MEM. AMER. ENT. SOC., 24 12 THE SATYRIDAE Biinae, Ragadiinae, Eritinae and Elymniinae (except the before- mentioned Mycalesini) have relatively uninflated cubital stems and 2A (Figs. 21, 29, 34, 49, 81, 90, 127, 134, for example). The pres- ence of a vestige of forewing vein 3A is characteristic of the Haeteri- nae (Figs. 2, 7, 8) and seems to connect them with the Ithomiidae. Hindwings. — Five measurements were made to delineate the shape of the hindwing and account for the majority of “tails”: the length from the base to the end of vein Rs; the length from the base to the end of Ms; the length from the base to the end of Cus; the length from the base to the end of 2A and the distance from the end of Rs to the end of 2A. The cell was delimited by two measure- ments: from the base to the origins of M: and Ms. Five measure- ments showed the relative placement of the veins along the cell: from the origin of Rs to the origin of Mi; from the origin of Mi: to the origin of Me; from the origin of M» to the origin of M3; from the origin of Ms to the origin of Cu: and from the origin of Cu to the origin of Cuz. In all Satyrinae except Dirini (Figs. 248, 255, 256) the hindwing cell averaged greater than half the length of the wing measured to the end of Ms (for example, Figs. 141, 171, 197, 212, 224, 234, 261, 296, 310), whereas in the other subfamilies the hindwing cell is half or less the length of the wing, again particularly short in the Elymniini (Figs. 90-92). The actual shape of the cell, which is not reflected in the raw measurements, has some phyletic significance. The cell is distally rounded in the Haeterinae (Figs. 2, 7, 8), perhaps representing a more generalized condition. In the Biinae a gradation may be seen between a rounded cell and one pro- duced at the origins of Ms and Cu: (Figs. 21, 26, 28, 29, 34, 40). The Elymniinae and the “Ypthimini complex” of the Satyrinae, as well as the Eritinae, have the cell produced at the origin of Ms (Figs. 49, 90, 99, 127, 171, 197, for example), whereas the Hypocysta series of the Hypocystini and the “Satyrini complex” of the Satyrinae, in general, display a migration of crossvein m2-m; outward at Me and a corresponding blunting of the cell (Figs. 141, 261, 296, 304, for ex- ample ). The Ragadiinae (Figs. 134, 139) do not fit another part of the classic definition of the Satyridae — the closure of the cells by tubu- lar veins. In males of this subfamily the hindwing cell is closed by vestigial, non-tubular veins, apparently only for the support of an LEE D. MILLER 13 androconial patch; some females have the cell completely open. Because of the migration of crossvein m2-m3, Me arises nearer to the origin of Ms than to the origin of Mi in some satyrine tribes; in other groups Me arises somewhat to much nearer Mi. Connate veins are generally considered an advanced characteristic of the Lepidop- tera, but in the Satyridae the opposite may be true. Hindwing veins Ms: and Cu: are frequently connate in the more primitive subfamilies, particularly the Haeterinae and the Elymniinae (for example, Figs. 2, 60, 90), but these veins are well separated in the Satyrinae (for example, Figs. 261, 296), which are more advanced from a number of other viewpoints, as detailed in the preceding pages. This situa- tion is emphasized chiefly as an illustration of the independence of characters and the danger of reliance on too few attributes, as has been suggested by some authors (for example, Warren, 1947). Hind- wing vein Ri of the hindwing characteristically is basally free in the Brassolinae (Figs. 9, 13, 14, 17, 19) and Elymniini (Figs. 90-92). Androconial Patches. — As noted previously the distribution of androconial patches has been used extensively in the establishment of genera (particularly by Moore, 1890-1893) — probably to too great an extent. A few general statements may be made concerning the distribution of these patches. There are several sites of the produc- tion of androconia on the forewing, the most common of which is the discal area just outside the cell from the inner margin to the origin of Ms. This patch may be highly modified to include all or any part of the area and generally consists of a patch of mealy scales. Hair patches are less frequent on the forewing and are usually restricted to the area below the cubital stem on the upper surface or along 2A on the under surface, particularly in the Biinae and Mycalesini; re- sultant distortion of the venation may occur in these groups. Several areas generate androconial patches on the hindwing. Mealy patches are generally found near the end of the cell in the area of Ms (for example, Fig. 54). Hair patches are found in three general areas: above the cell near the origins of Rs and Mi (for example, Fig. 73), within the cell proper (particularly characteristic of the Elymniini, Figs. 90-92, and Ragadiinae, Figs. 134, 139) and along the anal veins (for example, Fig. 121). Since androconia appear sporadi- cally within most groups they seem to have little phylogenetic signifi- MEM. AMER. ENT. SOC., 24 14 THE SATYRIDAE cance, though particular configurations may be characteristic of a few tribes. Male Genitalia. — Although these structures have been used ex- tensively by some authors for the establishment of genera (particu- larly by Forster, 1964), the genitalia are not considered in the pres- ent study because of the great variability shown in such genera as have not been split on genitalic grounds. While the genitalia are undoubtedly indispensable for the determination of intrageneric affin- ities, their unrestricted use as the basis for establishing genera is un- tenable, partly because doing so creates hierarchical difficuities in that genitalia often do not correlate with other structures, resulting in numerous monotypic genera. Too many monobasic taxa tend to obscure relationships within a larger group. SYSTEMATIC REVISION OF THE GENERA OF THE SATYRIDAE Before meaningful statements may be made concerning evolution, phylogeny and zoogeography. it is necessary to analyze relationships. No usable taxonomy presently exists for the Satyridae; the attempt in the following pages to apportion genera to subfamilies and tribes is designed to fill this need. This classification is based on many char- acters in the hope that it may more nearly approximate a “natural” classification than have some schemes in the past which were based on one or very few characteristics. It is the purpose of this revision to define, as closely as possible, the subfamilies and tribes which comprise the Satyridae, as well as to define the family itself. No single character can or should be ex- pected to be diagnostic of an entire group: evolution modifies char- acters indiscriminately, without regard to “sacred” taxonomic ones. That a single character — or even several — is atypical of the group in some genera is no reason to assign such genera to other tribes (Simpson, 1961la: 54-63). The practice of “splitting” across natural boundaries comes from the indiscriminate analysis of too few char- acters. Two other nymphaloid families are considered to be systemati- cally and phylogenetically close to the Satyridae: the Morphidae and the Ithomiidae. The Morphidae, which are represented in both the Old and New World tropics, have been regarded by many authors (e.g., Ehrlich, 1958) to be very near the Satyridae. With few exceptions the hind- LEE D. MILLER 15 wing cell of the morphids is open between veins M2 and M3;, whereas only satyrids of the subfamily Ragadiinae have the cell closed by other than fully functional tubular veins. The structure of the mor- phid legs is substantially that shown by the primitive satyrids, but larval foodplant preference will separate some morphids. All the Indo-Australian morphids feed as larvae on monocotyledons, as do the satyrids, but the Neotropical members feed predominantly on di- cotyledons. There are a few nymphalid genera that utilize mono- cotyledons, but Morpho is unique among nymphaloids in commonly using both monocots and dicots (Ehrlich and Raven, 1965). On the larval foodplant preference and on pattern morphology (see Schwan- witsch, 1924) the New World morphids appear to be very close to the condition apparent when the fundamental dichotomy of the nym- phaloids into monocotyledon and dicotyledon feeding groups tock place. Fox (1956: 18-19) considers the Ithomiidae to be as near or nearer the Satyridae than to the Danaidae, with which the ithomiids had formerly been placed. This conclusion was reached from the evidence of various characteristics, particularly of the adults and pu- pae. In many respects the ithomiids resemble the Haeterinae of the Satyridae, particularly as regards the presence of a vestige of fore- wing vein 3A (a characteristic also shared with the danaids) in both groups. The obviously similar tendency of the ithomiids and the haeterines to have hyaline wings is, therefore, supported by structural evidence. The vestige of forewing vein 3A will serve to separate the ithomiids from the remainder of the satyrids, and no discreet andro- conial areas are found in the haeterines, but these are usual (and taxonomically important) in the ithomiids. All Ithomiidae feed as larvae on the dicotyledonous family Solanaceae. The morphids ap- pear to be even more closely related to the satyrids than the ithomiids, but neither can be combined with the satyrids. Family SATYRIDAE Boisduval, 1836 Satyrides Boisduval, 1836: 166. = Hipparchiadae Kirby, 1837: 297. = Satyridae Swainson, 1840: 86. As mentioned in the preceding section, the Satyridae are related MEM. AMER. ENT. SOC., 24 16 THE SATYRIDAE to the nymphaloid families Morphidae and Ithomiidae. No single key character will serve to separate the satyrids from all other nymphaloids, but a general definition of the Satyridae may be formulated as follows: a nymphaloid family having a larva with a bifid tail (fleshy anal projections from the eleventh abdominal seg- ment) and feeding on monocotyledonous plants and generally having adults with the cells of the fore- and hindwings closed by tubular veins, forewing veins swollen at their bases and with more or less shaggy palpi. Only the habit of feeding on monocots and the bifid larval tail are universal in the Satyridae, and some morphids feed on monocotyledons. Many other nymphaloids (danaids, ithomiids, ac- raeines, etc.) have the cells of both wings closed by tubular veins, and the satyrid subfamily Ragadiinae is defined by not having the hindwing cell closed by tubular veins, often not closed at all. While many of the satyrids have shaggy palpi, many of the primitive ones have only very short hairs on the palpi. This part of the definition universally applies only to the temperate and arctic species. The forewing veins are usually, but not universally, swollen at their bases. The bases of the forewing veins of such satyrid genera as Melanitis, Lethe, Oeneis and Melanargia are less swollen than are those of such “true” nymphalid genera as Mestra, Bolboneura and Callicore (see Schatz and Rober, 1892). The Satyridae are nearly cosmopolitan, being found wherever but- terflies in general are found, except on some oceanic islands. These butterflies, like most others, find their greatest development and pro- liferation in the tropics, not in the temperate zone as claimed by Ehr- lich and Raven, (1965: 589). Seven subfamilies are recognized in this family. The most primi- tive of these are the Haeterinae, Brassolinae and Biinae; intermediate are the Elymniinae and Eritinae; the Ragadiinae and Satyrinae are advanced. In the following pages the individual subfamilies are characterized and diagnoses are provided for the tribes embodying the following parameters: (1) condition of the eyes; (2) relative length of the an- tennae; (3) characteristics of the antennal club; (4) relative lengths of the second and third segments of the palpus; (5) the development of the hairs of the palpus; (6) the development of the male foreleg; (7) the development of the female foreleg; (8) the relative lengths 17 Ajaja|du09 payeos ‘aTquiiea qniO poonpol Apeain coos poqqnys SI JI S}UOUL -sosqns ¢ jl “sjuc Wisasqns ¢ uvy} Jomo * % ica] =| = = OS YIM S posowojseuy ~ om . > w ©uu-Su 1e — psonpod 10 WysIes AT]eI Ajaqyo;dwos Ajayoi duos Ajajo|dui05 Ajajo|du05 Ajajo|du05 poyeos ‘qe poyeos poeos payeos ‘yeom poyeos ‘yeom -Iopoul qnjo "" “yeam qnyg “yeaa qni[D * Ajjesouas qnyiO * Ayjesouas qniD peonpol uoljonpel poonpol poonpol poonpol O9COO000 Apeoig Ap00000000 aque A 600000 Ajyeais 1ON 600000 Ajyeois 10N ntaleterste Ayyeois ION poqqnys os Apyeom poeqqnys Ayyeom =. JO poqqnyoun peqqnypoun peqqnyjoun ‘snojowejusg “ ‘snoisuejuodg ~ ‘snojouiejuodg © “shortourejyuod * “snojouejusd * ruruwAy Ul Ayyeseq 901J 9g WIM oS WM yng “OS YUM oS WM posowo}seuy * posowojseuy ~ poesowmojseuy ~ pasowojseuy *”" Ajjeseq 9014 SUIOA juRIIoge }YsIv1}S }YsIe.1}S }Ys1v.18 WYSsIeI}s Aq pasos A[eAsIp A]yeqstp Ayyeqsip Ayyeasip -sIp puv paso[g © Ajuo Jo ‘uedgQ “""" pue pasojD “"* pue pasojD " pur paso[D “"" Pue PEeso|) VZ YIM pasny “WZ YIM pas “WZ YM pasty “WT YUM pasny “VT WIM pasny “WT WM pasny IBUIIAJLS IVULIPLSe Y seul ovulluWwAlq oeullg dvVUI[OSse1g ‘seplLAIwg oy} JO sarpruueyqns dy} JO SIa}OBIVYD SsoUdeIP IWOg TZ ATAV EL aseq 38 Ajuo poyeos sates ‘yeom qnio poonpo ers ATIROIS JON poqqnjoun OS YIM a PosOUloO}sSvuy popunol Aj[eqsip meine es pue posso|[) do1y Ayjeseg SBUIIOJIV H “SnoIOWUeJusd G00600000 ruugqUy wereeeee Bd[910J °) SnsievjoIoy senceeee ceceeees ly UIDA SUIMPUTH uoneinsyuoo [[99 SuIMpulH ve UIDA SUIMIIOY Joyorieyo ENT. Soc., 24 AMER. MEM. 18 THE SATYRIDAE of the mid- and hindlegs, if significant; (9) the relative lengths of the midtibia and the first midtarsal subsegment; (10) the presence or ab- sence of dorsal spinosity on the midtibia; (11) the development of tibial spurs, if any; (12) the presence or absence of a dorsal, distal spine at the end of the midtibia; (13) the length and shape of the forewing cell; (14) the branching pattern of the forewing radial veins; (15) the relative positions of the origins of forewing veins Rs and M:; (16) the relative positions of the forewing medial veins; (17) the relative positions of forewing veins Ms, Cui and Cuz; (18) the patterns of inflation of the forewing vein bases; (19) the length and shape of the hindwing cell; (20) the relative lengths of hindwing veins Sct+R: and 3A; (21) the relative positions of hindwing veins Ms and Cur; (22) the relative positions of the hindwing medial veins; and (23) characteristic and aberrant superficial patterns and ihe dis- tribution of androconia. Table 2 gives the character states of certain key characteristics within the satyrid subfamilies. A key to these subfamilies foilows. KEY TO THE SUBFAMILIES OF THE SATYRIDAE 1. Hindwing cell open or closed only by aberrantly placed veins (Figs. 134, IRB IO) iran tontcr asciarenase abe brrice iene reraccodac acdc cecneoscriacoonecaoecdaeodue sccm pcanogey Ragadiinae Hindwing cell closed by tubular veins (Fig. 2, for example) ................-. 2 Forewing with a vestige of vein 3A at base (Figs. 2, 7, 8) ...... Haeterinae Forewing vein 3A fused with 2A throughout its entire length (Fig. 21, for (2.421100 0) (23) ea aera coer eae ce paca eeuonEpS SE aaac CoC ane R sOBencbCH eecabaccedccoscannaoacacencsncriondéiacdaq000 3 3. Hindwing vein R, free basally from Sc + R, (Figs. 9, 90, for example) 4 Hindwing vein R, completely fused with Sc (Fig. 21, for example) ...... 5 4. Forewing cell greater than half length of wing; Neotropical species .......... Ba da aseeactac tea Seca es ee cate a emen leew out pacaasus Eoeak cance Mane sb unreseusuegpaw coe naaeeeecsene Brassolinae Forewing cell much less than half length of wing; Paleotropical species .... BOR eR EN ar a GRO ADCOEEEE Elymniinae (Elymniin:) 5) Posttarsaleclaws bifida (igs 395747) meeeressesscecscossetcncene ses seseaecensssseencneneens 6 Posttarsal@claws simple iectesccsesecccecsessec seca cece sssonceeteoeees sate esccueseanseccneaosenneeee 7 6. Forewing cell more than half length of wing .............. Biinae (Melanitini) Forewing cell less than half length Of Wing ..............::ccccessessssseeeeeeensseeeeeeeeees Puss Tdd vase Tobe esuaaadeawtereretecwaes wasgewterees Elymniinae (Samanta) and Manataria 7. Female foretarsus with fewer than five subsegments ........ many Satyrinae Female foretarsus with five subsegments .............:.cccccccsccssceeeeeeeeeeeeeseeeeees 8 8. Female foretarsus strongly clubbed distad (Fig. 174, for example) .......... REECE CEE ECE EE SOE Sarr pana noobads bans acc bandsobecagesesecranscancraseec000 many Satyrinae Female foretarsus not so clubbed (Figs. 24, 52, 124, for example) ........ 9 N LEE D. MILLER 19 9. Forewing cell half length of wing or less ....................066 most Elymniinae Forewing cell greater than half length of wing ......................sseeeeseeeeeeees 10 10. Forewing cell produced at the origin of M, (Figs. 21, 29, for example); ING WAVWViOnl Gis peciesieer-ccescctece reece oeerer cee receeceeeeneaee New World Biinae Forewing cell produced at the origin of M, (Figs. 121, 127); Indo-Aus- EAL IAMS PECIESeeeanereree eee ec cree ecere renee eee ee aee eae oo sec cn Senses st chat esses Eritinae Subfamily HAETERINAE Herrich-Schaffer, 1864 Hetaerina [sic] Herrich-Schaffer, 1864: 124. = Pierellinae Clark, 1947: 149. The Haeterinae on several counts are the most primitive of the Satyridae. These butterflies cannot be considered close to any other subfamily but show more affinities (albeit slight ones) with the Biinae than with the Brassolinae or Elymniinae. From several aspects — the free proximal end of forewing vein 3A, the general elongate wing form, the seta-like modification of the wing scales and the preference for a deep forest habitat — this subfamily seems to form a connecting link with the Ithomiidae (Fox, 1956: 18-19), but, again, this is a very remote relationship, if not a completely spurious one arising from ecological adaptations. In itself the free proximal vestige of forewing vein 3A serves to distinguish the haeterines from all other satyrids. This characteristic is otherwise shown in the Nymphaloidea by the Ithomiidae, Danaidae, some Morphidae (Taenaris) and such nymphalid genera as Colaenis, Kallima, Apaturina and the very primitive Callinaga. The general shape of the wings (Figs. 2, 7, 8) is characteristic, as is the rounded hindwing cell. The combination of a dorsally spinose midtibia and the lack of tibial spurs (Fig. 6) is shown by no other satyrid. The lack of distal scaling on the antennae is also peculiar to the present subfamily. All members of the Haeterinae are restricted to the American tropics and are found only in the dense forests. A single tribe, the Haeterini, includes all members of the sub- family. The most comprehensive revision of the Haeterinae presently available is that of Brown (1942). MEM. AMER. ENT. SOC., 24 20 THE SATYRIDAE Tribe HAETERINI Herrich-Schaffer, 1864 (Figures 1-8) Hetaerina [sic] Herrich-Schaffer, 1864: 124. = Pierellinae Clark, 1947: 149. The diagnostic features of this tribe are those outlined for the subfamily above. The Haeterini are characterized as follows: The eyes are naked. The antennae vary little in length from slightly over one-third (Pierella) to just over two-fifths (Haetera) the length of the forewing costa. The antennal club is weakly de- veloped, generally less than twice the thickness of the shaft and oc- cupying the distal one-fourth to one-fifth of the antenna. The anten- nae of the haeterines are scaled on only the proximal few segments: the antennae of all other satyrids are fully scaled. The third segment of the palpus is usually less than one-fifth the length of the second, although the third segment is between one-third and one-fourth the length of the second in Haetera and Pseudohaetera. The hairs on the second segment of the palpus are generally about as long as the segment is wide (the hairs are twice as long as the segment is wide in Dulcedo). 0.4 x ML Haetera Cithaerias Pierella | Fig. 1. Haeterinae: Haeterini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements ob- tained from males, the bottom bar those from females. The male foreleg is well developed for a nymphaloid, the tibia is longer than the femur, although both are about equal in Pierella, and there is a single, unspined tarsal subsegment in all genera. The female — foreleg is also well developed with a pentamerous, unclubbed tarsus LEE D. MILLER 21 Figs. 2-8. Haeterinae: Haeterini. 2. Haetera piera (Linné), é venation. 3. H. piera, palpus (in this and all succeeding illustrations of palpi only the distal two segments are figured). 4. H. piera, é foreleg (in this and all suc- ceeding illustrations of the ¢ forelegs, unless otherwise indicated, the femur, tibia and tarsus are figured. 5. H. piera, 2 foretarsus. 6. H. piera, midleg (in this and all succeeding illustrations of the midlegs only the femur, tibia and tarsus are figured). 7. Pierella nereis (Drury), é venation. 8. Cithae- rias pireta (Cramer), ¢ venation. armed with spines on the second, third and fourth subsegments. The mid- and hindlegs are of about equal length. The midtibia is one and one-half to two times as long as the first midtarsal subsegment, is spiny dorsad and lacks tibial spurs or a spine at the dorsal, distal end. The relative development of the legs is shown in Fig. 1. The forewing cell varies from just greater than half to more than three-fifths the length of the forewing costa and is rounded distad in all genera. The forewing radial veins arise in three branches from the cell. Forewing veins Rs and M: are connate or approximate at MEM. AMER. ENT. SOC., 24 DD) THE SATYRIDAE their origins, vein Me arises midway between M: and Msz and vein Cu; arises midway between Mz and Cup, or slightly nearer the former. The forewing vein Sc is inflated in all genera, and the other veins may be: both the cubital stem and 2A in Pierella, only 2A in Haetera. The presence of a proximal vestige of forewing vein 3A is diagnostic in the Haeterinae. The hindwing cell is also distally rounded and is from two-fifths (Pierella) to three-fifths (Cithaerias) as long as the wing, as measured to the origin and end of Ms, respectively. Hindwing veins Sc+Ri and 3A are of about the same length. The origins of hindwing veins M:; and Cu; may be widely separated (Cithaerias, Fig. 8) to connate on a common stalk (Pierella, Fig. 7). Wein Me arises about midway between M, and Ms, or slightly nearer the latter. The pattern is distinctive (see Schwanwitsch, 1925): the scales of most species are reduced to narrow, hairlike vestiges, resulting in a hyaline or translucent appearance in most species. Relatively un- modified scales are displayed by some species, particularly of the genera Pierella and Haetera, with resultant opaque wings. Genera Included in the Haeterini Callitaera Butler, 1868b: 101. Type-species: “Papilio aurora Felder” (= Ci- thaerias phantoma Fassl, fide Clifton, in litt.), designated by Brown, 1942: 311. Cithaerias Hiibner, [1819] (1816-1826): 53. Type-species: “Papilio andro- meda Fabricius” (= Papilio pireta Cramer, according to Brown, 1942), designated by Scudder, 1875a: 143. Papilio menander Drury, often regarded as the type-species, was shown by d’Almeida (1951: 496) to be identical with Papilio pireta Cramer. Dulcedo d’Almeida, 1951: 501. Type-species: Haetera polita Hewitson, by original designation. Haetera Fabricius, 1807: 284. Type-species: Papilio piera Linné, designated by Butler, 1868a: 195. = Oreas Hiibner, [1807] (1806-1838): pl. [82]. Type-species: Pa- pilio piera Linné, by monotypy. Oreas is sunk to Haetera Fabri- cius in accordance with Opinion 137 (1942) of the International Commission of Zoological Nomenclature. Pierella Herrich-Schaffer, 1865: 65. Type-species: Papilio nereis Drury, desig- nated by Butler, 1868a: 195. Pseudohaetera Brown, 1942: 330. Type-species: Haetera hypaesia Hewitson, by original designation. LEE D. MILLER 23 Subfamily BRASSOLINAE Boisduval, 1836 Brassolides Boisduval, 1836: 166. The brassolines have long been considered a separate family somewhat intermediate between the satyrids and the morphids, but Ehrlich (1958: 351-352) placed the brassolines with the morphids. There is little except the large size of the butterflies to support this combination: the larvae of most Neotropical morphids feed on dicots, whereas the brassolines feed exclusively on monocots; brassoline lar- vae have bifid tails, but the tails are greatly reduced in the morphids. These two characters intimately associate the brassolines with the other satyrids. The cells of both wings are closed in the brassolines, as in the satyrids generally, whereas the hindwing cell is open in most of the Morphidae. Within the Satyridae the brassolines are allied to the New World Biinae, particularly through such genera as Narope. There seems little justification, in view of the above facts, for retain- ing the brassolines as a family separate from the Satyridae. The brassolines may be distinguished from all other New World Satyridae by the basal separation of hindwing veins Sc and Ri (Figs. 9, 13, 14, 17, 19). This characteristic is also shown by the Old — World Elymniini (Elymniinae), but the extremely short cell and vir- tually connate veins Mz and Cu: of the forewing immediately disting- uishes the elymniines. The Haeterinae show an approximate condi- tion of hindwing veins Sc + Ri and Rs, but there is no basal separation of Ri. The brassolines lack the vestige of forewing vein 3A which characterizes the Haeterinae. All brassolines are Neotropical, none being found further north than southern Mexico. Some species of Calizo are banana pests in Central America, and around the banana plantations these insects often may be seen flying just at dusk. Clark (1947, 1948) recognized two tribes as subfamilies of “Bras- solidae”, but no good evidence is found for splitting the brassolines into more than a single tribe, the Brassolini. Within the Brassolini groupings are possible, but not tribal ones. Tribe BRASSOLINI Boisduval, 1836 (Figures 9-19) Brassolides Boisduval, 1836: 166. = Caliginae Clark, 1947: 149. MEM. AMER. ENT. SOC., 24 24 THE SATYRIDAE Figs. 9-19. Brassolinae: Brassolini. 9. Brassolis sophorae (Linné), ¢ venation. 10. B. sophorae, palpus. 11. B. sophorae, 6 foreleg. 12. B. so- phorae, midleg. 13. Opsiphanes cassiae (Linné), ¢ venation (after Schatz and Rober, 1892). 14. Caligo atreus Kollar, é venation (after Schatz and Rober, 1892). 15. C. eurylochus (Cramer), 2° foretarsus. 16. C. eurylochus, midtarsus and distal portion of midtibia. 17. Dasyophthalma rusina (Godart), é venation. 18. D. rusina, 2 foretarsus. 19. Narope cyllastros Doubleday and Hewitson, ¢ venation. The characteristics of the subfamily are those of the tribe. Four groups may be noted within the tribe, but they are only loose as- semblages and tend to grade into one another. The diagnostic char- acters are summarized in Table 3. i) On LEE D. MILLER The Brassolis-series, sensu stricto, is composed of robust species with naked eyes, moderately well developed or absent tibial spurs and a basically brown upper surface with a transverse band of orange, yellow or red across the forewing. The Caligo-series includes large, broad-winged, but not robust- bodied butterflies with naked eyes, well developed tibial spurs and a basically iridescent purple upper surface with faint longitudinal light markings. The Dasyophthalma-series is comprised of a single genus of broad- winged species with hairy eyes, moderately well developed tibial spurs and a basically brown upper surface, with or without iridescence, and with longitudinal lighter markings. The Narope-series is also comprised of a single genus of medium- sized butterflies with acute forewing costal margins, naked eyes, mod- erately well developed tibial spurs and a brown upper surface with little or no maculation. TABLE 3 The character states of four characters in the genera of the Brassolinae: Brassolini. Length of fore- wing cell / Antennal Tibial length of Genus Eyes club spurs forewing costa Brassolis-series, s. str. Brassolis Naked Pronounced Present, ex. 3/5-2/3 B. sophorae Catoblepia Naked Gradual Present 3/5-2/3 Dynastor Naked Gradual Present 1/2-3/5 Opoptera Naked Gradual Present 3/5-2/3 Opsiphanes Naked Gradual Present 1/2-3/5 Penetes Naked Gradual Present 3/5-2/3 Selenophanes Naked Gradual Present 3/5-2/3 Caligo-series Caligo Naked Gradual Present 3/5-2/3 Ery phanis Naked Gradual Present 3/5-2/3 Dasyophthalma-series Dasyophthalma Hairy Gradual Present 3/5-2/3 Narope-series Narope Naked Gradual Present just >1/2 MEM. AMER. ENT. SOC., 24 26 THE SATYRIDAE The Brassolini are characterized as follows. — The eyes are naked in all genera but Dasyophthalma in which they are densely hairy. The antennae are short, one-third to two-fifths the length of the fore- wing costa, and the antennal club is weakly developed in most genera (just over twice the width of the shaft in Brassolis). The third seg- ment of the palpus is generally one-fourth to one-sixth the length of the second, although it is far shorter in such rather advanced genera as Brassolis and Opsiphanes and somewhat longer in the more primi- tive Narope. The hairs of the second segment of the palpus are gen- erally shorter than the segment is wide. The male foreleg is well-developed with a mono- to trimerous tar- sus; the femur is longer than the tibia in all genera. The female fore- leg is likewise well developed with a pentamerous, unclubbed tarsus bearing spines on at least the third and fourth (often the first and second, too) subsegments. The midtibia varies from just less than twice (Caligo and Eryphanis) to well over twice (Brassolis-series ) as long as the proximal midtarsal subsegment. The midtibia is spiny dorsad with well to moderately well developed tibial spurs (except in Brassolis sophorae, where the tibial spurs are totally wanting, Fig. 12); there is no spine at the dorsal, distal end of the midtibia. The forewing cell is produced at the origin of Ms, particularly in Caligo and Eryphanis. The cell is generally three-fifths to two-thirds the length of the forewing costa, although it is somewhat shorter in Dynastor, Opsiphanes and Narope. The forewing radial veins arise from the cell in three branches. Forewing veins Rs and Mi are well separated at their origins; vein Me usually arises nearer Mi than Ms, although Mp2 arises nearer M3 in Caligo and Dynastor; vein Cui arises nearer M; than Cuz, except in Dasyophthalma. The forewing veins are seldom inflated, though the base of Sc may be in some genera. The hindwing cell is produced marginad by a migration of cross- vein My-ms and is generally about three-fifths the length of the wing, measured to the origin and the end of Ms, respectively. Hindwing veins Sc and Ri are separate proximally in this tribe, as in the Elym- niini (Elymniinae); Sc+R: is longer than 3A. At their origins veins Ms and Cu; are well separated, and Mz arises nearer M; than Mi. The upper surface patterns have been described in the analysis of the generic series. On the under surface the wings are cryptically patterned, light brown with darker striations, much as in the Antir- LEE D. MILLER Di thini. In the brassolines, however, there are often large “eye-spots”, particularly on the hindwing, which have been credited with being protective markings (the wings are rapidly opened when the butter- fly is threatened by a predator, giving the impression of large eyes blinking). The development of these eye-spots gives the common name of the Caligo-series, the “owl butterflies’. Genera Included in the Brassolini The Brassolis-series, sensu stricto Brassolis Fabricius, 1807: 282. Type-species: Papilio sophorae Linné, desig- nated by Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 341. Catoblepia Stichel, 1902: 488, 491. Type-species: Brassolis amphiroe Hiibner, by original designation. Dynastor Westwood and Hewitson, 1849, in Doubleday, Westwood and Hewit- son, 1846-1852: pl. 58. Type-species: Dynastor napoleon Westwood and Hewitson, by monotypy. Opoptera Aurivillius, 1882: 75. Type-species: Brassolis syme Hubner, by origi- nal designation. Opsiphanes Westwood and Hewitson, 1849, in Doubleday, Westwood and Hew- itson, 1846-1852: pl. 57. Type-species: Opsiphanes sallei Westwood and Hewitson, designated by Scudder, 1875a: 233. Penetes Westwood and Hewitson, 1849, in Doubleday, Westwood and Hewit- son, 1846-1852: pl. 58. Type-species: Penetes pamphanis Westwood and Hewitson, by monotypy. Selenophanes Staudinger, 1888: 212. Type-species: Papilio cassiope Cramer, by original designation. The Caligo-series Caligo Hiibner, [1819] (1816-1826): 51. Type-species: Caligo eurylochus Hubner, designated by Scudder, 1875a: 129. Eryphanis Boisduval, 1870: 57. Type-species: Papilio automedon Cramer, by monotypy. Dasyophthalma-series Dasyophthalma Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 343. Type-species: Brassolis rusina Godart, designated by Scudder, 1875a: 155. MEM. AMER. ENT. SOc., 24 28 THE SATYRIDAE Narope-series Narope Westwood and Hewitson, 1849, in Doubleday, Westwood and Hewit- son, 1846-1852: pl. 50. Type-species: Narope cyllastros Westwood and Hewitson, by monotypy. Subfamily BIINAE Herrich-Schaffer, 1864 Biina Herrich-Schaffer, 1864: 124. The Biinae are the third of the very primitive satyrid groups and show no especially close relationship with any other subfamily. The Neotropical biines share some vague affinities with the haeterines and some firmer ones with the brassolines, particularly as regards vena- tion, whereas the Paleotropical biines grade toward the Lethini of the next subfamily. The biines are generally large to very large satyrids, often brightly colored above and always cryptically patterned beneath with few and usually poorly developed eye-spots on either wing surface. The long forewing cell, generally three-fifths the length of the wing except in some melanitines (Fig. 34), is characteristic, and the well separated hindwing veins Mz and Cu: serve to distinguish these butterflies from most elymniines. The wings are never translucent or transparent, as in the haeterines. TABLE 4 Some of the diagnostic characters of the tribes of the Biinae. Character Antirrhini Biini Melanitini MarsalliclawSiecesecesseeseseteeetes Simplerce..-ss-2 cee: Simpleteeses.00 Bifid Mibsalispurspeenees sees seceeessee Present, but ...... ADSENt ieee Present poorly devel- oped Length of 3rd seg. of palpus relative to length of second .............. Gills] AD ost ea tae Stine eee eee 1/4 Forewing radial veins .......... 3 branches ........ [branchigie.-< sc. 3 branches aberrant Inflation of forewing veins .. Only Sc, if any; .. Sc, Cu, 2A ........ Only Sc, slight or else v. slight greatly inflated LEE D. MILLER 29 All biines are tropical. The Antirrhini and Biini are found ex- clusively in the Neotropics and are forest dwellers, whereas the Mel- anitini are Paleotropical (from West Africa to Australia) and are found in less dense situations, though many of these species are cre- puscular. The Biinae are split, somewhat reluctantly, into three tribes, but Bia is far too aberrant to be referred to either of the other two. The diagnostic features of the three tribes are given in Table 4, and a key to these tribes is given below. KEY TO THE TRIBES OF THE BIINAE 1. Tarsal claws bifid (Fig. 39); Paleotropical species ..............006 Melanitini Marsaluclaws)simple:s NeotropicalispeCies ae-ccceeste-ne-cnccese-cceeesessessacneseeess 2 Tibial spurs present, though poorly developed (Fig. 25); third segment of palpus less than one-fourth length of second segment (Fig. 22) ............ nN OS O EOP CCC EEE ES Ce UGC ECG NSC OSSD SEO EEL CSOECCEEEE PERRET een eae CEE Antirrhini Tibial spurs absent (Fig. 33); third segment of palpus greater than half lensthvofssecondsseoments (ip s95,0))ieeessseessecccccccccccceececeseesrececceeeeee Bini The New World members of the Biinae are considered to be more primitive because of (1) their greater convergence toward the Bras- solinae and Haeterinae, (2) the greater relative size of the male fore- leg and (3) the unclubbed female foreleg. The Melanitini are the more advanced members of the subfamily and seem to show relation- ships with the more advanced Elymniinae. Tribe ANTIRRHINI, new tribe (Figures 20-28) The simple tarsal claws serve to separate this New World tribe from the Old World Melanitini. Many characteristics serve to sep- arate the Antirrhini from the Biini, notably the presence of tibial spurs, the more reduced third segment of the palpus, the arising of the forewing radial veins in three branches and the relative placement of several other veins on the cells. The Antirrhini are Neotropical insects, found from Guatemala to Paraguay and northern Argentina. All seem to be forest species, seldom venturing into open areas. The tribe Antirrhini is character- ized as follows: The eyes are naked in all genera but Sinarista, in which they are sparsely hairy. The antennae are about half as long as the forewing MEM. AMER. ENT. SOC., 24 30 THE SATYRIDAE 0.4xML FL ML HL |xML ANTIRRHINI Antirrhea Caerois Triteleuta Bia Melanitis Hipio Cyllogenes Parantirrhoea Fig. 20. Biinae: Antirrhini, Biini and Melanitini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all in- stances the ML value is unity. The top bar for each genus represents the mea- surements obtained from males, the bottom bar those from females. costa, and the antennal club is weakly developed (it is completely in- distinct in Triteleuta). When the club is distinct it occupies the dis- tal one-third to one-quarter of the antenna and is seldom more than twice as thick as the shaft. The third segment of the palpus is one- fourth to one-sixth the length of the second segment, and the hairs of the second segment are about as long as the segment is wide, although the hairs of Sinarista are about twice as long as the width of the seg- ment. The male foreleg is well developed with a monomerous, unspined tarsus, and the tibia is usually slightly longer than the femur, although the femur is much longer in Caerois. The female foreleg is also well ° developed with a pentamerous, unclubbed tarsus bearing spines on the first four subsegments. The midtibia is more than twice as long LEE D. MILLER 31 as the basal midtarsal subsegment, smooth dorsally and bearing weakly developed tibial spurs. There is no spine at the dorsal, distal end of the midtibia. The relative lengths of the legs are shown in Fig. 20. The forewing cell is square-cut in Caerois and Sinarista, but deeply excavate at Mi in the other genera. The cell is about three-fifths the length of the forewing costa. The radial veins arise in three branches from the forewing cell, and veins Rs and M:; are connate to approxi- mate at their origins. Forewing vein M2 arises midway between Mi and M3, whereas Cu: usually arises about midway between M3; and Cuz (nearer Mz in Caerois). The forewing veins are only slightly inflated, if at all; any thickening is at the base of Sc. The hindwing cell is highly variable, from very short and narrow in Antirrhea to over three-fifths the length of the wing, measured to the end of Ms, and rounded in Caerois. Hindwing vein Sc+R: is usually longer than 3A, although the two veins are of about the same length in Sinarista. Hindwing veins Ms; and Cu: are generally well separated at their origins, but they are approximate in Antirrhea; vein Mb arises midway between M; and Ms. Most of the species are rather brightly colored above with few ocelli, although some species, such as Sinarista adoptiva Weymer, have rows of submarginal ocelli. On the under surface all species are cryptically marked with many fine, dark striations on a tan to brown ground color. There are androconial hair patches on the under surface of the forewing along vein 2A in all genera but Sinarista, and these androconial areas frequently result in distortions of the venation. Genera Included in the Antirrhini Antirrhea Hiibner, [1822] (1806-1838): pl. 294. Type-species: Antirrhea ar- chaea Hubner, by monotypy. = Anchiphlebia Butler, 1868b: 106. Type-species: Antirrhea archaea Hubner, by original designation. Caerois Hiibner, [1819] (1816-1826): 56. Type-species: Papilio arcesilaus Cramer (= Papilio chorinaeus Fabricius), by monotypy. = Arpidea Duncan, 1837: 180. Type-species: Papilio chorinaeus Fabricius, by monotypy. Sinarista Weymer, 1909: 164. Type-species: Sinarista adoptiva Weymer, by monotypy. Triteleuta Strand, 1912: 44. Type-species: Antirrhea tomasia Butler, by origi- nal designation. Proposed as a subgenus of Antirrhea Hiibner. MEM. AMER. ENT. SOC., 24 32 THE SATYRIDAE 22 pee y trope Wh YY Wiley 7 pe Figs. 21-28. Biinae: Antirrhini. 21. Antirrhea archaea Hiibner, 4 vena- tion. 22. A. archaea, palpus. 23. A. archaea, ¢ foreleg. 24. A. archaea, 2 foretarsus. 25. A. archaea, midleg. 26. Caerois chorinaeus (Fabricius), ¢ venation. 27. C. chorinaeus, ¢ foreleg. 28. Sinarista adoptiva Weymer, ¢ venation. Tribe Bruni Herrich-Schaffer, 1864 (Figures 20, 29-33) Biina Herrich-Schaffer, 1864: 124. Bia actoriaena (Linné), the only species included in the Biini, is the most aberrant member of the Satyridae. It has been considered variously a brassoline (Weymer, 1912; Clark, 1947, 1948) or a per- fectly good satyrid (R6ber, 1892; Ehrlich, 1958). Bia is definitely LEE D. MILLER 33 a satyrid and shows relationships to other members of the Biinae, chiefly through pattern convergence, but it is far too aberrant to be included in either of the other two biine tribes. This species can be distinguished from all other biines by the lack of tibial spurs, in addi- tion to a multitude of other characteristics enumerated below in the description of the tribe. Bia is primitive, though highly specialized, and may be characterized as follows: The eyes are naked. The antennae are half as long as the fore- wing costa, and the club is poorly developed occupying the distal quarter of the antenna and about twice as thick as the shaft. The third segment of the palpus is very long, over half the length of the second segment, and the hairs of the second segment are not as long as the segment is wide. The male foreleg is well developed, with the femur longer than the tibia and a monomerous, unspined tarsus. The female foreleg is also well developed with a pentamerous, unclubbed tarsus bearing spines on the first four subsegments. The midtibia is smooth dorsally, more than twice as long as the proximal midtarsal subsegment and armed with neither tibial spurs nor a spine at the dorsal, distal end. The forewing cell, measured to the origin of Ms, is greater than three-fifths the length of the forewing costa, but the cell is deeply ex- cavate at the origins of Mi and Mb, and the cell is only about half the length of the forewing costa when it is measured to the origin of Mi. All the forewing radial veins arise from a single branch from the cell, and Ri anastomoses distally with Sc, resulting in a confused venational pattern along the costa. Forewing veins Rs and Mi: are well separated at their origins. Wein Me arises much nearer Mi; than M3, whereas Cu: arises midway between Mz; and Cus. Veins Sc, the cubital stem and 2A are all strongly inflated at their bases, the in- flation of the latter suggesting the pattern shown in the Mycalesini of the next subfamily. The hindwing cell is strongly produced at the origin of Mz and about three-fifths the length of the wing measured to the end of Ms. Hindwing vein Sc+Ri is longer than 3A. Veins Ms and Cu: are well separated at their origins, and Me arises nearer M: than Ms. The upper surface is dark brown with patches of orange and iri- descent blue scales. On the under surface Bia is marked like other members of the subfamily: tan with numerous, longitudinal brown MEM. AMER. ENT. SOC., 24 34 THE SATYRIDAE 30 3] 39 Figs. 29-33. Buinae: Biini, all Bia actoriaena (Linné). 29. é venation. 30. palpus. 31. ¢ foreleg. 32. 2 foretarsus. 33. midleg. striae. The androconial areas on the hindwing are characteristic: there is a patch of mealy scales on the upper end of the cell along crossvein r.-mi, and a long hair tuft lies along 2A. Genus Included in the Biini Bia Htibner, [1819] (1816-1826): 51. Type-species: Papilio actoriaena Linné, by monotypy. Tribe MELANITINI, new tribe (Figures 20, 34-40) The melanitines are the most advanced members of the Biinae, as noted above, particularly with regard to the more reduced male fore- leg and the slightly clubbed female foretarsus (Fig. 37). There are, in addition to the already-mentioned affinities to the Antirrhini, some rather suggestive ties between the Melanitini and the Lethini of the next subfamily, especially through the genera Parantirrhoea (Mela- nitini) and Ptychandra and Samanta (Lethini). The venation of Ptychandra and its pattern relate rather closely (but with modifica- tions) to those of the Melanitini, and Samanta displays the bifid claws characteristic of the melanitines. Bifid claws are the most character- _ istic feature of this tribe and serve to distinguish it from all other satyrids except Samanta and Manataria (a Neotropical genus of un- certain position). LEE D. MILLER 35 All members of this tribe are Paleotropical, being found from West Africa through the Malay Peninsula east as far as New Guinea and northern Australia. At least some of the species are crepuscular, and they do not appear to be so limited to the deep forest as are the Neotropical Antirrhini. The Melanitini are characterized as follows: The eyes are naked. The antennae are rather uniformly just over two-fifths the length of the forewing costa, and the antennal club is weakly developed, occupies the distal quarter of the antenna and is thickened twice to two and a half times the thickness of the shaft. The third segment of the palpus is about one-fourth the length of the second, and the hairs of the second segment are much shorter than the segment is wide. Figs. 34-40. Biinae: Melanitini. 34. Melanitis leda (Linné), 6 venation. 35. M. leda, palpus. 36. M. leda, 6 foreleg. 37. M. leda, 2 foretarsus. 38. M. leda, midleg. 39. M. leda, bifid tarsal claw. 40. Hipio constantia (Cramer), venation of proximal portion of 6 wings. The male foreleg is well developed with the femur and the tibia of about the same length (the femur is longer in Cyllogenes and Parantirrhoea) and bearing a single, usually unspined (weakly so in Cyllogenes) tarsal subsegment. The female foreleg is also well de- veloped with a pentamerous, unclubbed or weakly clubbed tarsus. The fourth tarsal subsegment always bears spines, and the spines are MEM. AMER. ENT. SOC., 24 36 THE SATYRIDAE present on the third and fourth subsegments in Gnophodes and Par- antirrhoea, on the second, third and fourth in Melanitis and Hipio. The midtibia is greater than twice as long as the proximal midtarsal subsegment, is smooth dorsad and bears moderately well developed tibial spurs but no spine at the dorsal, distal end. Relative lengths of legs are shown in Fig. 20. The forewing cell varies from just over half to nearly three-fifths as long as the forewing costa and is somewhat excavate at me2-ms. The forewing radial veins arise in three branches. Veins Rs and Mi are usually well separated at their origins, vein Mz arises much closer to M, than to Mz and Cui usually arises midway between M3 and Cup, although Cu: arises noticeably nearer Cuz in Melanitis. The forewing veins are little inflated; any inflation is restricted to the base of Sc. The hindwing cell is more or less rounded distad and is two-fifths to half the length of the wing measured to the end of Ms. The hind- wing vein Sc+R: is longer than 3A. Hindwing vein Mb arises mid- way between Mi and Ms, and veins M3 and Cu: are well separated at their origins. The pattern is similar to that of the Antirrhini with a shaded brown and orange upper surface and cryptic coloration on the under surface with a brown ground color and fine, dark striations. Genera Included in the Melanitini Cyllogenes Butler, 1868b: 6. Type-species: Melanitis suradeva Moore, by monotypy. Gnophodes Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 1852: 363. Type-species: Gnophodes parmeno Westwood, designated by Butler, 1868a: 194. Hipio Hiibner, [1819] (1816-1826): 56. Type-species: Papilio constantia Cra- mer, designated by Butler, 1867a: 279. Melanitis Fabricius, 1807: 282. Type-species: Papilio leda Linné, designated by Butler, 1868a: 194. = Cyllo Boisduval, 1832: 140. Type-species: Papilio leda Linné, designated by Scudder, 1875a: 151. Parantirrhoea Wood-Mason, 1880: 248. Type-species: Parantirrhoea mar- shallii Wood-Mason, by monotypy. Genus MANATARIA Kirby, 1908 (Figures 41-47) Manataria Kirby, 1908: 57. Type-species: Tisiphone hercyna Hubner, by monotypy. LEE D. MILLER 37 Manataria 47 Figs. 41-47. Manataria hercyna (Hiibner). 41. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling). The ML value is unity. The top bar represents the measurements obtained from the male, the bottom bar those from the female. 42. ¢ venation. 43. palpus. 44. ¢ foreleg. 45. 9 foretarsus. 46. midleg. 47. bifid tarsal claw. This South American genus has some morphological and pattern features which connect it to the Neorina-series of the Lethini. If this were its position, Manataria would be the only Neotropical lethine and not particularly closely related to the few Nearctic members of the tribe. The bifid tarsal claws, on the other hand, connect the pres- ent genus with the sympatric Antirrhini (although this is through the Paleotropical Melanitini), but in most other respects Manataria has little in common with the antirrhines. The paitern is quite typical of the Neorina-series, even to the basad displacement of the ocellus in hindwing space Rs-Mi of the under surface. Were it not for the vast geographical isolation and the bifid tarsal claws I would place Mana- taria in the Lethini, perhaps in a monotypic series, but it seems more advisable to call attention to this aberrant genus here without assign- ing it to a tribe, pending further information. Manataria is char- acterized as follows: MEM. AMER. ENT. SOC., 24 38 THE SATYRIDAE The eyes are naked. The antennae are about half the length of the forewing costa; the antennal club is poorly developed and less than twice as thick as the shaft. The third segment of the palpus is just under one-third the length of the second. The hairs of the second palpal segment are not as long as the segment is wide. The male foreleg is well developed with the tibia slightly longer than the femur and a monomerous, unspined tarsus. The female foreleg is also not significantly miniaturized with a pentamerous, un- clubbed tarsus which is spined on the second, third and fourth sub- segments. The midleg is slightly longer than the hindleg (Fig. 41). The midtibia is just less than twice as long as the proximal midtarsal subsegment, is slightly spinose dorsad, bears no spine at the dorsal, distal end and has well developed tibial spurs. The posttarsal claws are bifid. The relative lengths of the legs are shown in Fig. 41. The forewing cell is produced at the origin of Ms and about half as long as the forewing costa. The forewing radial veins arise in three branches from the cell, and Rs and M: are connate. Vein Me arises nearer M, than Ms, and Cuz arises slightly nearer M3 than Cue. Forewing veins Sc and the cubital stem are slightly inflated; 2A is not. The hindwing cell is produced at the origin of Ms and is slightly more than two-fifths the length of the wing measured to the end of M;. Veins Sc+Ri and 3A are of about the same length, and Ms and Cui are separate, but approximate, at their origins. Vein Me arises much nearer M: than Ms. The upper surface is dark brown with a transverse yellow spot- band on the forewing outside the cell. Below the pattern of the upper surface is repeated; in addition, there are pale scrawlings at the apex of the forewing and all over the hindwing and ocelli are developed in space Mi-Mb of the forewing and in all spaces from Rs-Mi to Cu2-2A of the hindwing. The ocellus in hindwing space Rs-Mi; is displaced basad as in most of the Lethini. Subfamily ELYMNIINAE Herrich-Schaffer, 1864 Eurytelidae Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 1852: 403 (in part). Elymniina Herrich-Schaffer, 1864: 124. = Enodiinae Clark, 1947: 149. = Lethinae Clark, 1948: 77. LEE D. MILLER 39 The Elymniinae are the now dominant primitive satyrid subfamily and apparently provided the source for the Satyrinae, Eritinae and probably the Ragadiinae — although the origin of the latter subfamily is still a problem. In some respects the elymniines may be connected to, and may have arisen from, the Melanitini (Biinae) through such genera as Ptychandra and Samanta (Lethini), as discussed under the melanitines. Although not all Elymniinae show this characteristic and some that are not elymniines do, most satyrids with the hindwing veins Ms and Cu: connate or closely approximate at their origins are probably referable to this subfamily. The form of the hindwing cell is also distinctive: it is usually greatly produced marginad at the origin of vein Ms, although this is not true of some Elymniini because of dis- placement of the venation by the androconial patch. The presence of androconial patches in specific locations is characteristic of the tribes. With the exception of a few lethine genera of the Pararge-series, the forewing cell is never more than half the length of the wing, in marked contrast with the preceding three subfamilies where the cell is seldom TABLE 5 Some of the diagnostic characters of the tribes of the Elymniinae. Character Lethini Zetherini Elymniini Mycalesini Tibial spurs .. Present ............ BTeSeM twee Absent ............ Present LENE Geooctensoreee Hairy, except .. Naked ............ INaked® c--: Naked or hairy in Neorina- series Forewing WEIN AN coo LHS, TE BLE” coococ Not inflated .... Not inflated .... Subquadrate all, inflated basal infla- tion, except two genera Forewing me- dian veins .. Ma, arises ........ Mg, arises much Mg, arises much M, arises much nearer M, nearer M, nearer M, nearer M, than Mz than M. than Ms than M. Forewing cell Rounded, ca. .. Slightly exca- .. Strongly exca- .. Strongly exca- 1/2 length vate, <1/2 vate, <1/2 vate, ca. 1/2 of forewing length of length of length of forewing forewing forewing MEM. AMER. ENT. SOC., 24 40 THE SATYRIDAE much less than three-fifths the length of the forewing costa. The male foreleg is moderately well developed in all genera and is characteristi- cally tapered to a rather sharp point. The metropolis of the Elymniinae is the Paleotropical area, and three of the four tribes are found more or less exclusively there. The other tribe, the Lethini, reaches its greatest development in the Indo- Malayan region but has representatives in Africa, the Palearctic and two genera in North America. The tribes of the Elymniinae all more or less grade into one an- other. They are the Lethini, Zetherini, Elymniini and Mycalesini. It is difficult to determine which of the tribes is primitive and which is derived, if any are, but the Aeropetes-series of the Lethini are defin- itely primitive, whatever the condition of the other members of the tribe; the other tribes have about equal status. A key to the tribes of the Elymniinae is given below and an analysis of them in Table 5. KEY TO THE TRIBES OF THE ELYMNIINAE 1) Tibial spurs absent (Bigs 96) ee weeccceereest se erences sen ene eee Elymniini Tibial spurs present (Fig. 53, for example)... ee eeeceeeeeessseeeeeeeeees Z 2. Forewing vein 2A inflated at base to a subquadrate thickening (Figs. 99, itil aya) (0) eat=e. @-0 01) 0) (OD on pacarecetot ced tanosassoedaconcsr ger osaandesoodsocas sae most Mycalesini Forewing vein 2A, if thickened, not subquadrate at base ................. 3 3. Eyes hairy; forewing cell frequently rounded distad (Fig. 49, for example) eA Maectcchiscae ovtre sei ar NeMederamall uns Nten na Cae Lethini (except Neorina-series) Eyes naked; forewing cell frequently excavate (Fig. 90, for example) .. 4 4. Base of forewing vein Sc about three times the thickness of any other Stalks (Bip l06) teense Mycalesini (Orsotriaena, Bletogona) Bases of forewing veins little inflated, Sc seldom over twice the thickness Of any. Other Ve cee eeiiiecdecc cus cacdecehones vente ctecs sees skiceegiossee aaa ease eae 5 5. Forewing vein M, arising much nearer M, than Ms (mj,-m, one-fourth or less as long as Moy-ms; Figs. 81, 86, 87) oc... ceceeeeeeeseeeeeeeeeeeeeees Zetherini Forewing arising nearer, but not much nearer, M, than Mz ......... eee ses esi ah aden cul eae Heer ye asa Sab TMU tomas ee Oe ee a Lethini (Neorina group) Tribe LETHINI Clark, 1948 (Figures 48-80) Enodiinae Clark, 1947: 149. Lethinae Clark, 1948: 77. “Lethinae” is used in preference to the older “Enodiinae” for a variety of reasons. Enodia is generally considered at best a subgenus LEE D. MILLER 41 of Lethe, but this is an invalid reason for the suppression of a family- group name, according to Art. 40 of the International Code of Zoo- logical Nomenclature (1961). Part (a), however, of the same Ar- ticle states, “If a family-group name, changed before 1961 because of such synonymy, has won general acceptance, it is to be maintained in the interests of stability.” This is precisely applicable to the ques- tion at hand. Dos Passos (1964: 99) uses the collective name “Lethinae” in preference to the older “Enodiinae”. To now replace Lethinae (or in this instance, “Lethini”) with Enodiinae (Enodiini) would not serve the interests of stability of nomenclature. Lethe is further to be preferred, all other things being equal, inasmuch as this genus is recognized throughout much of the world; Enodia, where it is even recognized today, is known only in the eastern part of North America. The Lethini may be the focal point of much of the evolution of the higher categories of the Satyridae. As mentioned earlier in the discussion of the Melanitini, there are some vaguely but intriguingly indicated connections between that group and the lethines through such lethine genera as Ptychandra and Samanta. The venation of the former genus, while highly modified by androconial displacement of the forewing veins, is suggestive of the pattern in the melanitines (Fig. 57). Samanta, while a perfectly typical lethine from almost all other aspects, does show the bifid tarsal claws which characterize the Melanitini and the Neotropical aberrant genus Manataria. With- in the Elymniinae, too, some interesting gradations may be seen. The line separating the Zetherini from the Lethini is a thin one, based chiefly on the relative positions of some veins and naked versus hairy eyes, respectively. The zetherines, in turn, have many characteristics in common with the Elymniini, as will be shown below. The genus Mandarinia is a puzzle: the distribution of androconia suggest that it is a mycalesine, but the lack of inflation of the forewing vein 2A and the distally rounded forewing cell are lethine characteristics: this genus is provisionally placed in the Lethini (Fig. 73). The African mem- bers of the Aeropetes-series show yet another relationship. If one considers these butterflies and members of the satyrine tribe Dirini together, one can almost obtain a graded series from perfectly good Lethini to typical Satyrinae. This is probably convergence, as will be shown later. MEM. AMER. ENT. SOC., 24 42 THE SATYRIDAE The Lethini are generally characterized by the hairy eyes which separate them from the Elymniini and the zetherines, the presence of tibial spurs (a characteristic of the Elymniini is the lack of tibial spurs) and by the more or less rounded forewing cell, never exca- vate as in the elymniines and most mycalesines. Most Lethini show a characteristic basad displacement (diastoma) of the hindwing ocel- lus in space Mi-M». There are a number of generic series, however, so it is difficult to distinguish all lethines by a single set of criteria. The Aeropetes-series, consisting of two genera, is confined to South Africa and characterized by the large size, extremely primitive pattern (Schwanwitsch, 1924), hairy eyes, connate hindwing veins Mp» and Cui, extremely spiny midtibia and lack of androconial patches (Figs. 67-72). The Lethe-series, sensu stricto, has twenty-five nomenclatorially, though not necessarily biologically, valid genera, twenty-two of which are Indo-Australian or sub-Palearctic, one East African and two North American. These insects are medium to large-sized, have a rather primitive pattern on the under surface, though not necessarily on the upper, hairy eyes, connate to separate hindwing veins M3 and Cu, dorsally smooth to only slightly spiny midtibia and frequently with androconial patches in the usual positions (Figs. 49-59, 66). The Pararge-series has nine Palearctic genera which differ from the Lethe-series chiefly in the length of the forewing cell: in all other groups the cell is about half the length of the wing, whereas in Par- arge and its allies it is about three-fifths as long (Figs. 60-65). The Mandarinia-series is monotypic, and is characterized by its Mycalesis-like androconial area and is found exclusively in China (Figs. 73-76). The Neorina-series is composed of four Indo-Australian genera. These butterflies are characterized by their large size, highly modified pattern, naked eyes, spiny midtibia and lack of androconia (Figs. 77-80). The Lethini are characterized as follows: The eyes are hairy in all members except those of the Neorina- series, which have naked eyes. The antennae are generally from two- fifths to nearly half as long as the forewing costa, but the antennae are slightly longer than half the costa in such genera as Hermias, © Tansima and Pararge. The antennal club is moderately well devel- LEE D. MILLER 43 0.4xML, FL ML HL [xML Aeropetes | series Aeropetes iil oo Paralethe Lethe |series, S. Ss. Lethe Aphysoneura Satyrodes Enodia Zophoessa Ptychandra | | / See | Pararge| Series | Pararge Orinoma Kirinia | Mandarinia | series Mandarinia Neorina| series Neorina Anadebis a Fig. 48. Elymniinae: Lethini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements ob- tained from males, the bottom bar those from females. MEM. AMER. ENT. SOC., 24 44 THE SATYRIDAE oped, occupying the distal quarter of the antenna and inflated two to three times the thickness of the shaft. The relative lengths of the second and third segments of the palpus are highly variable within the tribe: the third segment is less than one-sixth the length of the second in Crebeta and between one-third and one-half in Hermias; in most genera the third palpal segment is one-fourth to one-fifth the length of the second. The length of the hairs of the second segment of the palpus is also variable: the hairs are shorter than the segment is wide in Hermias, but four times as long as the segment is wide in such genera as Magula and Lopinga; in most genera the hairs are about three times as long as the second segment is wide. The development of the male foreleg is also variable, although the leg itself is relatively unminiaturized. There is a great variation in the relative length of the forefemur and tibia, and the tarsus has one to three subsegments and is usually unspined, although there are many tarsal spines in such genera as Ethope and Paralethe. The female foreleg is likewise well developed and slightly clubbed at the end of the tarsus. The tarsus is always pentamerous with spines on at least the second, third and fourth subsegments (also on the first subsegment in such genera as Lethe and Ptychandra). The mid- and hindlegs are of about the same length. The midtibia is always less than twice the length of the first midtarsal subsegment. In the South African Aeropetes and Paralethe the midtibia is extremely spinose dorsad (Fig. 71), but the midtibia is smooth in other genera, such as Lethe and Satyrodes (Fig. 53). The tibial spurs are always present and well developed, but there is no spine at the dorsal, distal end of the midtibia. The relative lengths of the legs are shown in Fig. 48. The following cell is square-cut or rounded distad, never excavate as in other elymniine tribes, and about half as long as the forewing costa (Fig. 49, for example )—but this characteristic is by no means universa!. In Ptychandra the cell is only a third the length of the wing (Fig. 57), and in most members of the Pararge-series the cell is about three-fifths the length of the costa (Figs. 60, 64, 65). The forewing radial veins always arise in three branches from the cell, and veins Rs and M; are usually well separated at their origins, although they may arise approximate to one another. Forewing vein Mb» arises nearer, though not much nearer, M: than Ms; and Cu; arises midway LEE D. MILLER 45 = SEES ( S - Figs. 49-56. Elymniinae: Lethini. 49. Lethe europa (Fabricius), $ vena- tion. 50. L. europa, palpus. 51. L. europa, & foreleg. 52. L. europa, 2 fore- tarsus. 53. L. europa, midleg. 54. Rangbia scanda (Moore), 4 venation. 55. Rhaphicera satricus (Westwood and Hewitson), ¢ venation. 56. Tansima satyrina (Butler), ¢ venation. aS 96 between M; and Cuz in most genera, but Cu: arises nearer Cue in several genera (e.g., Lethe, Fig. 49, Ptychandra, Fig. 57, and Pararge, Fig. 60). Forewing vein Sc is often somewhat inflated, the cubital stem seldom inflated, although it is much inflated in Ptychandra, and 2A is never much swollen. The hindwing cell is produced at the origin of vein M; and is MEM. AMER. ENT. SOC., 24 46 THE SATYRIDAE two-fifths to three-fifths the length of the wing measured to the end of Ms. Hindwing vein Sc+Ry is of equal length or shorter than 3A. Veins Ms and Cu: are usually connate, but they are well separated in such aberrant genera as Mandarinia, Ptychandra and Rhaphicera. Vein Mp» usually arises nearer M;i than Ms, though not much nearer. In Aeropetes My» arises midway between Mi and Ms, slightly nearer Ms in the Pararge-series and much nearer Ms in Mandarinia. The pattern is highly variable, the main unifying feature being the diastoma of the hindwing ocelli in space Mi-Mz2 which was mentioned 9] Figs. 57-59. Elymniinae: Lethini. 57. Ptychandra lorquinii Felder and Felder, ¢ venation. 58. Enodia portlandia (Fabricius), 6 venation. 59. Saty- rodes eurydice (Linné), 6 venation. earlier. This characteristic is by no means universal, though, and is lacking in many genera, either because of the loss of many ocelli or extensive realignment of all the hindwing spots. There are several sites of androconial generation in the Lethini. Forewing patches are uncommon and restricted to the area between the origin of the cubitus and the inner margin. These forewing patches are rarely hair tufts (Ptychandra, Fig. 57) and are more frequently small patches of mealy scales under the origin of Cu:. The hindwing androconial patches are much more common and usually lie just outside the cell between Ms; and Cup, although Mandarinia has a Mycalesis-like hair tuft along | vein Rs (Fig. 73). Hindwing hair tufts are commoner than mealy patches, but both are present throughout the tribe. LEE D. MILLER 47 Genera Included in the Lethini Aeropetes-series Aeropetes Billberg, 1820: 79. Type-species: Papilio tulbaghia Linné, desig- nated by Hemming, 1943: 23. = Meneris Westwood, 1850, in Doubleday, Westwood and Hewitson, 1846-1852: 296. Type-species: Papilio tulbaghia Linné, by mono- typy. Paralethe van Son, 1955: 51. Type-species: Satyrus dendrophilus Trimen, by original designation. Lethe-series, sensu stricto Aphysoneura Karsch, 1894: 190. Type-species: Aphysoneura pigmentaria Karsch, by original designation. = Rhaphiceropsis Sharpe, 1894: 336. Type-species: Rhaphiceropsis pringlei Sharpe (= Aphysoneura pigmentaria Karsch), by original designation. Archondesa Moore, 1892 (1890-1893): 270. Type-species: Lethe lanaris But- ler, by original designation. Charma Doherty, 1886: 117. Type-species: Zophoessa baladeva Moore, by original designation. Proposed as a subgenus of Lethe Hiibner. = Putlia Moore, 1892 (1890-1893): 287. Type-species: Zophoessa baladeva Moore, by original designation. Proposed to replace Charma Doherty, wrongly believed to be preoccupied. Choranesa Moore, 1892 (1890-1893): 270. Type-species: Lethe trimacula Leech, by original designation. Debis Doubleday and Hewitson, 1849, in Doubleday, Westwood and Hewitson, 1846-1852: pl. 61. Type-species: Debis samio Doubleday and Hewit- son, by monotypy. Dionana Moore, 1892 (1890-1893): 271. Type-species: Lethe margaretae Elwes, by original designation. Enodia Hiibner, [1819] (1816-1826): 61. Type-species: Enodia andromacha Hubner (= Papilio portlandia Fabricius), designated by Scudder, 1872: 26. Hanipha Moore, 1880 (1880-1881): 18. Type-species: Lethe sihala Moore (= Lethe dynsate Hewitson), by original designation. This name is commonly misspelled “Hanifa’. Harima Moore, 1892 (1890-1893): 299. Type-species: Neope callipteris But- ler, by original designation. Hermias Fruhstorfer, 1912 (1912-1915): 324. Type-species: Satyrus verma Kollar, by monotypy. Proposed as a “subgenus or species-group” of Lethe Hubner. Kerrata Moore, 1892 (1890-1893): 285. Type-species: Lethe tristigmata E\|- wes, by original designation. Kirrodesa Moore, 1892 (1890-1893): 237. Type-species: Debis sicelis Hew- MEM. AMER. ENT. SOC., 24 48 THE SATYRIDAE Figs. 60-72. Elymniinae: Lethini. 60. Pararge aegeria (Linné), ¢ vena- tion. 61. P. aegeria, palpus. 62. P. aegeria, é foreleg. 63. P. aegeria, ° foretarsus. 64. Crebeta deidamia (Eversmann), é venation. 65. Lopinga dumetorum (Oberthtir), venation of ¢ forewing. 66. A physoneura pigmentaria Karsch, venation of ¢ forewing. 67. Aeropetes tulbaghia (Linné), ¢ venation. 68. A. tulbaghia, palpus. 69. A. tulbaghia, 6 foreleg. 70. A. tulbaghia, 2 foretarsus. 71. A. tulbaghia, midleg. 72. Paralethe dendrophilus (Trimen), 6 venation. itson, by original designation. : Lethe Hiibner, [1819] (1816-1826): 56. Type-species: Papilio europa Fabri- cius, by monotypy. = Tanaoptera Billberg, 1820: 79. Type-species: Papilio europa Fab- LEE D. MILLER 49 ricius, designated by Hemming, 1933: 199. Magula Fruhstorfer, 1912 (1912-1915): 313. Type-species: Zophoessa jala- urida deNicéville, designated by Hemming, 1935: 1. Preoccupied by Magula Scudder (Megerle MS.), 1882, but no replacement name has been proposed. Nemetis Moore, 1892 (1890-1893): 237. Type-species: Papilio minerva Fab- ricius, by original designation. Neope Butler, 1867d: 166. Type-species: Lasiommata (?) bhadra Moore, designated by Butler, 1868b: 112. = Enope Moore, 1857: 228. Type-species: Lasiommata (?) bhadra Moore, designated by Moore, 1892 (1890-1893): 299. Preoc- cupied by Enope Walker, 1854. = Blanaida Kirby, 1877: 699. Type-species: Lasiommata (?) bha- dra Moore, by original designation. Proposed to replace Enope Moore. Ninguta Moore, 1892 (1890-1893): 310. Type-species: Pronophila schrenkii Ménétriés, by original designation. = Aranda Fruhstorfer, 1909: 134. Type-species: Pronophila schrenkii Ménétriés, by original designation. Patala Moore, 1892 (1890-1893): 305. Type-species: Zophoessa yama Moore, by original designation. Placilla Moore, 1892 (1890-1893): 253. Type-species: Lethe christophi Leech, by original designation. Ptychandra Felder and Felder, 1861: 304. Type-species: Ptychandra lorquinii Felder and Felder, by monotypy. Rangbia Moore, 1892 (1890-1893): 232. Type-species: Debis scanda Moore, by original designation. Samanta Moore, 1880: 166. Type-species: Mycalesis malsara Moore, by original designation. Satyrodes Scudder, 1875b: 242. Type-species: Papilio eurydice Linné, by original designation. = Argus Scopoli, 1777: 432. Type-species: Papilio eurydice Linné, designated by Scudder, 1872: 27. Preoccupied by Argus Bohadsch, 1761. Sinchula Moore, 1892 (1890-1893): 275. Type-species: Debis sidonis Hew- itson, by original designation. Tansima Moore, 1881: 305. Type-species: Lethe satyrina Butler, by mono- typy. Zophoessa Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 1852: 362. Type-species: Zophoessa sura Westwood, by monotypy. Pararge-series Chonala Moore, 1893 (1890-1893): 14. Type-species: Debis (Tansima) masoni Elwes, by original designation. MEM. AMER. ENT. SOC., 24 50 THE SATYRIDAE Crebeta Moore, 1893 (1890-1893): 11. Type-species: Pararge deidamia Evers- mann, by original designation. Kirinia Moore, 1893 (1890-1893): 14. Type-species: Lasiommata epimen- ides Ménétriés, by original designation. Lasiommata Westwood, 1841: 65. Type-species: Papilio megera Linné, desig- nated by Scudder, 1875a: 202. = Amecera Butler, 1867d: 162. Type-species: Papilio megera Linné, designated by Butler, 1868b: 123. Lopinga Moore, 1893 (1890-1893): 11. Type-species: Pararge dumetorum Oberthiir, by original designation. Orinoma Gray, 1846: 14. Type-species: Satyrus (?) damaris Doubleday (nec. Gray), by monotypy. Pararge Hubner, [1819] (1816-1826): 59. Type-species: Papilio aegeria Linné, designated by Butler, 1868a: 195. This name has been frequently mis- spelled as ‘‘Pararga” or “‘Parage” in the literature. Rhaphicera Butler, 1867d: 164. Type-species: Lasiommata satricus Westwood and Hewitson, designated by Butler. 1868b: 158. Tatinga Moore, 1893 (1890-1893): 5. Type-species: Satyrus thibetanus Ober- thiir, by original designation. Mandarinia-series Mandarinia Leech, 1892: 9. Type-species: Mycalesis regalis Leech, by original designation. Neorina-series Ethope Moore, 1865: 770. Type-species: Mycalesis (2?) himachala Moore, by monotypy. = Theope Moore, 1857: 234. Type-species: Mycalesis (?) hima- chala Moore, by original designation. Preoccupied by Theope Doubleday and Hewitson, 1847. = Anadebis Butler, 1867c: 50. Type-species: Mycalesis (?) hima- chala Moore, by monotypy. Euploeamima Holland, 1887: 113. Type-species: Zethera diademoides Moore, by original designation. Hermianax Fruhstorfer, 1912 (1912-1915): 326. Type-species: Neorina lati- picta Fruhstorfer (= Neorina lowi Doubleday and Hewitson), by origi- nal designation. Proposed as a subgenus of Neorina Westwood. Neorina Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 369. Type-species: Neorina hilda Westwood, by original designation. Figs. 73-87. Elymniinae: Lethini (Figs. 73-80) and Zetherini (Figs. 81- 87). 73. Mandarinia regalis (Leech), ¢ venation. 74. M. regalis, palpus. 75. M. regalis, 2 foretarsus. 76. M. regalis, midleg. 77. Neorina hilda West- - wood, 6 venation. 78. N. hilda, 6 foreleg. 79. N. hilda, 2 foretarsus. 80. LEE D. MILLER Syl 85 86 84 N. hilda, midleg. 81. Zethera pimplea (Erichson), é venation. 82. Z. pim- plea, palpus. 83. Z. pimplea, é foreleg. 84. Z. pimplea, 2 foretarsus. 85. Z. pimplea, midleg. 86. Amechania incerta Hewitson, é venation. 87. Cal- large sagitta (Leech), é venation. MEM. AMER. ENT. SOC., 24 Nn Nw THE SATYRIDAE Tribe ZETHERINI, new tribe (Figures 81-88) The zetherines have been considered part of the Elymniini by such authors as Gaede (1931), and while these two tribes share such features as close forewing veins Mz and Cui, they are abundantly distinct. Perhaps the closest relatives of the zetherines are certain of the more primitive lethines such as Aeropetes, but the differences discussed below also serve to separate these two tribes. The pattern of the zetherines is highly aberrant: several species are white with brown interveinal markings and either no ocelli or poorly developed ones, whereas Zethera pimplea (Erichson) looks like the Nearctic 0.4xML, FL ML HL IxML, YA 8 _ j Zethera Amechania Callarge Fig. 88. Elymniinae: Zetherini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of the genera. In all instances the ML value is unity. The top bar for each genus represents the measurements obtained from males, the bottom bar those from females. Limenitis weidemeyeri (Edwards) (Nymphalidae: Limenitinae), a black species with a wide white central band crossing both wings. The absence of inflation of the forewing veins serves to distinguish this tribe from the Mycalesini, while the spiny midtibia and the well developed tibial spurs separate the Zetherini from the Elymniini. Most Lethini, with the exception of the Neorina-series, have hairy eyes, whereas all zetherines have naked eyes. The relative position of the forewing veins M3, Cu: and Cup serve to distinguish members of this tribe from the Neorina-series of the Lethini. All the members of the Zetherini are Indo-Australian. The zetherines are character- ized as follows: The eyes are naked. The antennae are just over two-fifths the LEE D. MILLER 53 length of the forewing costa. The antennal club is weakly developed and gently tapered, never more than twice the thickness of the shaft. The relative lengths of the second and third segments of the palpus are variable: in Callarge the third segment is about one-seventh as long as the second, whereas in Zethera it is one-third as long. The hairs of the second segment of the palpus are always shorter than the segment is wide. The male foreleg is well developed with the femur longer than the tibia and a mono- to trimerous tarsus which may or may not bear spines. The female foreleg is also well developed with a pentamerous, slightly clubbed tarsus bearing spines on the first four subsegments. The hindlegs are equal to or slightly longer than the midlegs (Fig. 88). The mid-tibia is long twice to two and a half times as long as the first midtarsal subsegment—and the midtibia is spiny dorsad, bearing moderately well developed tibial spurs, but no spine at the dorsal, distal end. The forewing cell is slightly excavate and two-fifths to half as long as the forewing costa. The forewing radial veins arise from the cell in three branches, and veins Rs and M; are approximate, but not connate, at their origins. Vein Mz arises much nearer M: than Ms, and Cu; arises much nearer Ms than Cuz. None of the forewing veins are much inflated basally. The hindwing cell is strongly produced at the origin of M:; and is one-third to half the length of the wing measured to the end of Ms. Hindwing vein Sc+R; is longer than 3A. Veins Mz and Cui are connate, or nearly so, and Mb arises nearer (or in some cases, much nearer) M; than Ms. The pattern of all the species is aberrant. Callarge is cream- colored with dark veins. Amechania is whitish with dark veins, some dark interveinal markings and a few ocelli. Zethera is blackish- brown with broad discal white bands on all wings. There are no defined androconial patches on any zetherine. Genera Included in the Zetherini Amechania Hewitson, 1861 (1856-1876): 87. Type-species: Amechania in- certa Hewitson, by monotypy. Callarge Leech, 1892: 57. Type-species: Zethera sagitta Leech, by original designation. MEM. AMER. ENT. SOC., 24 54 THE SATYRIDAE Zethera C. Felder, 1861: 26. Type-species: Cynthia pimplea Erichson, by monotypy. Tribe ELYMNIINI Herrich-Schaffer, 1864 (Figures 89-97) Eurytelidae Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 1852: 403 (in part). Elymniina Herrich-Schaffer, 1864: 124. The Elymniini are a homogeneous group and show some affinities within the subfamily. There is little to connect the elymniines with the Lethini directly, but the positions of certain of the veins show a definite relationship between these butterflies and the zetherines, which in turn do relate to the Lethini. The deeply excavate forewing cell of these insects is approached in the Mycalesini, and many of the latter groups have androconial patches placed very like those which characterize the Elymniini. 0.4xML, FL ML HL IxML Elymnias eee Elymniopsis Melynias Mimadelias Bruasa Fig. 89. Elymniinae: Elymniini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements ob- tained from males, the bottom bar those from females. This tribe may be distinguished from all others within the sub- family by the lack of tibial spurs in all species. Many of the genera also have extremely short cells on both wings, but there are a few exceptions. LEE D. MILLER 55 All elymniines are Paleotropical, most being found in the Indo- Australian region, but a couple of species range as far west as West Africa. The Elymniini are characterized in the following paragraphs. The eyes are naked. The antennae are about two-fifths as long are the forewing costa, although they may be as much as 0.46 times as long as the costa. The antennal club is very gentle and seldom more than twice the width of the shaft, except in the genus Melynias where it is over twice the thickness of the shaft. The third segment of the palpus is from one-fifth (Melynias) to one-third (Dyctis) the length of the second segment. The hairs of the second palpal seg- ment are about as long as the segment is wide. The male foreleg is well developed with the femur longer than the tibia and with a monomerous, unspined tarsus. The female fore- leg is also well developed with a pentamerous, unclubbed tarsus bearing a double set of spines on the third and fourth subsegments (also a double set on the second subsegment in Elymniopsis). The mid- and hindlegs are of about the same length (Fig. 89). The midtibia is generally less than twice the length of the first midtarsal subsegment and is smooth dorsad, without tibial spurs or a spine at the dorsal, distal end. The forewing cell is usually deeply excavate (Figs. 90-92) and is always less than half the length of the forewing costa. Frequently the cell is less than a third as long as the costa. The forewing radial veins arise in three branches from the cell, and Rs and M;, are well separated at their origins. Vein Me arises much nearer M; than Ms, and Cu: is almost connate with Ms, never near Cuz. Forewing vein Sc is usually inflated basad; the other stems are not. The hindwing cell is more or less square-cut because of the dis- placement marginad of M: and is from one-third to almost half as long as the wing measured to the end of Ms. Hindwing vein Sc+Ri is always much shorter than 3A. Veins Ms and Cu: are always connate, or nearly so, and Mp» arises much nearer Mi: than Ms. The pattern is quite variable and has been considered a mimetic one. Many of the species resemble various Danaidae, particularly of the genus Euploea. For a description of most of the species see Moore, 1893 (1890-1893). MEM. AMER. ENT. SOC., 24 56 THE SATYRIDAE 93 Figs. 90-97. Elymniinae: Elymniini. 90. Elymnias hypermnestra (Linné), é venation. 91. Bruasa penanga (Westwood and Hewitson), ¢ venation. 92. Dyctis agondas Boisduval, 6 venation. 93. E. hypermnestra, palpus. 94. E. hypermnestra, é foreleg. 95. E. hypermnestra, 2 foretarsus. 96. E. hy- permnestra, midleg. 97. Elymniopsis lise Hemming, palpus. Genera Included in the Elymniini Agrusia Moore, 1893 (1890-1893): 144. Type-species: Melanitis esaca West- wood and Hewitson, by original designation. Bruasa Moore, 1893 (1890-1893): 144. Type-species: Melanitis penanga Westwood and Hewitson, by original designation. Didonis Hiibner, [1819] (1816-1826): 17. Type-species: Papilio vitellia Cramer, designated by Scudder, 1875a: 156. LEE D. MILLER Nn —i Dyctis Boisduval, 1832: 138. Type-species: Dyctis agondas Boisduval, by monotypy. Elymnias Hiibner, 1818 (1818-1825): 12. Type-species: Elymnias jynx Hiib- ner (= Papilio hypermnestra Linné), designated by Hemming, 1943: 24. Scudder (1875a: 162) designated Papilio lais Fabricius as type, based on Elymnias Hiibner, [1819] (1816-1826): 37. This designation was invalidated when the appropriate sections of the “Zutrage” were found to be older than those of the “Verzeichniss”. Elymniopsis Fruhstorfer, 1907: 171, 173-174. Type-species: Papilio phegea Fabricius (= Elymniopsis lise Hemming), designated by Hemming, 1943: 24. Melynias Moore, 1893 (1890-1893): 144. Type-species: Papilio lais Cramer (= Papilio nesaea Linné), by original designation. Mimadelias Moore, 1893 (1890-1893): 144. Type-species: Elymnias vasu- deva Moore, by original designation. Tribe MYCALESINI, new tribe (Figures 98-119) The mycalesines show rather close relationships to both the Lethini and the Elymniini. Such mycalesine genera as Orsotriaena and Bletogona lack the characteristic inflation of the veins, but do have the excavate forewing cell and the distinctive androconial patches, whereas the lethine Mandarinia regalis (Leech) has the androconial patch but lacks the inflation of forewing vein 2A or the excavate forewing cell. The excavate forewing cell and the presence in some genera of a hair tuft on the upper surface of the hindwing at the anterior margin of the cell strongly suggests a relationship between the Mycalesini and the Elymniini. The characteristic subquadrate shape of the basal inflation of forewing vein 2A serves to distinguish the vast majority of all mycalesines. In the Hypocystini (Satyrinae) this inflation of 2A is suggested, but in that tribe the inflation is in the form of a rounded knot, rather than a subquadrate thickening, as in the present tribe. The Mycalesini are general throughout the Old World tropics and form a homogeneous group. There is some regional correlation of genera in the tribe, and for that reason they are arranged geo- graphically here. It is interesting to note that although hairy or naked eyes do not characterize the tribe, all mainland African Myca- lesini have naked eyes, all those from Madagascar have hairy eyes and both states are found in the Indo-Australian genera. This MEM. AMER. ENT. SOCc., 24 58 THE SATYRIDAE 0.4 XML, FL ML HL IXML, Indo- Malayan | genera Mycalesis Satoa Orsotriaena Bletogona | African | genera Bicyclus eieienatanentoneeaeaan fo Dichothyris Admiratio Heteropsis Masoura Fig. 98. Elymniinae: Mycalesini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hind legs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements ob- tained from males, the bottom bar those from females. situation will be discussed further in the section on “Evolution and Zoogeography.” The Mycalesini are characterized as follows: The eyes are naked or hairy: the Indo-Australian species show both conditions, those from mainland Africa all have naked eyes and those from Madagascar have hairy eyes. The antennae are from two-fifths to half the length of the forewing costa. The antennal club is moderately well developed, occupying the distal one-third to one- fifth of the antenna and thickened from less than twice (Orsotriaena) to more than three times (such genera as Dalapa and Martanda) the . thickness of the shaft. The third segment of the palpus is usually one-third to one-fifth the length of the second segment, but the third LEE D. MILLER 59 segment is more than half as long as the second in Dalapa. The hairs of the second palpal segment are generally from as long as the seg- ment is wide to twice the width of the segment. The male foreleg is well developed with the femur longer than the tibia and one or (rarely) two unspined or weakly spined tarsal subsegments. The female foreleg is moderately well developed with a pentamerous, lightly clubbed (less so in genera from Madagascar, Fig. 118) tarsus bearing spines on at least the fourth subsegment, often also on the second and third and occasionally on the first (such genera as Martanda). The mid- and hindlegs are not significantly unequal. The midtibia is less than twice as long as the first midtarsal subsegment. The midtibia is usually unspined dorsad, although it is weakly spined in Orsotriaena (Fig. 108), etc.; tibial spurs are always present and well developed and there is no spine at the dorsal distal end of the midtibia. The forewing cell is excavate and varies little from half the length of the forewing costa. The forewing radial veins arise in three branches from the cell, and veins Rs and Mi arise separate, but approximate, from the cell. Vein Mes arises much nearer M: than M3, and Cu: arises nearer Mz than Cus. The pattern of basal infla- tion of the forewing veins is characteristic, except in Orsotriaena (Fig. 106) and Bletogona: veins Sc and the cubital stem are inflated normally, but markedly, and 2A is inflated near its base as a sub- quadrate knot. The hindwing cell is produced at Ms and is one-third to half as long as the wing measured to the end of M3. Hindwing vein Sc+Ri is as long as, or shorter than, 3A. Veins Ms and Cui are connate or approximate at their origins, except in Culapa (Fig. 104) where they are widely separated, and Mb» arises much nearer M; than Ms. The pattern is highly variable. Many genera show the greatest development of the ocelli in forewing space Cu:-Cuz. Most of the species are dark brown on both surfaces, hence the common name “bushbrowns”, but some of the African species are iridescent blue or purple above and the Malgache species Masoura masoura (Hewit- son) is white with a yellow patch on the forewing, superficially more resembling a pierid than a satyrid. The distribution of androconia is characteristic: there is a polished area along the inner margin of the forewing on the under surface, and other areas of androconial concentration are in the area between Cue and 2A of the forewing MEM. AMER. ENT. SOC., 24 60 THE SATYRIDAE 100 101 1 “102 ES : 106 107 *108 Figs. 99-108. Elymniinae: Mycalesini. 99. Mycalesis francisca (Stoll), é venation. 100. M. francisca, palpus. 101. M. francisca, & foreleg. 102. M. francisca, 2 foretarsus. 103. M. francisca, midleg. 104. Culapa mnasicles (Hewitson), ¢ venation. 105. Mydosama fuscum (Felder and Felder), ¢ venation. 106. Orsotriaena medus (Fabricius), $ venation. 107. O. medus, 2 foretarsus. 108. O. medus, midleg. and along the hindwing cell on both margins. Hair tufts are more frequent than mealy patches in this tribe. Genera Included in the Mycalesini Indo-Australian genera Bletogona Felder and Felder, 1867 (1864-1867): 465. Type-species: Bleto- LEE D. MILLER 6i gona mycalesis Felder and Felder, by monotypy. Calysisme Moore, 1880 (1880-1881): 20. Type-species: Papilio drusia Cra- mer, designated by Moore, 1891 (1890-1893): 172. Celebina Fruhstorfer, 1898: 263. Type-species: Celebina inga Fruhstorfer, designated by Hemming, 1935: 1. Culapa Moore, 1878: 825. Type-species: Mycalesis mnasicles Hewitson, by monotypy. Dalapa Moore, 1880: 158. Type-species: Mycalesis sudra Felder, by original designation. Hamadryopsis Oberthtir, 1899: 17. Type-species: Hamadryopsis drusillodes Oberthiir, by monotypy. = Drusillopsis Fruhstorfer, 1908: 217. Type-species: Hamadryopsis drusillodes Oberthitir, by monotypy. Indalasa Moore, 1880: 166. Type-species: Mycalesis moorei Felder, by mono- typy. Jatana Moore, 1880: 164. Type-species: Mycalesis mynois Hewitson, by monotypy. Kabanda Moore, 1880: 168. Type-species: Mycalesis malsarida Butler, by original designation. Loesa Moore, 1880: 177. Type-species: Mycalesis oroatis Hewitson, by mono- typy. Lohora Moore, 1880: 175. Type-species: Mycalesis dexamenus Hewitson, by original designation. Martanda Moore, 1880: 169. Type-species: Mycalesis janardana Moore, by original designation. Monotrichtus Hampson, 1891: 179. Type-species: Mycalesis safitza Hewitson, by original designation. Originally proposed as a subgenus of Myca- lesis Hiibner. Mycalesis Hiibner, 1818 (1818-1825): 17. Type-species: Papilio francisca Stoll, designated by Hemming, 1937: 149. Butler, 1868a: 196, desig- nated Papilio evadne Cramer as type, based on Mycalesis Hubner, [1819] (1816-1826): 55. This designation was invalidated when the appropriate sections of the ‘“Zutrage” were found to be older than those of the “Verzeichniss”. = Gareris Moore, 1880: 156. Type-species: Mycalesis sanatana Moore (= Papilio francisca Stoll), by original designation. Mydosama Moore, 1880: 170. Type-species: Dasyomma fuscum Felder and Felder, by original designation. = Dasyomma Felder and Felder, 1860: 401. Type-species: Dasy- omma fuscum Felder and Felder, by monotypy. Preoccupied by Dasyomma Macquart, 1841. Myrtilus deNicéville, 1891: 341. Type-species: Mycalesis (Myrtilus) mystes deNicéville, by original designation. Proposed as a subgenus of Myca- lesis Htibner. MEM. AMER. ENT. SOC., 24 62 THE SATYRIDAE Sie 111 : : | 114 | | bs 115 118 Figs. 109-119. Elymniinae: Mycalesini. 109. Bicyclus hewitsonii (Dou- met), ¢ venation. 110. B. hewitsonii, palpus. 111. B. hewitsonii, 8 foreleg. 112. B. hewitsonii, 2 foretarsus. 113. Dichothyris sambulos (Hewitson), ¢ venation. 114. Heteropsis drepana Westwood, antennal club. 115. H. dre- pana, 6 venation. 116. H. drepana, palpus. 117. H. drepana, ¢ foreleg. 118. H. drepana, 2 foretarsus. 119. Admiratio paradoxa (Mabille), ¢ vena- tion. Nasapa Moore, 1880: 176. Type-species: Mycalesis aramis Hewitson, by monotypy. Nebdara Moore, 1880: 173. Type-species: Mycalesis tagala Felder, by origi- nal designation. Nissanga Moore, 1880: 169. Type-species: Mycalesis patnia Moore, by origi- nal designation. Orsotriaena Wallengren, 1858: 79. Type-species: Papilio medus Fabricius (= Papilio hesione Cramer), by monotypy. Pachama Moore, 1880: 165. Type-species: Mycalesis mestra Hewitson, by monotypy. LEE D. MILLER 63 Physcon deNicéville, 1898: 133. Type-species: Mycalesis pandoea Hopfler, by original designation. Proposed as a subgenus of Mycalesis Hiibner. Sadarga Moore, 1880: 157. Type-species: Mycalesis gotama Moore, by origi- nal designation. Samundra Moore, [1890-1892] (1890-1893): 162. Type-species: Mycalesis anaxioides Marshall and deNicéville, by monotypy. Satoa Moore, 1880: 157. Type-species: Mycalesis maianeas Hewitson, by monotypy. Sevanda Moore, 1880: 174. Type-species: Satyrus duponcheli Guérin, by original designation. Suralaya Moore, 1880: 159. Type-species: Mycalesis oreseis Hewitson, by monotypy. Telinga Moore, 1880: 167. Type-species: Satyrus adolphei Guérin, by mono- typy. Virapa Moore, 1880: 156. Type-species: Mycalesis anaxias Hewitson, by origi- nal designation. African genera Bicyclus Kirby, 1871la: 47. Type-species: Idiomorphus hewitsonii Doumet, designated by Kirby, 1871b: 363. = Idiomorphus Doumet, 1861: 174. Type-species: Idiomorphus hewitsonii Doumet, by monotypy. Preoccupied by Idiomorphus Chaudoir, 1846. Dichothyris Karsch, 1893: 203. Type-species: Mycalesis sambulos Hewitson, designated by Hemming, 1935: 1. Hallelesis Condamin, 1960: 1257. Type-species: Mycalesis asochis Hewitson, designated by Condamin, 1961: 783. Malgache genera Admiratio Hemming, 1964b: 137. Type-species: Smithia paradoxa Mabille, by original designation. = Smithia Mabille, 1880: 173. Type-species: Smithia paradoxa Mabille, by monotypy. Preoccupied by Smithia Milne Edwards and Haime, 1851. | Henotesia Butler, 1879: 228. Type-species: Henotesia wardii Butler, by origi- nal designation. Heteropsis Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 1852: 323. Type-species: Heteropsis drepana Westwood, by monotypy. Houlbertia Oberthiir, 1916: 199. Type-species: Erebia passandava Ward, des- ignated by Hemming, 1964a: 120. Masoura Hemming, 1964b: 138. Type-species: Melanitis masoura Hewitson, by original designation. = Gallienia Oberthir, 1916: 205. Type-species: Melanitis masoura Hewitson, designated by Hemming, 1964a: 120. MEM. AMER. ENT. SOC., 24 64 THE SATYRIDAE Subfamily ERITINAE, new subfamily It is obvious that the two genera in this subfamily are intermediate between the Elymniinae (and closest to the Lethini of that subfamily ) and the Satyrinae, but perhaps the present genera should not be com- bined into a single sub-family or tribe. Nevertheless, these genera seem closest to one another, and I should be reluctant to erect two monotypic taxa for them. The female foretarsus is pentamerous and unclubbed (Figs. 125, 131) and more primitive than that of the Satyrinae. The pattern is rather more primitive than that of most satyrines (Schwanitsch, 1924) and more closely approximates the pattern of the lethines. The male foreleg of the genus Coelites is much reduced over the situation in the lethines, however, and the forewing cell of Erites is much longer than that of any elymniine. Both genera are found on the Malay peninsula and in Indonesia. A single tribe, the Eritini, includes all species. Tribe ERITINI, new tribe (Figures 120-132) The diagnostic features of the subfamily Eritinae and of the only tribe, the Eritini, are as follows: The eyes are naked. The antennae are about two-fifths the length 0.4x ML, FL ML Erites Coelites Fig. 120. Eritinae: Eritini. Relative lengths of the femur + tibia + tar- sus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of the genera. In all instances the ML value is unity. The top bar for each genus represents the measurements obtained from males, the bottom bar those from females. of the forewing costa. The antennal club is indistinct (Coelites, Fig. 122) to gradual (EFrites, Fig. 128) and never more than twice | as thick as the shaft. The third segment of the palpus is about LEE D. MILLER 65 one-fifth as long as the second, and the development of the hairs of the second segment is variable: in Coelites they are about as long as the segment is wide, whereas in Erites the hairs are two and a half times the width of the segment. The male foreleg is variably developed: that of Erites is slightly miniaturized with subequal femur and tibia and a monomerous, unspined tarsus (Fig. 130); whereas that of Coelites is greatly reduced, the femur longer than the tibia and the tarsus represented only by a bump at the tip of the tibia (Fig. 124). The female foreleg is moderately well developed with a pentamerous, unclubbed tarsus bearing spines on the third and fourth subsegments (Erites, Fig. 131) or on the first four subsegments (Coelites, Fig. 125). The midlegs are approximately equal to, or slightly longer than, the hindlegs (Fig. 120). The midtibia is just over twice as long as the first midtarsal subsegment and is smooth dorsad with moderately well developed tibial spurs and no spines at the dorsal, distal end. The forewing cell is produced at the origin of M: and one-half (Coelites, Fig. 121) to three-fifths (Erites, Fig. 127) as long as the forewing costa. The forewing radial veins arise in three branches from the cell, and Rs and M: are approximate. Vein Mb arises slightly nearer (Erites) to much nearer (Coelites) Mi than Ms, and Cui arises somewhat nearer Mz; than Cus. Only forewing vein Sc is inflated basad. The hindwing cell is produced at the origin of Ms and is less than half (Coelites, Fig. 121) to three-fifths (Erites, Fig. 127) as long as the wing measured to the end of Ms. Hindwing vein Sc+ Ri is longer than 3A, and veins Mz and Cu: may be connate (Coelites ) to widely separated (Erites) at their origins. Vein Mz arises slightly nearer M; than Ms. Erites is an olive-brown insect above with the lines of the under surface faintly indicated and ocelli in forewing spaces Mi-M2, Mo-Ms and a large one in Cui-Cuz and hindwing spaces Rs-M; through Cui-Cue. On the under surface the pattern is the same but there are dark striations and transverse ochre lines in the medial areas of both wings. Coelites is uniform gray-brown above with a basal purplish sheen and a patch of oily hairs between hindwing veins 2A and 3A. The under surface is dark brown basad, gray-brown mar- MEM. AMER. ENT. SOC., 24 66 THE SATYRIDAE 128 129 130 | 131 132 Figs. 121-132. Eritinae: Eritini. 121. Coelites nothis Westwood and Hewitson, 6 venation. 122. C. nothis, antennal club. 123. C. nothis, palpus. 124. C. nothis, 6 foreleg. 125. C. nothis, 2 foretarsus. 126. C. nothis, mid- leg. 127. Erites madura (Horsefield), ¢ venation. 128. E. madura, antennal club. 129. E. madura, palpus. 130. E. madura, ¢ foreleg. 131. E. madura, 2 foretarsus. 132. E. madura, midleg. ginad with ocelli in hindwing spaces Rs-M; through Cui-Cur. Genera Included in the Eritini Coelites Westwood and Hewitson, 1850, in Doubleday, Westwood and Hewit- son, 1846-1852: pl. 66. Type-species: Coelites nothis Westwood and Hewitson, designated by Butler, 1868a: 195. Erites Westwood, 1850, in Doubleday, Westwood and Hewitson, 1846-1852: 392. Type-species: Hipparchia madura Horsefield, by monotypy. LEE D. MILLER 67 Subfamily RAGADIINAE Herrich-Schaffer, 1864 Ragadiina Herrich-Schaffer, 1864: 124. It is difficult to place the Ragadiinae in a satisfactory systematic position, but for reasons to be enumerated below, this subfamily appears to occupy a position intermediate between the Elymniinae and the Satyrinae. In common with the former, hindwing veins Mz; and Cu: arise from a common point on the cell, but the abortion of the forelegs of both sexes (Figs. 136, 137) is a characteristic shared with the Satyrinae and Coelites of the Eritinae. In other respects, however, the ragadiines stand apart from all of the other subfamilies. The configuration of the hindwing cell (Figs. 134, 139) is dis- tinctive. In males the cell is closed by narrow vestigial veins, whereas in the females of some species the cell is completely open between the origins of veins Me and Ms. All members of the Ragadiinae are native to the Indo-Australian region and constitute a single tribe, the Ragadiini. Rober (1892) split this subfamily, referring Acrophtalmia to the Ypthima group and Ragadia to the Mycalesis group, but the characteristic venation of the hindwing cannot be ascribed merely to the convergence of two different phyletic lines. Tribe RAGADIINI Herrich-Schaffer, 1864 (Figures 133-139) Ragadiina Herrich-Schaffer, 1864: 124. 0.2xML, FL ML HL XML Ragadia Acrophtalmia Fig. 133. Ragadiinae: Ragadiini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements ob- tained from males, the bottom bar those from females. MEM. AMER. ENT. SOC., 24 68 THE SATYRIDAE 135 136 138 139 137 Figs. 134-139. Ragadiinae: Ragadiini. 134. Ragadia crisia (Geyer), 4 venation. 135. R. crisia, palpus. 136. R. crisia, é foreleg, showing also the trochanter and the distal portion of the coxa. 137. R. crisia, 2 foretarsus. 138. R. crisia, midleg. 139. Acrophtalmia artemis Felder and Felder, ¢ vena- tion. The diagnostic features of the subfamily Ragadiinae and its only tribe, the Ragadiini, are as follows: The eyes are naked. The antennae are no more than half the length of the forewing costa. The antennal club is moderately well developed, occupies the distal one-fourth to two-fifths of the antenna and is thickened to two and a half to three times the width of the shaft. The third segment of the palpus is from less than a quarter (Ragadia) to slightly less than one-third (Acrophtalmia) the length of the second segment. The hairs of the second segment of the palpus are shorter than the segment is wide. The male foreleg is greatly reduced with but a bump representing the tarsus in some species (Ragadia, Fig. 136, for example); the tarsus is completely fused with the tibia and indistinguishable from it in Acrophtalmia. The femur and tibia are of about the same length. The female foreleg is also miniaturized and the tarsus clubbed with five subsegments bearing spines on the first four. The mid- and hindlegs are of about the same length. The midtibia is less than twice the length of the first midtarsal subsegment, is smooth dorsad and bears weakly developed tibial spurs, but no spine at the dorsal, . distal end. The relative lengths of the legs are shown in Fig. 133. The forewing cell is excavate near the origin of M» and about LEE D. MILLER 69 three-fifths the length of the forewing costa. The forewing radial veins arjse in two branches from the cell, and veins Rs and M: are nearly connate. Vein Mb» arises nearer Mi than M3, and Cuz arises much nearer M3; than Cuz. Forewing vein Sc is inflated basad, but the other veins are not. The hindwing cell is distinctive. It is open in some females and at most closed by small, aberrant veins (Figs. 134, 139). The cell is from less than half (Acrophtalmia) to more than three-fifths (Ragadia) as long as the wing measured to the end of Ms. The relative lengths of hindwing veins Sc+Ri and 3A vary: Sc+Ri is longer in Acrophtalmia, and 3A is longer in Ragadia. Veins Ms and Cu: are connate, whereas Mp» arises basad of M: and much closer to it than to Ms. The pattern is highly modified, either brown with a broad central white band or with alternating brown and off-white stripes. The ocelli are poorly developed, except in Ragadia which has a full com- plement on the under surface. An androconial hair tuft lies on the aberrant veins of the hindwing cell (Figs. 134, 139) and is charac- teristic of the tribe. Genera Included in the Ragadiini Acrophtalmia Felder and Felder, 1861: 305. Type-species: Acrophtalmia ar- temis Felder and Felder, by monotypy. Many authors, following Fel- der and Felder, 1867 (1864-1867): 486, have misspelled this name “Acrophthalmia’. Acropolis Hemming, 1934b: 77. Type-species: Acrophthalmia (sic.) thalia Leech, by original designation. = Pharia Fruhstorfer, 1912 (1912-1915): 295. Type-species: Ac- rophthalmia (sic.) thalia Leech, by monotypy. Preoccupied by Pharia Gray, 1840. Ragadia Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 376. Type-species: Euptychia crisia Geyer, by monotypy. Proposed as a subgenus of Neonympha Hubner. Subfamily SATYRINAE Boisduval, 1836 Satyrides Boisduval, 1836: 166; Satyridae Swainson, 1840: 86. = Hipparchiadae Kirby, 1837: 297. = Maniolinae Hampson, 1918: 385. The Satyrinae are the most evolved members of the family and MEM. AMER. ENT. SOC., 24 THE SATYRIDAE 70 podojaaop A|3u0.1}8 podojoaop wen eee “A 0} AJTYBOM eee e cere eeenee ALIBI AA ¢ Ajarel ‘sways Z| Ajared ‘suuays 7 VC A][euo1sed50 “Te WZ ‘Ng ‘9g " ‘skemye Ng ‘dg yjoowig “"* yyoous ‘uaH poonpol “13 ‘snoJOwesjo} peqqnys “IO snOIOWOUOW * ‘snoroweUdg * aSiereruear: Z< wag Cf T-oy Bsn00seceer C/ET/P 6 /EZ/T TurydurAuous09 tunyoAydng podojaaop podojaaop se eeeenees A[snorse A sence eeene [Jo UdIH ogo swio}s Z IO | “* Swd}s € 10 Z BIIUIS SNOLIVA UI SI9Y}O see eeeee Ajuo no ‘OS seen eeneee ‘sAVmye oS . 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AMER. ENT. SOC., 24 2 THE SATYRIDAE have become the dominant group throughout most of the world. Fairly close relationships with three other subfamilies, the Elymniinae, Eritinae and Ragadiinae, are indicated by analysis of structures, and there are three instances of gradation between the elymniines and the satyrines. There is probably only convergence between the Aero- petes-series of the Lethini and the Dirini of the present subfamily. Far more significant evolutionarily, however, are such genera as Lamprolenis in the primitive satyrine tribe Hypocystini. These but- terflies show certain similarities to some Mycalesini and Lethini (Mandarinia) of the Elymniinae, particularly as regards pattern and the placement of the androconial areas. This is probably the stock from which the Satyrinae evolved. The Lethini are connected to the Satyrinae through the sub-family Eritinae, as indicated in the discussion of the eritines. This evidence seems to place the Lethini as the closest present-day ancestral stock of the Satyrinae. The relationship with the ragadiines is obscure and more in the nature of an association because of the greatly reduced forelegs of both sexes. Any satyrid with fewer than five subsegments on the female fore- tarsus 1s a member of this subfamily. Even when there are five subsegments on the female foreleg of a member of the present sub- family, the tarsus is formed into a clublike structure totally unlike anything found in the Haeterinae, Brassolinae, Biinae, Elymniinae or Eritinae, but hinted at in the Ragadiinae. Comparative female fore- tarsi of the pentamerous type are shown in Figs. 5 (Haeterinae), 15 (Brassolinae), 24, 32 (Biinae), 52, 84, 95, 102 (Elymniinae), 131 (Eritinae), 137 (Ragadiinae) and 145, 174, 200 (Satyrinae). A greatly reduced male foreleg will serve to separate the Satyrinae from all other satyrid subfamilies except the Ragadiinae and Coelites of the Eritinae. None of the satyrines have the hindwing veins Ms and Cur connate, nor the open hindwing cell of the ragadiines. The hindwing cell, measured to the origin of vein Ms, is greater than half as long as the wing to the end of Ms, except in some Dirini. The antennal club is always moderately well and frequently highly developed. The Satyrinae are virtually cosmopolitan, with representatives in all parts of the world where butterflies are found, except some oceanic islands. Unlike the other subfamilies, the Satyrinae are highly developed in temperate regions. Ten tribes are recognized in the Satyrinae. The most primitive LEE D. MILLER US tribe, the Hypocystini, contains genera which span the gap between the other two groups. The Ypthimini-section, characterized more or less by their smaller size, frailer appearance and an ecological prefer- ence for grasslands, contains the tribes Ypthimini, Euptychiini, Coe- nonymphini, Maniolini and Erebiini. The Satyrini-section, larger and sturdier insects with woodland or scrubland preferences, is com- posed of the tribes Dirini, Pronophilini, Satyrini and Melanargiini. A key to the tribes of the Satyrinae is given below, and a diagnostic chart of these tribes is provided in Table 6. KEY TO THE TRIBES OF THE SATYRINAE 1. Hindwing cell half or less the length of the wing; forewing cell rounded distad (Figs. 248, 255, 256); South Africa ....................s0sesss2+0000- Dirini Hindwing cell greater than half the length of the wing; forewing cell SQUIANE=CULTORZER CAV ALC rel ccvesere: semen ees uSneanah Onc leatauvealss tea ce tnacacetesecscd 2 2. Midtibia with a long spine at the dorsal distal end (Fig. 301); Holarc- LL Cee poet ee ee ae ene eer cnr a une Sune enee Minace eens Aabestaadeene onesere most Satyrini Mirdtibiaawithoutssuchwayspim eicerc.cseetee eset eeene crore see soreoeers sens teers 3 3. Hindwing cell prolonged by a distad displacement of crossvein m:2-m: alonm Wenn IM (eis, BOil, WO SxeiTNON)) coconoouesonocacovcdboopssbocnapossneqb0D¢aGoGee 4 Hindwing cell more or less square cut with crossvein mms in line with nena (ene, SOL), Ore SEIS) concsssosnccosonoooenoscecodcognéadoacoonagdadacigsaaneccddo0ou 8 4. Female foretarsus with fewer than five subsegments ................::::eeereeees 5 FERIA TOREIESUS: [OEMIAITIEITOUS scccconscoscososscoscobeoocada93te09000900000000000000000000000 6 5. Third segment of palpus less than one-fourth the length of the second (ETI, SKOSYS I SIOIENKCINE Gensseecosoncbotenchodosacabedarooseoe5a Satyrini (Oeneis-series ) Third segment of palpus longer than one-fourth the length of the second Ge 82) PAINeo tropical east e irae ete cote ccaseeeee oot some Pronophilini ©; IMIMCKOVEA GPO INISOUROOKES. cooccocbonecoocaosadopcocoqseoscscbuceecd600: most Pronophilini Midi rae smoother cscs ses mee. tree sec sete ens cesses ceeinaes gy acne ceneaiane coneeueeaave si caseneiee 7 Ys INI@OWROVNCAll GHSSE]S -ccccocoosesseconcanssssacosoadabascdecagadcooncecéanga0cs a few Pronophilini ANUCUANIEIN TEGO, GSSCIES shovssosoccocoonocnsosqcnosvespesdeoosbNeccodescoacabaeee Hypocystini 8. White (or cream-colored) species, marbled with black#giPal Cancti chars. eter cee ee eee ek hae Melanargiini Generally brown or orange species, if white (YOLE” Tay FO eC A oe ea ceacinateeecconedoounddnarceaoteecu ec oeteneacdinal SosoecSn une see CasusaEcR eee tes 9 9. Female foretarsus with fewer than five subsegments ................::eeee 10 EMMIS WOTABNUS PXSTUETNETROWS - jcoccosconsco.ccanaceasasdegssenc0qaacensacoCUAdeceeceoHOAFE i, 10. Forewing veins only slightly inflated (Figs. DS AOA (i) MEO LAR Chi Cece ace sere oe cate rete eo cai dicoehcttuetene Erebiini Forewing veins inflated (Fig. 224, for example) ................::::::eeeeeeeee 11 MEM. AMER. ENT. SOC., 24 74 THE SATYRIDAE 11. Midtibia twice or more the length of the first midtarsal subsegment; Holarctic ............. cc eeeeeessereeeeees Coenonymphini Midtibia less than twice as long as the first midtarsal subsegment; Holarctic ................::ssesscsscececececeeeeeeeeees Maniolini 12. Forewing with a double-pupilled ocellus in space Mia=Mcvor IMb=Mis-s@ldi Worl dieteeeessevenscoseesesercereree see eee 13 Forewing with no ocellus in space M:-Ms, or if present only single-pupilled; New World ..............:ccccccceeeseeeees Euptychiini Wis Feed Gis eS 01.216 bree tonsaneGoacboacecoacescsorctioacsonsccco oncdesacoacaededccuaccussptccone Ypthimini | BA (eRSIN) NEE i (eomaeciee osac ecb boa eapdee eas oodeacdt oanodsecbaobooedagsokeacddebene genus Palaeonympha Tribe HypocysTINI, new tribe (Figures 140-169) The Hypocystini are here considered to be the most primitive members of the Satyrinae, primarily because of the configurations of the male and female forelegs. Although these butterflies are not closely allied to any other tribe within the Satyrinae, they seem to connect the Ypthimini- and Satyrini-sections. The Xenica-series in the present tribe connects with the Satyrini-section by pattern and structural similarities, whereas the Hypocysta-series exhibits an even closer approach to the Ypthimini-section. The New Guinean hypo- cystine genus Lamprolenis shows many characteristics in common with the elymniine tribes Mycalesini and Lethini. This genus has the typical mycalesine androconial patch along the anterior portion of the hindwing, and the general facies are reminiscent of the mycalesines or Of Mandarinia. This similarity may offer an explanation of the evolution of the Satyrinae and will be discussed in greater detail later. Parallel evoijution is shown very well by the New Zealand mem- bers of this tribe, with the exception of Dodonidia. The other members, living in montane situations, have evolved into butterflies very like the Holarctic genus Erebia—indeed, several were placed in that genus by early authors. Because the New Zealand “coppers” are placed in the Holarctic genus Lycaena Fabricius (H. K. Clench, per- sonal communication), the possibility that Argyrophenga and espe- cially Erebiola and Percnodaimon are indeed referable to Erebia had to be examined, but there is no morphologic evidence to support their inclusion in the Holarctic genus. The New Zealand “Erebia’ . are highly modified Hypocystini, most nearly related to the Australian members of the tribe. LEE D. MILLER iD) 0.4x ML FL ML HL lx ML Hypocysta|series, S. Ss. | Hypocysta Platypthima Zipaetis Hyalodia 3 | XenicajSeries WH) Xenica Yl Heteronympha 7 Dodonidia Argynnina Argyrophenga Fig. 140. Satyrinae: Hypocystini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip- pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements obtained from males, the bottom bar those from females. Members of the New Guinean hypocystine fauna (Platypthima, Erycinidia, Pieridopsis and Lamprolenis) have evolved into bewil- dering butterflies. In fact, in wing shape and general appearance Erycinidia is a classic riodinid, and Pieridopsis is a fine pierid “mimic”. The less miniaturized forelegs in both sexes serves to distinguish the hypocystines from all other Satyrinae. The forewing cell is usually deeply excavate in this tribe, a characteristic shared with such Ypthimini-section tribes as the Ypthimini and Coenonymphini, but the hindwing crossvein mz-m; is produced marginad along Mb, as in much of the Satyrini-section. Forewing vein 2A is usually inflated basad, recalling the Mycalesini, but the inflated part is not sub- MEM. AMER. ENT. SOC., 24 76 THE SATYRIDAE quadrate in the hypocystines. A few of the Xenica-series have little or no inflation of forewing vein 2A. All members of the Hypocystini are found in the Australian region, a few being found as far west as Indonesia. Most of the species are found in New Zealand, Australia, New Guinea and the associated oceanic islands. Two series of genera are recognized. 1 I 142 { 144 ke h 146 = § Figs. 141-149. Satyrinae: Hypocystini. 141. Hypocysta euphemia West- wood and Hewitson, ¢ venation. 142. H. euphemia, antennal club. 143. H. euphemia, palpus. 144. H. euphemia, 6 foreleg. 145. H. euphemia, 2 fore- tarsus. 146. H. euphemia, midleg. 147. Erycinidia gracilis Rothschild and Jordan, é venation. 148. Lamprolenis nitida Godman and Salvin, é vena- tion. 149. Pieridopsis virgo Rothschild and Jordan, ¢ venation. The Hypocysta-series, sensu stricto, most of which are exclusively tropical insects, are characterized by the generally absent tibial spurs, LEE D. MILLER HY the longer third segment of the palpus and hindwing vein Mb arising nearer M3 than Mi. The Xenica-series is restricted to the temperate part of the region and characterized by the development of the tibial spurs, the shorter third segment of the palpus and (in many genera) hindwing vein Me, which arises midway between M: and Ms. The Hypocystini are characterized as follows: The eyes are generally naked, although they are hairy in such diverse genera as Argynnina, Heteronympha, Nesoxenica and Pla- typthima. The antennae are generally two-fifths to half the length of the forewing costa, but the antennae of Argyronympha are three- fifths the length of the costa. The antennal club is well developed in most genera, occupying the distal one-fifth to one-third of the antenna and expanded to three to four times the width of the shaft (just over twice the width of the shaft in Lamprolenis). The club is eccen- trically developed in some New Zealand genera (Figs. 163, 166, 169). The third segment of the palpus is variously one-half to one-third the length of the second in the Hypocysta-series (Fig. 143) and one-fourth to one-fifth the length of the second in the Xenica- series (Fig. 157). The hairs of the second segment of the palpus vary from less than the width of the segment to more than twice as long as the segment is wide in the Hypocysta-series to over four times the width of the segment in some of the New Zealand genera. 195 \ t \ \Y 152 153 151 150 154 Figs. 150-155. Satyrinae: Hypocystini, all Zipaetis saitis Hewitson. 150. 6 venation. 151. antennal club. 152. palpus. 153. 6 foreleg. 154. 9 foretarsus. 155. midleg. MEM. AMER. ENT. SOC., 24 78 THE SATYRIDAE The male foreleg is moderately well developed (Fig. 140); the tibia is usually longer than the femur, and the tarsus is usually mono- merous, but may have as many as four subsegments. The female foreleg is also well developed (Fig. 140) with a pentamerous, only slightly clubbed tarsus bearing spines on the first four subsegments (Fig. 145, for example )—there are more than one pair of spines on each of the first four subsegments in Dodonidia (Fig. 164). The midlegs are as long as, or longer than, the hindlegs (Fig. 140). The midtibia is from less than twice to more than twice the length of the first midtarsal subsegment, and the tibia is smooth dorsad. Tibial spurs are generally well developed in the Xenica-series (Fig. 160), including the New Zealand genera; but the spurs are generally absent in the Hypocysta-series (Fig. 146), but they are weakly developed in Pieridopsis and moderately developed in Zipaetis (Fig. 155). No spine is developed on the dorsal, distal end of the midtibia. The forewing cell is excavate, often deeply so, and is from more than half to more than three-fifths the length of the forewing costa. In the Hypocysta-series, Argyrophenga and Dodonidia the forewing radial veins arise in two branches from the cell; in the other genera the radials arise in three branches. Forewing veins Rs and Mi: are connate in Argyrophenga, Erebiola and Percnodaimon, and separate, though often approximate, in the other genera. Vein Mb arises nearer, often much nearer, Mi than Ms, and Cu; arises a little nearer Ms; than Cus. Vein Sc is characteristically inflated in all genera but Dodonidia (Fig. 162), and the cubital stem and 2A are inflated in such diverse genera as Argyrophenga (Fig. 167), Xenica (Fig. 156), Geitoneura (Fig. 161) and most of the Hypocysta-series (Fig. 141, for example). The hindwing cell is usually produced at m2-ms, but it is more blunted in Dodonidia (Fig. 162) and Zipaetis (Fig. 150), and the cell is usually half to three-fifths the length of the wing measured to the origin and end of Ms, respectively. The relative lengths of the hindwing veins Sc +R: and 3A are variable. Veins Ms and Cu: arise separately, and Ms commonly arises midway between Mi and Ms, but nearer the latter in the New Zealand “Erebia” and the Hypocysta- Figs. 156-169. Satyrinae: Hypocystini. 156. Xenica achanta (Donovan), 4 venation. 157. X. achanta, palpus. 158. X. achanta, ¢ foreleg. 159. X. achanta, @ foretarsus. 160. X. achanta, midleg. 161. Geitoneura klugii LEE D. MILLER 719 (Guérin), ¢ venation. 162. Dodonidia helmsii Butler, 6 venation. 163. D. helmsii, antennal club. 164. D. helmsii, 2 foretarsus. 165. Percnodaimon pluto (Fereday), & venation. 166. P. pluto, antennal club. 167. Argy- rophenga antipodum Doubleday, 4 venation. 168. Erebiola butleri Fereday, 6 venation. 169. E. butleri, antennal club. MEM. AMER. ENT. SOC., 24 80 THE SATYRIDAE series. The pattern is variable and highly modified. Many of the species are illustrated by Waterhouse and Lyell (1914) and by Fruhstorfer (1912[1912-1915]). There is a remarkable resemblance between Percnodaimon, Erebiola and Argyrophenga and the Holarctic Erebia. Genera Included in the Hypocystini Hypocysta-series, sensu stricto Argyronympha Mathew, 1886: 346. Type-species: Argyronympha_ pulchra Mathew, designated by Hemming, 1943: 23. Erycinidia Rothschild and Jordan, 1905: 457. Type-species: Erycinidia gra- cilis Rothschild and Jordan, by monotypy. Harsiesis Fruhstorfer, 1912(1912-1915): 299. Type-species: Hypocysta hy- geia Hewitson, by monotypy. Hyalodia Jordan, 1924: 285. Type-species: Hypocysta tenuisquamosa Joicey and Talbot, by monotypy. Hypocysta Westwood and Hewitson, 1850, in Doubleday, Westwood and Hewitson, 1846-1852: pl. 67. Type-species: Hypocysta euphemia Westwood and Hewitson, by monotypy. Lamprolenis Godman and Salvin, 1880: 610. Type-species: Lamprolenis nitida Godman and Salvin, by monotypy. Pieridopsis Rothschild and Jordan, 1905: 457. Type-species: Pieridopsis virgo Rothschild and Jordan, by original designation. Platypthima Rothschild and Jordan, 1905: 458. Type-species: Platypthima ornata Rothschild and Jordan, by original designation. Zipaetis Hewitson, 1863(1856-1876): [100]. Type-species: Zipaetis saitis Hewitson, designated by Butler, 1868a: 194. This genus is exception- ally aberrant and is only provisionally placed here. X enica-series Argynnina Butler, 1867d: 165. Type-species: Lasiommata hobartia Westwood and Hewitson, designated by Butler, 1868a: 196. Argyrophenga Doubleday, 1845: 307. Type-species: Argyrophenga antipodum Doubleday, by monotypy. Dodonidia Butler, 1884: 172. Type-species: Dodonidia helmsii Butler, by original designation. Erebiola Fereday, 1879: 128. Type-species: Erebiola butleri Fereday, by original designation. Geitoneura Butler, 1867d: 164. Type-species: Satyrus klugii Guérin, desig- nated by Butler, 1868b: 166. LEE D. MILLER 81 Heteronympha Wallengren, 1858: 78. Type-species: Papilio merope Fabricius, designated by Butler, 1868a: 195. =Hipparchioides Butler, 1867c: 125. Type-species: Papilio merope Fabricius, designated by Butler, 1868b: 99. Nesoxenica Waterhouse and Lyell, 1914: 35. Type-species: Lasiommata leprea Hewitson, by original designation. = Xeniconympha Noricky, 1923: 60. Type-species: Lasiommata leprea Hewitson, by original designation. Oreixenica Waterhouse and Lyell, 1914: 41. Type-species: Lasiommata (?) lathionella Westwood, by original designation. Paratisiphone Watkins, 1928: 615. Type-species: Lasiommata lyrnessa Hewit- son, by original designation. Percnodaimon Butler, 1876: 152. Type-species: Erebia pluto Fereday, by original designation. Tisiphone Hibner, [1819](1816-1826): 60. Type species: Papilio abeona Donovan, designated by Butler, 1868b: 71. Xenica Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 387. Type-species: Papilio achanta Donovan, designated by Scudder, 1875a: 289. Tribe YPTHIMINI, new tribe (Figures 170-188) This tribe lies at the base of the Ypthimini-section of the Satyrinae and seems in some respects to connect with the Hypocysta-series of the last tribe. In its turn the Ypthimini appears to have given rise to the Holarctic Erebiini and Coenonymphini and the Neotropical Euptychiini; the Maniolini were evolved later from a coenonymphine stock. The Ypthimini are generally characterized by a pattern feature: the forewing ocellus in space Mi-Ms3 has a double pupil of white or bluish-white scales. Forewing vein 2A is not inflated in the ypthi- mines, whereas in the Coenonymphini and many Euptychiini this vein is greatly distended. While the hindwing crossvein m2-ms 1s produced at M2 in the Hypocysta-series, the crossveins are aligned in this tribe and in other members of the Ypthimini-section, resulting in a straight cell. The female foreleg is pentamerous in the present tribe, a situation which is never found in the coenonymphines and very rare in either the Maniolini or the Erebiini. MEM. AMER. ENT. SoOc., 24 82 THE SATYRIDAE 0.2X ML, FL ML Al 1X ML, Ypthima Thympia Xois = oo Dallacha i — ee Kolasa Melampias Pseudonympha | AE mae : Strabena | Callerebia|series | Soe oe — Callerebia A ee : Lip svi fo ce Paralasa ? i — if Es Loxerebia Fig. 170. Satyrinae: Ypthimini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements obtained from males, the bottom bar those from females. All ypthimines are found in the Old World, and most genera are exclusively tropical. Most species fly in open country; few are found in scrub country, much less in the deep forest. Three major series are recognized in the Ypthimini, as follows: The Ypthima-series, sensu stricto, characterized generally by their small size, brown coloration and diffuse androconia. This series is found throughout the Paleotropics. LEE D. MILLER 83 The Callerebia-series, characterized by large size, brown color- ation and androconia arranged in rows on the discal portion of the forewing (Fig. 184). These species are generally distributed in the temperate mountains of the Middle and Far East. The Melampias-series, characterized generally by reddish shades on the upper surface, more elongate wings, small size and either diffuse androconia or none. These butterflies are found in South Africa and Madagascar. The Ypthimini are characterized as follows: The eyes are naked. The antennae are short, often less than two-fifths the length of the forewing costa (only one-fourth as long in Mashuna) and never as long as half the length of the costa (nearly half in Paralasa and Dallacha). The antennal club is variously developed: in such genera as Physcaeneura, Periplysia and Dallacha the club is less than twice as thick as the shaft, whereas in genera like Mashuna the club is almost four times as thick as the shaft. The third segment of the palpus is usually from one-fourth to slightly over one-third the length of the second, but the third segment is more than half as long as the second segment in Physcaeneura. The hairs of the second segment of the palpus are very long, usually three to five times as long as the segment is wide. The male foreleg is extremely reduced (Fig. 170), with the tarsus represented in most genera by a small knob on the end of the tibia (Fig. 173, for example). In such genera as Mashuna (Fig. 177), Strabena and Pandima, the tarsus is completely fused to the tibia, but in the Callerebia-series (Fig. 185) the tarsus is moderately well represented. The forefemur is as long as, or longer than, the fore- tibia. The female foreleg is also miniaturized, but it retains five tarsal subsegments, but the tarsus itself is significantly clubbed (Fig. 174, for example). There are spines on the second, third and fourth subsegments. The midleg is subequal to the hindleg in the Callerebia- and Melampias-series, but significantly shorter in the Ypthima-series (Fig. 170). The midtibia is less than twice the length of the first midtarsal subsegment, is smooth dorsad, without a spine at the dorsal, distal end and with well to moderately well developed tibial spurs, except in Physcaenura (Fig. 179). The forewing cell is usually produced at the origin of Ms and not excavate (except in Periplysia, Fig. 180); the cell is about half MEM. AMER. ENT. SOC., 24 84 THE SATYRIDAE 173 175 178 Figs. 171-180. Satyrinae: Ypthimini. 171. Ypthima philomela (Linné), é venation. 172. Y. philomela, palpus. 173. Y. philomela, ¢ foreleg, in- cluding trochanter and distal portion of coxa. 174. Y. philomela, 2 foretarsus. 175. Y. philomela, midleg. 176. Mashuna mashuna (Trimen), ¢ venation. 177. M. mashuna, 8 foreleg, including trochanter and distal portion of coxa. 178. Xois sesara Hewitson, venation of ¢ hindwing. 179. Physcaenura panda (Boisduval), midleg. 180. P. panda, ¢ venation. as long as the forewing costa. The forewing radial veins arise in one or two branches from the cell, and veins Rs and M: are well separated at their origins. Vein Me usually arises midway between M: and Ms, but nearer Mi, in the Melampias-series; Cu: arises nearer, or much nearer, M3 than Cus. Forewing vein Sc is always much thickened at its base, the cubital stem is generally inflated and 2A never is. LEE D. MILLER 85 The hindwing cell is straight, longest at the origin of Ms and is about three-fifths the length of the wing measured to the end of Ms. Hindwing vein Sc+Ri, is as long as, and more commonly longer than, 3A. Veins Ms and Cu: arise separately, but often approximate, and Mp arises somewhat nearer M; than M3, or midway between the two veins. As noted in the general description, the pattern is rather constant. For illustrations of the various species see Moore (1893[1890-1893]) and van Son (1955). Genera Included in the Y pthimini Y pthima-series, sensu stricto Dallacha Moore, 1893(1890-1893): 94. Type-species: Yphthima (sic.) hya- griva Moore, by original designation. Hemadera Moore, 1893(1890-1893): 107. Type-species: Yphthima (sic.) narasingha Moore, by original designation. Kolasa Moore, 1893(1890-1893): 82. Type-species: Satyrus chenui Guerin, by original designation. Lohana Moore, 1893(1890-1893): 92. Type-species: Yphthima (sic.) inica Hewitson, by original designation. Mashuna van Son, (July) 1955: 159. Type-species: Ypthima mashuna Tri- men, by original designation. =Ypthimorpha Overlaet, (Dec.) 1955: 23. Type-species: Ypthima mashuna Trimen, by original designation. Nadiria Moore, 1893(1890-1893): 85. Type-species: Ypthima bolanica Mar- shall, by original designation. Pandima Moore, 1893(1890-1893): 86. Type-species: Satyrus nareda Kollar, by original designation. Thympia Moore, 1893(1890-1893): 58. Type-species: Papilio ba!dus Fabri- cius, by original designation. Xois Hewitson, 1865: 282. Type-species: Xois sesara Hewitson, by monotypy. Ypthima Hiibner, 1818(1818-1825): 17. Type-species: Papilio philomela Linné, by monotypy. This name is frequently misspelled in the litera- ture as “Yphthima’. Hemming (1964c: 157-158) makes a strong case for “philomela” of Hiibner being a misidentification of Ypthima hueb- neri Kirby. Should the International Commission on Zoological No- menclature rule that this is the case, Y. huebneri will become the type- species. Y pthimomorpha van Son, 1955: 158. Type-species: Ypthima itonia Hewit- son, by original designation. MEM. AMER. ENT. SOC., 24 86 THE SATYRIDAE 183 184 ¢ 185 186 Figs. 181-188. Satyrinae: Ypthimini. 181. Melampias hyperbius (Lin- né), 6 venation. 182. Cassionympha cassius (Godart), é venation. 183. C. cassius, ° foretarsus. 184. Callerebia scanda (Kollar), ¢ venation. 185. C. scanda, 6 foreleg. 186. C. scanda, 2 foretarsus. 187. C. scanda, mid- leg. 188. Boeberia parmenio (Boeber), ¢ venation. Melampias-series Callyphthima Butler, 1880: 335. Type-species: Pseudonympha wardii Butler, by original designation. Cassionympha van Son, 1955: 96. Type-species: Satyrus cassius Godart, by original designation. Coenyropsis van Son, 1958: 6. Type-species: Satyrus natalii Boisduval, by original designation. LEE D. MILLER 87 Neita van Son, 1955: 101. Type-species: Pseudonympha neita Wallengren, by original designation. Melampias Hiibner, [1819](1816-1826): 63. Type-species: Papilio hyperbius Linné, designated by Scudder, 1875a: 214. Neocoenyra Butler, 1885: 758. Type-species: Neocoenyra duplex Butler, by monotypy. Periplysia Gerstacker, 1871: 358. Type-species: Periplysia leda Gerstacker, by monotypy. Pseudonympha Wallengren, 1857: 31. Type-species: Papilio hippia Cramer, designated by Butler, 1868a: 194. Physcaeneura Wallengren, 1857: 32. Type-species: Satyrus panda Boisduval, by monotypy. Strabena Mabille, 1887: 10, 22. Type-species: Satyrus tamatavae Boisduval, by original designation. Stygionympha van Son, 1955: 137. Type-species: Pseudonympha_ vigilans Trimen, by original designation. Callerebia-series Argestina Riley, 1922: 469. Type-species: Callerebia waltoni Elwes, by origi- nal designation. Boeberia Prout, 1901: 233. Type-species: Papilio parmenio Boeber, by origi- nal designation. =Erebomorpha Elwes, 1899: 351. Type-species: Papilio parmenio Boeber, by original designation. Preoccupied by Erebomorpha Walker, 1860. Callerebia Butler, 1867e: 217. Type-species: Erebia scanda Kollar, by origi- nal designation. Loxerebia Watkins, 1925: 237. Type-species: Callerebia pratorum Oberthir, by original designation. Paralasa Moore, 1893(1890-1893): 103. Type-species: Erebia kalinda Moore, by original designation. Genus PALAEONYMPHA Butler, 1871 (Figures 189-195) Palaeonympha Butler, 1871: 401. Type-species: Palaeonympha opalina But- ler, by original designation. The original description of Palaeonympha pointed to its similarity to the North American Neonympha (Euptychiini). If Palaeonympha is a member of the Euptychiini it would be the first known Old World representative of that tribe. It is tempting to place the present genus there, since it is superficially very similar to the American Megisto cymela (Cramer), except for the androconial distribution in opalina. MEM. AMER. ENT. SOC., 24 88 THE SATYRIDAE Palaeonympha 193 195 ig} 192 194 Figs. 189-195. Satyrinae: tribe uncertain, Palaeonympha opalina Butler. 189. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diago- nal lines), midlegs (ML, light stipp'ing) and hind'egs (HL, heavy stippling). The ML value is unity. The top bar represents the measurements obtained from the male, the bottom bar those from the female. 190. ¢ venation. 191. antennal club. 192. palpus. 193. ¢ foreleg. 194. 2 foretarsus. 195. midleg. There are certain similarities between the Euptychiini and the Ypthi- mini—indeed, they probably arose from the same basic stock—and it is also tempting to place Palaeonympha in the latter tribe on purely geographical considerations, but if the present genus is one of the ypthimines, it is the only one with hairy eyes. Certainly Palaeo- nympha is either an ypthimine or a euptychiine, or it may represent an intermediate tribe, but I cannot place the genus in either tribe with certainty, and I hesitate to erect a monotypic tribe at this time. Palaeonympha is characterized as follows: The eyes are slightly hairy. The antennae are just less than half as long as the forewing costa. The antennal club is moderately well developed, occupying the distal quarter of the antenna and two and a half to three times as thick as the shaft. The third segment of the palpus is long, about two-fifths the length of the second. The. hairs of the second segment of the palpus are three and a half times as long as the segment is wide. LEE D. MILLER 89 The male foreleg is greatly reduced (Figs. 189, 193): the femur is longer than the tibia, and the tarsus is represented only by a bump at the end of the tibia. The female foreleg is also miniaturized (Figs. 189, 194) with a pentamerous tarsus, the fifth subsegment of which is greatly reduced, bearing spines on the second and third subseg- ments. The hindlegs are slightly longer than the midlegs (Fig. 189). The midtibia is less than twice the length of the first midtarsal sub- segment, smooth dorsad, with no spine at the dorsal, distal end and with well developed tibial spurs. The forewing cell is very slightly excavate between M: and M2 and about half as long as the forewing costa. The forewing radial veins arise from the cell in a single branch, and Rs and Mi; are separate at their origins. Vein Mb» arises slightly nearer M; than Ms, and Cu: arises nearer M; than Cuy. Forewing vein Sc and the cubital stem are both much inflated basad, but 2A is not. The hindwing cell is somewhat produced at the origin of Ms and is about three-fifths the length of the wing measured to the end of Ms;. Hindwing vein Sct+R: is longer than 3A, and Ms; and Cu: are well separated at their origins. Vein M2 arises nearer M; than Ms. The single species is brown above with a double-pupilled ocellus in M:i-Me of the forewing and single-pupilled ones in Rs-M;i and M:-Cui of the hindwing. There is an androconial patch of mealy scales along the boundary of the cell and extending out the veins from M, to 2A (Fig. 190). The under surface is light brown with discal and extradiscal transverse reddish-brown bands and well de- veloped ocelli as on the upper surface, plus very small ones in forewing spaces Ms-Cu; through Cus-2A and better developed ones in hindwing spaces Mi-Me, Me-M: and Cuz-2A. Tribe EUPTYCHIINI, new tribe (Figures 196-210) This tribe shows the greatest affinities to the Old World Ypthi- mini and was probably derived from a common stock with it. The euptychiines are less closely related to the Maniolini and Coeno- nymphini. The Euptychiini are separable from the Ypthimini on the basis of the slightly longer and more square-cut forewing cell and the ocellus in forewng space Mi-M3; and differ from the coenonymphines MEM. AMER. ENT. SOC., 24 90 THE SATYRIDAE 0.4XML FL ML HL 1x ML Euptychia Taygetis Pindis Paramecera Oressinoma | Fig. 196. Satyrinae: Euptychiini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip- pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measure- ments obtained from males, the bottom bar those from females. in that the latter have an excavate forewing cell and a greater abor- tion of the female foreleg, as well as a greater distention of forewing vein 2A. All of the genera definitely assigned to the euptychiines are found in the New World, from southern Canada through the Neotropics to temperate South America. Most are woods or forest dwellers, in contrast to most members of the Ypthimini-section. The most com- prehensive review of the tribe to date is included in Forster (1964), but, unfortunately, it is by no means complete. The Euptychiini are characterized below. The eyes are variously naked or hairy. The antennae are short, just under to just over two-fifths the length of the forewing costa. The antennal club is usually weakly developed and about twice the thickness of the shaft (about three times as thick in Cyllopsis). The third segment of the palpus is about a third as long as the second segment. The hairs of the second segment of the palpus are variable: in the tropical groups they are about three times as long as the segment is wide, but in the Nearctic Megisto cymela (Cramer) the hairs are almost five times the width of the segment. The male foreleg is reduced, with a monomerous, unspined tarsus; the femur and tibia are of about the same length. The female foreleg LEE D. MILLER 9] 199 iy. Figs. 197-205. Satyrinae: Euptychiini. 197. Euptychia mollina Hubner, é venation. 198. E. mollina, palpus. 199. E. mollina, 6 foreleg. 200. E. mollina, 2 foretarsus. 201. Cissia penelope (Fabricius), 2 foretarsus. 202. E. mollina, midleg. 203. Megisto cymela (Cramer), midleg. 204. Cyllopsis hedemanni R. Felder, 4 venation. 205. Taygetis mermeria (Cramer), 4 venation. is also reduced, with a pentamerous, clubbed tarsus bearing spines on the second, third and fourth subsegments (also on the first in at least Neonympha). The midtibia is one and a half to one and three- quarters times the length of the proximal midtarsal subsegment. The midtibia is variously smooth dorsad in most genera to slightly spiny in such genera as Neonympha and spiny in such genera as Cissia. The tibial spurs are present and well developed in most species, but MEM. AMER. ENT. SOC., 24 92 THE SATYRIDAE absent in Euptychia mollina Hiibner, and there is no spine at the dorsal, distal end of the midtibia. The relative lengths of the legs of selected Euptychiini are shown in Fig. 196. The forewing cell is generally square-cut and varies little from half the length of the forewing costa. The forewing radial veins arise from the cell in two branches (rarely one branch), and veins Rs and M; arise well separate. Vein Mz arises somewhat nearer Mi than Ms, and Cu: arises nearer M3 than Cuz. Forewing veins Sc and the cubital stem are basally inflated in all genera, and 2A is inflated in such genera as Taygetis and Oressinoma (Figs. 205, 210, respec- tively). The latter genus displays an unusual form of inflation of the veins. The hindwing cell is straight and produced at the origin of Ms; the cell is half to three-fifths the length of the wing measured to the end of Ms. Hindwing veins Sc+R: and 3A are of about the same length, Ms and Cu: arising well separate and Mp» arising nearer, occasionally much nearer, M: than Ms. The pattern is reasonably constant. Most species are brown above—they may be white or iridescent purple, blue in part—with or without ocelli. The under surface is usually brown with trans- verse darker markings and a more or less complete complement of ocelli. Oressinoma typhla Westwood is aberrant: white above and below bordered with blackish brown and with reddish lunules in the dark border. Genera Included in the Euptychiini \ ‘Names preceded by asterisks (*) are genera proposed too late for de- tailed study, but from the original descriptions they are judged to be members of this tribe. *Archeuptychia Forster, 1964: 80. Type-species: Papilio c’uena Drury, by original designation. *Caeruleuptychia Forster, 1964: 92. Type-species: Euptychia caerulea But- ler, by original designation. *Capronnieria Forster, 1964: 73. Type-species: Euptychia abretia Capron- nier, by original designation. *Cepheuptychia Forster, 1964: 96. Type-species: Euptychia cephus Butler, by original designation. *Chloreuptychia Forster, 1964: 119. Type-species: Papilio chloris Cramer,. by original designation. Cissia Doubleday, 1848b: 33. Type species: Papilio clarissa Cramer (=Pa- LEE D. MILLER 93 208 -~F Se 207 909 210 206 Figs. 206-210. Satyrinae: Euptychiini. 206. Paramecera xicaque (Rea- kirt), ¢ venation. 207. P. xicaque, palpus. 208. P. xicaque, 6 foreleg. 209. P. xicaque, 2 foretarsus. 210. Oressinoma typhla Westwood, 8 venation. pilio penelope Fabricius), by monotypy. =Argyreuptychia Forster, 1964: 123. Type-species: Papilio penel- ope Fabricius, by original designation. *Coeruleotaygetis Forster, 1964: 73. Type-species: Euptychia periboea God- man and Salvin, by original designation. Cyllopsis R. Felder, 1869: 474. Type-species: Cyllopsis hedemanni R. Felder, by monotypy. “Erichthodes Forster, 1964: 118. Type-species: Euptychia erichtho Butler, by original designation. Euptychia Hiibner, 1818 (1818-1825): 20. Type-species: Euptychia mol- lina Hiibner, designated by Hemming, 1937: 150. Butler (1868a: 194) designated Papilio herse Cramer as the type, based on Euptychia Hiibner, [1819](1816-1826): 54, but this designation was invalidated when the appropriate portions of the “Zutrage” were found to be older than those of the “Verzeichniss”’. *Euptychoides Forster, 1964: 97. Type species: Euptychia saturnus Butler, by original designation. *“Godartiana Forster, 1964: 118. Type-species: Satyrus byses Godart, by original designation. “Harjesia Forster, 1964: 78. Type-species: Taygetis blanda Moschler, by original designation. *“Haywardina Forster, 1964: 109. Type-species: Satyrus necys Godart, by original designation. Preoccupied by Haywardina Aczél, 1952, but no replacement name is proposed. *“Hermeuptychia Forster, 1964: 87. Type-species: Papilio hermes Fabricius, by original designation. MEM. AMER. ENT. SOC., 24 94 THE SATYRIDAE *“Magneuptychia Forster, 1964: 125. Type-species: Papilio libye Linné, by original designation. *Megeuptychia Forster, 1964: 122. Type-species: Nymphalis autonoe Cramer, by original designation. Megisto Hiibner, [1819](1816-1826): 54. Type-species: Papilio eurytus Fa- bricius (=Papilio cymela Cramer), designated by Butler, 1868b: 14. *Moneuptychia Forster, 1964: 92. Type-species: Euptychia soter Butler, by original designation. Neonympha Hiibner, 1818(1818-1825): 8. Type-species: Papilio areolatus Smith and Abbot, designated by Hemming, 1937: 150. Oressinoma Westwood, 1852, in Doubleday, Westwood and Hewitson, 1846- 1852: 371. Type-species: Oressinoma typhla Westwood, by monotypy. =Ocalis Westwood, 1852, in Doubleday, Westwood and Hewitson, 1846-1852: 371. Type-species: Oressinoma typhla Westwood, by monotypy. Listed in synonymy of Oressinoma Westwood. Paramecera Butler, 1868b: 98. Type-species: Neonympha xicaque Reakirt, by monotypy. *Parataygetis Forster, 1964: 79. Type-species: Taygetis albinotata Butler, by original designation. *Pareuptychia Forster, 1964: 83. Type-species: Papilio hesione Sulzer, by original designation. *Parypththimoides Forster, 1964: 106. Type-species: Neonympha_poltys Prittwitz, by original designation. *Pharneuptychia Forster, 1964: 91. Type-species: Satyrus phares Godart, by original designation. Pindis R. Felder, 1869: 475. Type-species: Pindis squamistriga R. Felder, by monotypy. *Posteuptychia Forster, 1964: 137. Type-species: Euptychia mycalesoides Felder, by original designation. *Posttaygetis Forster, 1964: 74. Type-species: Papilio penelea Cramer, by original designation. *Praefaunula Forster, 1964: 137. Type-species: Euptychia armilla Butler, by original designation. *Pseudeuptychia Forster, 1964: 86. Type-species: Euptychia languida Butler, by original designation. *Pseudodebis Forster, 1964: 75. Type-species: Papilio valentina Cramer, by original designation. *Rareuptychia Forster, 1964: 87. Type-species: Euptychia clio Weymer, by original designation. *Satyrotaygetis Forster, 1964: 73. Type-species: Euptychia satyrina Bates, by original designation. *Splendeuptychia Forster, 1964: 128. Type-species: Euptychia ashna Hew- itson, by original designation. *Taygetina Forster, 1964: 77. Type-species: Taygetis banghaasi Weymer, by original designation. LEE D. MILLER 95 Taygetis Hiibner, [1819](1816-1826): 55. Type-species: Papilio mermeria Cramer, designated by Westwood, 1852, in Doubleday, Westwood and Hewitson, 1846-1852: 355. Hemming (in press) accepts the designa- tion of Papilio virgilia Cramer as the type-species by Butler, 1868a: 194, but the intent of Westwood to designate mermeria as the type-species is unmistakable. *Vareuptychia Forster, 1964: 125. Type-species: Euptychia similis Butler, by original designation. *Weymerana Forster, 1964: 108. Type-species: Euptychia viridicans Wey- mer, by original designation. *Y pthimoides Forster, 1964: 100. Type-species: Neonympha yphthima Felder, by original designation. *Zischkaia Forster, 1964: 116. Type-species: Euptychia fumata Butler, by original designation. Tribe COENONYMPHINI, new tribe (Figures 211-222) The Coenonymphini are intimately related to the ypthimines and probably replace them in the north temperate zone savannas of both hemispheres. There are some relationships suggested with the maniolines, the latter group occupying scrub country environments: probably both evolved from a basic ypthimine stock. There is little to connect the coenonymphines with the Euptychiini or the Erebiini, two isolated tribes of the Ypthimini-section. The excavate forewing cell serves to distinguish most coeno- nymphines from most ypthimines, and the very long midtibia (over twice as long as the first midtarsal segment in all genera but A phan- topus) is unique in the Ypthimini-section. The female foreleg of the Ypthimini and the Euptychiini is fully formed, having five tarsal sub- segments, whereas that of the Coenonymphini is lacking at least one of the subsegments. The Coenonymphini are exclusively Holarctic and are grassland species. Almost all are temperate, few (e. g., Coenonympha mix- turata Alpheraky) being found so far north as the subarctic. The Coenonymphini are characterized as follows: The eyes are naked. The antennae are always shorter than half the length of the forewing costa, and those of Lyela and Triphysa are less than two-fifths the length of the forewing. The antennal club is variable in its development: in Aphantopus the club is only about MEM. AMER. ENT. SOC., 24 96 THE SATYRIDAE 0.2xML, FL ML HL IxML 4 : ss : Coenonympha Wit 'Z Lyela Z | ee Triphysa | == = | a Aphantopus | | Fig. 211. Satyrinae: Coenonymphini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip- pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measure- ments obtained from males. the bottom bar those from females. two and a half times as thick as the shaft, five times as thick in Tri- physa (Fig. 220) and seven times the thickness of the shaft in Lyela. The third segment of the palpus is generally half to one-third as long as the second, but the third segment is less than one-sixth the length of the second in Triphysa and Lyela. The hairs of the second seg- ment of the palpus are about four times as long as the segment is wide. The male foreleg is weakly developed with a monomerous, un- spined (weakly spined in Lyela) tarsus. The forefemur and fore- tibia are of about the same length; in some genera slight variations may be apparent. The female foreleg is reduced and is of two gen- eral types: in such genera as Coenonympha there are four tarsal sub- segments which are spined on at least the third and fourth ones, whereas in other genera like Chortobius, Lyela and Triphysa there is but one, unspined tarsal subsegment. With the exception of Aphantopus, the midtibia is very long, greater than twice as long as the first midtarsal subsegment. The tibial spurs are present and well developed in all genera but Triphysa, in which they are totally absent. The midtibia is smooth dorsad and has no spine at the dorsal, distal end. The hindlegs are significantly longer than the midlegs (Fig. PAs The forewing cell is deeply excavate and varies little from half - the length of the forewing costa. The forewing radial veins generally arise in two branches from the cell, but in three branches in A phan- LEE D. MILLER 97 topus; veins Rs and M;, are separate at their origins, although they may be approximate. Vein Mbp arises nearer M: than M3, and Cur arises midway between Mz; and Cup, or slightly nearer M;. Fore- 218 Figs. 212-222. Satyrinae: Coenonymphini. 212. Coenonympha oedippus (Fabricius), ¢ venation. 213. C. oedippus, antennal club. 214. C. oedippus, palpus. 215. C. oedippus, 4 foreleg. 216. Chortobius pamphilus (Fabricius), 2 foretarsus. 217. C. oedippus, midleg. 218. C. oedippus, 2 foretarsus. 219. Triphysa phryne (Pallas), é venation. 220. T. phryne, antennal club. 221. T. phryne, midleg. 222. Aphantopus hyperantus (Linné), é venation. wing veins Sc, the cubital stem and 2A are all inflated, though the latter may be only slightly thickened. The hindwing cell is square-cut distad and half to three-fifths the length of the wing to the end of Ms. Hindwing vein Sc+R: is MEM. AMER. ENT. SOC., 24 98 THE SATYRIDAE as long as, or longer than, 3A. Veins Ms and Cu: are separate, but close together, at their origins, and Mb» arises nearer, or much nearer, M; than Ms. The pattern is fairly constant. The upper surface is brown to tawny with few, if any, ocelli. The under surface of the forewings is likewise brown to tawny, and the hindwings are brown to grayish- olive, often with a light mesial band, with a more or less developed series of ocelli on both wings. Triphysa phryne (Pallas) is the only species to show marked sexual dimorphism: the male is brown with a more complete set of ocelli than is usual in the tribe, and the female is white with a full complement of ocelli. Genera Included in the Coenonymphini Aphantopus Wallengren, 1853: 30. Type-species: Papilio hyperantus (mis- spelled “hyperanthus”) Linné, by monotypy. Chortobius [Dunning and Pickard], 1858: 5. Type-species: Papilio pamphilus Linné, designated by Moore, 1893 (1890-1893): 51-52. Coenonympha Hiibner, [1819](1816-1826): 65. Type-species: Coenonympha oedipe Hiibner (=Papilio oedippus Fabricius), designated by Butler, 1868a: 194 (as Papilio geticus Esper). Dubierebia Muschamp, 1915b: 20. Type-species: Coenonympha myops Staudinger, by original designation. Lyela Swinhoe, 1908: 60. Type-species: Lyela macmahoni Swinhoe, by origi- nal designation. Sicca Verity, 1953: 83. Type-species: Papilio dorus Esper, by original desig- nation. Proposed as a subgenus of Coenonympha Hubner. Triphysa Zeller, 1850: 308. Type-species: Papilio tircis Stoll (=Papilio phryne Pallas), designated by Butler, 1868a: 194. =Phryne Herrich-Schaffer, 1844 (1843-1856): 90. Type-species: Papilio tircis Stoll (=Papilio phryne Pallas), by monotypy. Pre- occupied by Phryne Meigen, 1800. Tribe MANIOLINI Hampson, 1918 (Figures 223-232) Maniolinae Hampson, 1918: 385; Maniolidi Verity, 1953: 228, 237. This tribe is most closely related to the Coenonymphini; probably both tribes evolved in the Holarctic from a basic ypthimine stock. The maniolines are much more distantly related to either the Eupty- . chiini or the Erebiini. The maniolines are distinguished from other members of the LEE D. MILLER 99 Ypthimini-section by forewing vein Mz arising midway between Mi and M:; (in other tribes M» arises nearer M:), and by the distribu- tion of androconia. Whereas the Euptychiini and the Ypthimini E 0.2ML, FL ML HL IxML Maniola Cercyonis Fig. 223. Satyrinae: Maniolini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements obtained from males, the bottom bar those from females. have a pentamerous female foreleg, this tribe shows leg abortion in all genera, like the Coenonymphini. Cercyonis is considered by many authors (e.g., Hemming, in press) to be biologically synonymous with Minois Hiibner. The latter genus, however, is a member of the Satyrini, as shown by the pro- duction of the hindwing cell along vein Mz and the typically satyrine configuration of the midleg (Fig. 301). The midleg of Cercyonis is typical of the Maniolini (Fig. 228) and has nothing in common evolutionarily with Minois. The proper assignment of Cercyonis again demonstrates the necessity of looking beyond mere superficial similarities. The maniolines are found throughout the north temperate zone, and a few members are also found in the Arctic. These butterflies seem to replace the grassland Coenonymphini in scrublands. The Maniolini are characterized as follows: The eyes are naked. The antennae are greater than two-fifths, but less than half, the length of the forewing costa. The antennal club is weakly developed, from less than twice to about two and a half times the thickness of the shaft, and occupying the distal third to a quarter of the antenna. The third segment of the palpus is about one-third of the length of the second. The hairs of the second seg- ment of the palpus are long, about four to five times as long as the segment is thick. The male foreleg is greatly reduced, the femur and the tibia are MEM. AMER. ENT. SOC., 24 100 THE SATYRIDAE of approximately the same length and the tarsus is monomerous and unspined. The female foreleg is also greatly reduced with a mono- or dimerous tarsus without spines. The mid- and hindlegs are of 232 229 Figs. 224-232. Satyrinae: Maniolini. 224. Maniola jurtina (Linné), é venation. 225. M. jurtina, palpus. 226. M. jurtina, 6 foreleg. 227. M. jurtina, 2 foretarsus. 228. M. jurtina, midleg. 229. Pyronia tithonus (Lin- né), ¢ venation. 230. Cercyonis pegala (Fabricius), é venation. 231. C. pegala, & foreleg. 232. C. pegala, 2 foretarsus. about the same length (Fig. 223). The midtibia is somewhat less than twice as long as the basal midtarsal subsegment, without spines dorsad or at the dorsal distal end and with well developed tibial spurs. The forewing cell is somewhat excavate and rather uniformly half LEE D. MILLER 10] as long as the forewing costa. The forewing radial veins arise in three branches from the cell, and veins Rs and M: are separate at their origins. Vein Me arises about midway between Mi and Ms. and Cu: arises nearer M; than Cus. Forewing vein Sc is always inflated basally; the cubital stem and 2A may be greatly inflated (Maniola, Fig. 224) or not noticeably inflated (Cercyonis, Fig. 230). The hindwing cell is produced at the origin of Ms; and varies from less than half to as long as three-fifths the length of the wing measured to the end of Ms. Vein Sc+Ri is longer than 3A, and Ms and Cur are well separated at their origins. Vein Mp arises nearer Mi; than Ms. The upper surface is either tawny with brown margins or brown with or without a yellow discal patch on the forewing with the ocelli, if they are present, in the forewing spaces Mi-M; and Cus-2A and on the hindwing at the tornus. Beneath the wings are light brown with darker brown striae and moderately well developed ocelli on both wings. Genera Included in the Maniolini Cercyonis Scudder, 1875b: 241. Type-species: Papilio alope Fabricius (= Papilio pegala Fabricius), by original designation. Hyponephele Muschamp, 1915a: 156. Type-species: Papilio lycaon Rottem- burg, by monotypy. Idata deLesse, 1952: 72. Type-species: Pyronia cecilia Vallantin, by original designation. Proposed as a subgenus of Pyronia Hiibner. Maniola Schrank, 1801: 152. Type-species: Maniola lemur Schrank (=Papilio jurtina Linné), designated by Scudder, 1875a: 211. =Epinephele Hiibner, [1819](1816-1826): 59. Type-species: Pa- pilio janira Linné (=Papilio jurtina Linné), designated by Butler, 1868a: 194. Note that many authors have misspelled this name as “Epinephile” or “Epinephila’. Pasiphana de Lesse, 1952: 72. Type-species: Hipparchia bathseba Fabricius, by original designation. Proposed as a subgenus of Pyronia Hubner. Pyronia Hibner, [1819](1816-1826): 59. Type-species: Pyronia_ tithone Hubner (=Papilio tithonus Linné), designated by Scudder, 1875a: 261. Tribe EREBIINI Tutt, 1896 (Figures 233-246) Erebiinae Tutt, 1896: 87, 402. This tribe shows certain affinities with the Ypthimini and to the Satyrini-section, but is closer to the former; the similarity to the latter MEM. AMER. ENT. SOC., 24 102 THE SATYRIDAE is probably convergence. Structurally no group is exceptionally close to the erebiines, except the Callerebia-series of the Ypthimini. Erebia Medusia Phorcis Gorgo Atercoloratus Fig. 233. Satyrinae: Erebiini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements obtained from males, the bottom bar those from females. The erebiines may be distinguished from other satyrines of the Ypthimini-section by the extremely short third palpal segment, which is less than a fourth as long as the second segment in all genera. The facies of the species are characteristic, but, as indicated in the dis- cussions of the Dirini, the Hypocystini and the Pronophilini, some other genera outside the present tribe share the dark coloration, rounded wings and alpine habits with these butterflies. The very slight inflation of the forewing veins, even of Sc, is atypical of the Y pthimini-section. All of the erebiines are Holarctic, and many species are found in the high Arctic, as well as in alpine situations; few are at home in truly temperate climates. Warren (1936) has admirably mono- graphed this group. The Erebiini are characterized as follows: The eyes are naked. The antennae are from just over two-fifths to greater than half the length of the forewing costa. The antennal club is moderately well to very well developed, generally occupying the distal quarter of the antenna and inflated as much as five times the thickness of the shaft. The third segment of the palpus is very short, from one-quarter to one-sixth the length of the second. The LEE D. MILLER 103 246 244 245 Figs. 234-246. Satyrinae: Erebiini. 234. Erebia ligea (Linné), 6 vena- tion. 235. E. ligea, antennal club. 236. E. ligea, palpus. 237. E. ligea, 4 foreleg. 238. E. ligea, 2 foretarsus. 239. E. ligea, midleg. 240. Phorcis epistygne Hiibner, ¢ venation. 241. P. epistygne, stygne, 2 foretarsus. 243. P. epistygne, midleg. (Bang-Haas), ¢ venation. 245. A. alini, palpus. cluding trochanter and distal portion of coxa. 6 foreleg. 242. P. epi- 244. Atercoloratus alini 246. A. alini, $ foreleg, in- MEM. AMER. ENT. SOC., 24 104 THE SATYRIDAE hairs on the second segment of the palpus are long, between three and five times as long as the segment is wide. The male foreleg is reduced with a mono- or dimerous, unspined or weakly spined tarsus; the femur is generally as long as, or longer than, the tibia. The development of the female foreleg is variable, though the leg itself is always greatly miniaturized (Fig. 233). There are two to five tarsal subsegments with spines on all but the last one. The midleg is about as long as, and frequently longer than, the hindleg (Fig. 233). The midtibia is from just under to well over one and a half times the length of the first midtarsal sub- segment. The midtibia never bears a spine at its dorsal, distal end and is usually smooth dorsad, but slightly spiny in Phorcis (Fig. 243). The tibial spurs are present and well developed. The forewing cell is slightly excavate and varies little from half the length of the forewing costa. The forewing radial veins arise in three branches from the cell, and veins Rs and M; are well separated at their origins. Vein Me» arises much nearer M; than Ms, and Cur arises nearer Ms; than Cus. Forewing vein Sc is always more or less inflated, the cubital stem is somewhat inflated and 2A hardly at all. The hindwing cell is produced at the origin of Ms and is half to three-fifths as long as the wing measured to the end of Ms. Vein Sc+Ri is longer than 3A, and Ms and Cu: are well separated at their origins. Vein Me arises nearer M; than Ms. These are dark insects with very little pattern on either surface except scattered ocelli and a forewing tawny to russet shade; either or both pattern features may be absent. [Illustrations of the species are given by Warren (1936). Genera Included in the Erebiini Atercoloratus Bang-Haas, 1938: 178. Type-species: Coenonympha alini Bang- Haas, by original designation. Erebia Dalman, 1816: 58. Type-species: Papilio ligea Linné, by original designation. =Epigea Hiibner, [1819](1816-1826): 62. Type-species: Papilio ligea Linné, designated by Hemming, 1933: 198. Gorgo Hiibner, [1819](1816-1826): 64. Type-species: Papilio ceto Hubner, designated by Hemming, 1933: 198. Marica Hiibner, [1819](1816-1826): 63. Type-species: Papilio stygne Ochs- enheimer, designated by Hemming, 1933: 198. Medusia Verity, 1953: 179. Type-species: Papilio medus Schiffermiller, by original designation. Proposed as a subgenus of Erebia Dalman. LEE D. MILLER 105 Oreina Westwood, 1841: 76. Type-species: Papilio cassiope Fabricius, desig- nated by Butler, 1868a: 194. Phorcis Hiibner, [1819](1816-1826): 62. Type-species: Phorcis epistygne Hubner, designated by Hemming, 1933: 198. Simplica Verity, 1953: 194. Type-species: Papilio epiphron Knoch, by origi- nal designation. Proposed as a subgenus of Erebia Dalman. Syngea Hiibner, [1819](1816-1826): 62. Type-species: Papilio pronoe Esper, designated by Hemming, 1933: 198. Triariia Verity, 1953: 186. Type-species: Papilio triarius de Prunner, by original designation. Proposed as a subgenus of Erebia Dalman. Truncaefalcia Verity, 1953: 188. Type-species: Papilio aethiops Esper, by originial designation. Proposed as a subgenus of Erebia Dalman. Tribe DiRINI, new tribe (Figures 247-259) This tribe is not as primitive as the hypocystines, as may be seen form examination of the forelegs of both sexes, even though it seems to connect the remainder of the Satyrini-section with the South Afri- can members of the Lethini—a convergence which is probably ecological and with little phylogenetic significance. The forelegs of both sexes are greatly reduced, about as much so as in any tribe; the dirines are nevertheless primitive members of the Satyrini-section. There is little to connect the Dirini—or any member of the Satyrini- section, for that matter—with the Hypocystini except the configura- tion of the hindwing cell (Figs. 248, 254, 257). In common with the Satyrini, Pronophilini (usually to a lesser extent) and Melanar- giini are the greatly reduced forelegs, the spiny midtibiae and the well developed tibial spurs of the present tribe. Some species have a well developed dorsal, distal spine at the end of the midtibia (Fig. 259), a characteristic shared with the Satyrini and indicating a close rela- tionship between these two tribes. The Dirini are characterized chiefly by the short forewing cell, always less than half the length of the wing; in other Satyrinae the cell is half the length of the wing or greater. The forewing cell is also rounded and reminiscent of that of the Lethini. All members of this tribe are restricted to South Africa and are well illustrated by van Son (1955). The Dirini are characterized as follows: The eyes are hairy in all genera. The antennae are approximately two-fifths the length of the forewing costa. The antennal club is MEM. AMER. ENT. SOC., 24 106 THE SATYRIDAE Dira Tarsocera Torynesis Fig. 247. Satyrinae: Dirini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements ob- tained from males, the bottom bar those from females. abrupt, occupying the distal one-fifth to one-third of the antenna and three to five times as thick as the shaft (Fig. 249). The third segment of the palpus is usually about a third the length of the second, but is half as long in Cassus (Fig. 254) and only one-ninth as long in Torynesis (Fig. 257). The hairs of the second palpal segment are two or three times as long as the segment is wide. The male foreleg is greatly reduced (Fig. 247), particularly as regards the tibia and tarsus. The femur is always more than twice as long as the tibia, and the tarsus has one or two unspined sub- segments (Figs. 251, 258). The female foreleg is also greatly re- duced, frequently as much so as that of the male, with a mono- or dimerous, unspined tarsus (Fig. 252). The midleg is as long as, and occasionally much longer than (Torynesis), the hindleg (Fig. 247). The midtibia is almost twice as long as the first midtarsal subsegment in some genera such as Dira (Fig. 253), but in Tory- nesis (Fig. 259) and Dingana the midtibia is only slightly longer than is the proximal midtarsal subsegment. The midtibia is spiny dorsad and bears well developed tibial spurs; in most genera there is no spine at the dorsal, distal end of the midtibia, but the spine is weakly developed in Dingana. The configuration of the midleg in Dingana and Torynesis strongly suggests that of the Satyrini. The forewing cell is generally rounded distad and is greater than two-fifths, but less than half, the length of the forewing costa. The forewing radial veins arise from the cell in three branches, and Rs’ and M:; are well separated at their origins. Wein Me arises nearer Mi than Ms, and Cu; arises nearer Ms than Cuz. Only forewing vein LEE D. MILLER 107 250 252 254 256 X 255 Figs. 248-259. Satyrinae: Dirini. 248. Dira clytus (Linné), é venation. 249. D. clytus, antennal club. 250. D. clytus, palpus. 251. D. clytus, & fore- leg. 252. D. clytus, 2 foretarsus. 253. D. clytus, midleg. 254. Cassus cassus (Linné), palpus. 255. C. cassus, é venation. 256. Torynesis mintha (Geyer), 6 venation. 257. 7. mintha, palpus. 258. T. mintha, 8 foreleg. 259. T. mintha, midleg. Sc is inflated basad, and it is only slightly thickened. The hindwing cell is somewhat produced at the origin of Ms; and about half the length of the wing measured to the end of Ms. Hindwing veins Sc+R: and 3A are of about the same length, and veins Mz and Cu: arise separately, though they may be approximate. Vein Mz arises much nearer M; than Ms. MEM. AMER. ENT. SOC., 24 108 THE SATYRIDAE These butterflies are dark colored insects on the upper surface with ocelli in forewing spaces Mi-Ms; and Cus-2A. On the under surface the forewing ocelli are repeated and the hindwings are pat- terned by pale mottling and veinal markings. The general aspect is that of the Erebiini or the Satyrini. The species are figured by van Son (1955). Genera Included in the Dirini Cassus van Son, 1955: 79. Type-species: Papilio cassus Linné, by original designation. Dingana van Son, 1955: 70. Type-species: Leptoneura dingana Trimen, by original designation. Dira Hubner, [1819](1816-1826): 60. Type-species: Dira clyte Hiibner (= Papilio clytus Linné), designated by Scudder, 1875a: 157. =Leptoneura Wallengren, 1857: 31. Type-species: Papilio clytus Linné, by monotypy. Tarsocera Butler, 1899: 903. Type-species: Leptoneura cassina Butler, by monotypy. Torynesis Butler, 1899: 903. Type-species: Dira mintha Geyer, by monotypy. =Mintha van Son, 1955: 76. Type-species: Dira mintha Geyer, by original designation. Tribe PRONOPHILINI Clark, 1947 (Figures 260-294) Pronophilinae Clark, 1947: 149. This tribe was derived from a basic Satyrini-like stock of Old World origin but is more primitive than any present-day Satyrini, particularly as regards the forelegs, hence the pronophilines are placed before the satyrines in this discussion. In addition to the affinity with the Satyrini, the Pronophilini share some characteristics (though not particularly important ones) with the South African Dirini, which are, in turn, the most primitive tribe within the Satyrini-section. The pronophilines are only very distantly related to the Xenica-series of the Hypocystini; hence, any trans-Antarctic distribution of the family is doubtful. As in the Hypocystini and Dirini, convergent evolution toward the Holarctic Erebia is shown by several high-altitude pronophilines of the Lymanopoda-series. Several members of the Elina-series of. the present tribe show resemblance to members of the Holarctic LEE D. MILLER 109 0.4%ML, FL ML HL XML, | Pronophila|series, s. s. | Pronophila SS Daedalma Eteona Steroma Gyrocheilus Ba ocr Elina | series Elina Argyrophorus Neomaenas Nelia Lymanopoda Homoeonympha Sarromia Fig. 260. Satyrinae: Pronophilini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip- pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measure- ments obtained from males, the bottom bar those from females. Maniolini; indeed, they were long considered members of Maniola (= Epinephele), but on structural grounds these butterflies are per- fectly valid pronophilines. Many aberrant patterns have evolved in the great adaptive radiation of this tribe in the Neotropics: the Chilean MEM. AMER ENT. SOC., 24 110 THE SATYRIDAE species Argyrophorous argenteus Blanchard is silver above, while Lymanopoda samius Westwood is caerulean blue on the upper surface and members of Drucina resemble nymphalids or danaids more than they do other satyrids. The Pronophilini are the only Neotropical satyrids which have the hindwing crossvein mi-me produced distad at M2 and hindwing vein Sc+R: completely fused throughout its entire length. The present tribe can be distinguished from the Satyrini by the longer midtibia—never less than one and a half times the length of the first midtarsal subsegment—and the lack of a dorsal, distal midtibial spine. The more aborted forelegs serve to distinguish these butterflies from the Hypocystini. All members of the Pronophilini are Neotropical with the excep- tion of a single genus (Gyrocheilus) which penetrates the southern- most Nearctic. Many genera, particularly those of the Elina-series, are found in the temperate parts of South America. Three groups are recognized within the pronophilines: The Pronophila-series, sensu stricto, characterized by the gen- erally broad wings and rounded margins of the hindwings, except where modified by tails. The vast majority of these butterflies are found in tropical forests. The Elina-series, characterized by broad wings, but the hindwings are usually produced in the anal region. The forelegs are more reduced than in the Pronophila-series. Most of these butterflies are found in the South Temperate region. The Lymanopoda-series, characterized by the narrow wings, rounded hindwing margins and reduced forelegs. Most of these species are found in the Andean highlands. The Pronophilini are characterized as follows: The eyes are usually hairy, but are naked in such diverse genera as Argyrophorus, Amphidecta, Auca, Spinantenna, Manerebia, Idio- neurula and Gyrocheilus. The antennae are generally two-fifths to half the length of the forewing costa. The antennal club is well developed, occupying the distal one-fifth to one-third of the antenna and between two and five times the thickness of the shaft. The third segment of the palpus is long for the Satyrini-section, never less than one-fourth as long as the second segment and frequently over half. as long as the second in such genera as Lasiophila and Idioneurula. LEE D. MILLER iit 262 263 264 265 a. So —! —s 212 Figs. 261-272. Satyrinae: Pronophilini. 261. Pronophila thelebe Double- day and Hewitson, ¢ venation. 262. P. thelebe, antennal club. 263. P. the- lebe, palpus. 264. Drucina leonata Butler, palpus. 265. Amphidecta pignera- tor Butler, palpus. 266. P. thelebe, 6 foreleg. 267. P. thelebe, 2 foretarsus. 268. P. thelebe, midleg. 269. A. pignerator, midleg. 270. Pedaliodes poesia (Hewitson), ¢ venation. 271. P. poesia, $ foreleg. 272. P. poesia, 2 fore- tarsus. The hairs on the second palpal segment are highly variable in their lengths: in such genera as Eteona and Dioriste the hairs are less than twice as long as the segment is wide, whereas in other genera, MEM. AMER. ENT. SOC., 24 A THE SATYRIDAE such as Gyrocheilus and Manerebia, the hairs may be as much as five times the width of the segment. The male foreleg is moderately well developed in most genera with one (rarely two) tarsal subsegments that are devoid of spines 278 Figs. 273-279. Satyrinae: Pronophilini. 273. Catargynnis pholoe (Stau- dinger), ¢@ venation. 274. C. pholoe, palpus. 275. Corades enyo Hewitson, é venation. 276. Steroma bega Westwood, ¢ venation. 277. Daedalma dinias Hewitson, 6 venation. 278. D. dinias, palpus. 279. Argyrophorous | argenteus Blanchard, ¢ venation. LEE D. MILLER 113 (except in such genera as Cheimas, Gyrocheilus and Pronophila). The tibia is usually equal to, or longer than, the femur. Members of the Lymanopoda-series have greatly reduced forelegs with a mono- merous, unspined tarsus, and the femur is longer than the tibia. The female foreleg is usually moderately well developed with a penta- merus, clubbed tarsus bearing spines on the first four subsegments, but the foreleg of Gyrocheilus is greatly reduced with two unspined tarsal subsegments (Fig. 283). The relative lengths of the mid- and hindlegs are variable (Fig. 260). The mid-tibia is rarely less than twice as long as the first mid-tarsal subsegment, and it is gen- erally spiny dorsad, but smooth in such diverse genera as Amphidecta and Penrosada. The tibial spurs are present and well developed, and there is no spine at the dorsal, distal end of the midtibia. The forewing cell is usually excavate and half to three-fifths the length of the forewing costa. The forewing radial veins arise in three branches from the cell in most genera, but in such genera as Calisto, Steroma and Argyrophorus the radials arise in two, or even a single, stems; veins Rs and Mi arise connately (Jdioneurula, etc.) to widely separate (as Gyrocheilus and Pedaliodes). Vein M2 usually 82 283 Figs. 280-283. Satyrinae: Pronophilini. 280. Proboscis propylea (Hew- itson), ¢ venation. 281. Gyrocheilus patrobas (Hewitson), ¢ venation. 282. G. patrobas, palpus. 283. G. patrobas, 2 foretarsus. MEM. AMER. ENT. SOCc., 24 114 THE SATYRIDAE arises midway between Mi and M3 (but much nearer Mi: in the Lymanopoda-series), and Cui arises nearer M; than Cus in some genera, midway between the two veins in some and nearer Cuz in others. Forewing vein Sc is always inflated, often greatly distended, and the other stalks may also be—all stalks are inflated in such genera as Steroma and Spinantenna, for example. The hindwing cell is produced by the distad displacement of Mez-m; and half to over three-fifths the length of the wing measured to the end of Ms. Vein Sc+R: usually is longer than 3A, and M; and Cu: are well separated at their origins. Vein Me arises nearer, gen- erally much nearer, Ms; than Mu. The pattern is variable and highly modified, and there is con- vergence between some of the Lymanopoda-series and the Holarctic Erebia. Most of the species are figured by Weymer, 1912 (1910- LOM Dye Genera Included in the Pronophilini 2 * Several genera have not been seen or were described too recently for critical inclusion in this revision. From the original descriptions it has, how- ever, been possible to place them in their taxonomic positions. Such genera are preceded by an asterisk (*). Pronophila-series, sensu stricto * Altopedaliodes Forster, 1964: 148. Type-species: Pronophila tena Hewitson, by original designation. Amphidecta Butler, 1867a: 404. Type-species: Amphidecta pignerator But- ler, by monotypy. * Antopedaliodes Forster, 1964: 151. Type-species: Pedaliodes antonia Stau- dinger, by original designation. Calisto Hiibner, 1816(1806-1838): 16. Type-species: Calisto zangis Hubner, designated by Butler, 1868a: 194. Catargynnis R6éber, 1892: 284. Type-species: Daedalma pholoe Staudinger, designated by Hemming, 1943: 23. =Pseudomaniola Réber, 1892: 222. Type-species: Daedalma pholoe Staudinger, by original designation. Preoccupied by Pseudomani- ola Weymer, 1890. Cheimas Thieme, 1906: 175. Type-species: Oxeoschistus opalinus Staudinger, by original designation. Corades Doubleday, 1848a: 115. Type-species: Corades enyo Hewitson, by monotypy. *Corderopedaliodes Forster, 1964: 155. Type-species: Pedaliodes corderoi Dognin, by original designation. LEE D. MILLER 115 ( 285 291 292 Figs. 284-294. Satyrinae: Pronophilini. 284. Elina vanessoides Blanch- ard, ¢ venation. 285. E. vanessoides, palpus. 286. E. vanessoides, 3 fore- leg. 287. E. vanessoides, 2 foretarsus. 288. E. vanessoides, midleg. 289. Spinantenna tristis (Guérin), é venation. 290. Lymanopoda samius West- wood, é venation. 291. L. samius, antennal club. 292. L. samius, palpus. 293. Idioneurula erebioides (Felder and Felder), ¢ venation. 294. Manerebia cyclopina Staudinger, é venation. MEM. AMER. ENT. SOC., 24 116 THE SATYRIDAE Daedalma Hewitson, 1858(1856-1876): [85]. Type-species: Daeda!ma dinias Hewitson, designated by Butler, 1867f: 268. Dioriste Thieme, 1906: 171. Type-species: Pronophila tauropolis Westwood, by original designation. Drucina Butler, 1872: 72. Type-species: Drucina leonata Butler, by monotypy. Eretris Thieme, 1905: 131. Type-species: Pronophila decorata Felder and Felder, designated by Hemming, 1943: 24. Eteona Westwood, 1850, in Doubleday, Westwood and Hewitson, 1846-1852: 254. Type-species: Euterpe tisiphone Boisduval, by monotypy. Gyrocheilus Butler, 1867f: 267. Type-species: Pronophila patrobas Hewitson, by original designation. This name was misspelled “Geirocheilus” by Holland, 1898: 211. Lasiophila Felder and Felder, 1859: 325. Type-species: Lasiophi'a cirta Felder and Felder, designated by Scudder, 1875a: 203. *Muscopedaliodes Forster, 1964: 153. Type-species: Pedaliodes muscosa Thieme, by original designation. Oxeoschistus Butler, 1867f: 268. Type-species: Pronophila puerta Hewitson, by original designation. Panarche Thieme, 1906: 228. Type-species: Corades tricordata Hewitson, by original designation. *Panyapedaliodes Forster, 1964: 157. Type-species: Pronophila panyasis Hewitson, by original designation. *Parapedaliodes Forster, 1964: 153. Type-species: Pronophila parepa Hew- itson, by original designation. Pedaliodes Butler, 1867f: 267. Type-species: Pronophila poesia Hewitson, by original designation. *Pherepedaliodes Forster, 1964: 149. Type-species: Pedaliodes pheretiades Smith and Kirby, by original designation. *Physcopedaliodes Forster, 1964: 152. Type-species: Pronophila physcoa Hewitson, by original designation. Polymastus Thieme, 1906: 138. Type-species: Daedalma doraete Hewitson, by original designation. *Praepedaliodes Forster, 1964: 152. Type-species: Pronophila phanias Hew- itson, by original designation. *Praepronophila Forster, 1964: 182. Type-species: Pedaliodes emma Stau- dinger, by original designation. Proboscis Thieme, 1906: 168. Type-species: Pronophila propylea Hewitson, by original designation. Pronophila Doubleday and Hewitson, 1849, in Doubleday, Westwood and Hew- itson, 1846-1852: pl. 60. Type-species: Pronophila thelebe Doubleday and Hewitson, designated by Butler, 1867f: 266. =Mygona Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852; 357. Type-species: Pronophila thelebe Doubleday, designated by Hemming, 1939: 133. Pseudosteroma Weymer, 1912(1910-1912): 241. Type-species: Steroma pro- LEE D. MILLER I A17/ nophila Felder and Felder, designated by Hemming, 1943: 25. *Punapedaliodes Forster, 1964: 148. Type-species: Pedaliodes albopunctata Weymer, by original designation. Steremnia Thieme, 1905: 137. Type-species: Pedaliodes (?) polyxo Godman and Salvin, designated by Hemming, 1943: 25. Steroma Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846-1852: 400. Type-species: Steroma bega Westwood, by monotypy. *Steromapedaliodes Forster, 1964: 148. Type-species: Pedaliodes albonotata Godman, by original designation. Thiemeia Weymer, 1912(1910-1912): 267. Type-species: Pronophila phoro- nea Doubleday, designated by Hemming, 1943: 25. Elina-series Argyrophorus Blanchard, 1852: 30. Type-species: Argyrophorus argenteus Blanchard, by monotypy. Auca Hayward, 1953: 30. Type-species: Satyrus pales Philippi, by original designation. Elina Blanchard, 1852: 28. Type-species: Elina vanessoides Blanchard, desig- nated by Butler, 1868a: 194. *Haywardella Herrera, 1966: 71. Type-species: Satyrus thione Berg, by origi- nal designation. Nelia Hayward, 1953: 42. Type-species: Satyrus nemyroides Blanchard, by original designation. Neomaenas Wallengren, 1858: 78. Type-species: Neomaenas servilaea Wal- lengren, by monotypy. =Stibomorpha Butler, 1874: 204. Type-species: Stibomorpha deco- rata Butler (=Neomaenas servilaea Wallengren), by original des- ignation. Neosatyrus Wallengren, 1858: 79. Type-species: Neosatyrus ambiorix Wal- lengren, by monotypy. Pampasatyrus Hayward, 1953: 28. Type-species: Epinephele gyrtone Berg, by original designation. *Pseudomaniola Weymer, 1890: 107. Type-species: Pseudomaniola euripides Weymer, designated by Hemming, 1943: 25. =Neomaniola Hayward, 1949: 156. Type-species: Pseudomaniola euripides Weymer, by original designation. Proposed to replace Pseudomaniola Weymer, wrongly believed to be preoccupied. *Quilaphoethosus Herrera, 1966: 69. Type-species: Satyrus monachus Blan- chard, by original designation. Spinantenna Hayward, 1953: 38. Type-species: Satyrus tristis Guerin, by original designation. Stuardosatyrus Herrera and Etcheverry, 1965: 74. Type-species: Argyro- phorus williamsianus Butler, by original designation. MEM. AMER. ENT. SOC., 24 118 THE SATYRIDAE Lymanopoda-series Chillanella Herrera, 1966: 71. Type-species: Faunula stelligera Butler, by original designation. Cosmosatyrus Felder and Felder, 1867(1564-1867): 495. Type-species: Cos- mosatyrus leptoneuroides Felder and Felder, designated by Butler, 1868b: 59. Herrera (1965: 70) lists the type of this genus as “Cos- mosatyrus chiliensis chiliensis C. et R. Felder”, but chiliensis is a Guérin name. In any event, the type was properly designated by Butler. Etcheverrius Herrera, 1965: 62. Type-species: Satyrus chiliensis Guérin, by original designation. Faunula Felder and Felder, 1867(1864-1867): 488. Type-species: Faunula leucoglene Felder and Felder, by monotypy. Homoeonympha Felder and Felder, 1867(1864-1867): 487. Type-species: Homoeonympha pusilla Felder and Felder, by monotypy. Idioneurula Strand, 1942: 389. Type-species: Idioneura erebioides Felder and Felder, by original designation. =Idioneura Felder and Felder, 1867(1864-1867): 474. Type-species: Idioneura erebioides Felder and Felder, by monotypy. Preoccu- pied by Idioneura Selys, 1860. Lymanopoda Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 1852: 401. Type-species: Lymanopoda samius Westwood, designated by Butler, 1868a: 196. Manerebia Staudinger, 1897: 139. Type-species: Manerebia cyclopina Stau- dinger, designated by Hemming, 1943: 24. Palmaris Herrera, 1965: 67. Type-species: Hipparchia monticolens Butler, by original designation. Penrosada Brown, 1944: 255. Type-species: Lymanopoda leaena Hewitson, by original designation. *Sabatoga Staudinger, 1897: 143. Type-species: Sabatoga mirabilis Stau- dinger, by monotypy. Sarromia Westwood, 1851, in Doubleday, Westwood and Hewitson, 1846- 1852: 402. Type-species: Sarromia obsoleta Westwood, by monotypy. Stygnolepis Strand, 1942: 389. Type-species: Stygnus humilis Felder and Felder, by original designation. =Stygnus Felder and Felder, 1867(1864-1867): 487. Type-species: Stygnus humilis Felder and Felder, by monotypy. Preoccupied by Stygnus Perty, 1833. Tetraphlebia Felder and Felder, 1867 (1864-1867): 489. Type-species: Tetra- phlebia germainii Felder and Felder, by monotypy. Zabirnia Hewitson, 1877: 92. Type-species: Zabirnia zigomala Hewitson (= Lymanopoda acraeida Butler), by monotypy. =Trophonina Rober, 1892: 222. Type-species: Lymanopoda acrae- ida Butler, by original designation. LEE D. MILLER 119 Tribe SATYRINI Boisduval, 1836 (Figures 295-308) Satyrides Boisduval, 1836: 166. =Hipparchiadae Kirby, 1837: 297. This tribe is at the center of its section, showing relationships with the Dirini by virtue of the general facies and the presence of the heavy spine at the dorsal, distal end of the midtibia, and with the Pronophilini and much of the Hypocystini in the shape of the hindwing cell. Through Davidina there is a progressive gradation from typical Satyrini to the Melanargiini. 0.2xML, FL ML HL IxML, Satyrus Hipparchia Neominois Minois Karanasa Davidina Oeneis Oeneis Fig. 295. Satyrinae: Satyrini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stippling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measurements ob- tained from males, the bottom bar those from females. The weakly developed forelegs‘ of both sexes serve to distinguish this tribe from the Hypocystini and much of the Pronophilini, and the production of the hindwing crossvein m2-ms; at M2 will distinguish the satyrines from Melanargiini, Dirini and Erebiini. The most char- MEM. AMER. ENT. SOC., 24 120 THE SATYRIDAE acteristic features, however, of the Satyrini are on the midleg: all genera except Chionobas and Oeneis have a heavy spine at the dorsal, distal end of the midtibia, and in all genera but the Oeneis-series and Davidina the midtibia is less than one and one-fourth times as long as (and often shorter than) the first midtarsal subsegment. All members of the Satyrini are Holarctic. The Oeneis-series is found throughout the Holarctic, but the Satyrus-series is almost exclusively Palearctic: only Neominois is known from North America. There is a very interesting pattern similarity between the Euro- pean genus Minois and the North American Cercyonis, but while the European butterflies belong to the present tribe, Cercyonis is a member of the Maniolini. Several authors (e.g., Hemming, in press) have considered Cercyonis synonymous with Minois, but structural examination of the two genera does not confirm this conclusion, as noted in the discussion of Maniolini. Davidina is intriguing. Several authors have claimed it was a pierid, but examination of the tarsal claws shows Davidina to be a satyrid. Pierid tarsal claws are characteristically bifid (see Schatz and Rober, 1892), whereas those of Davidina are simple, as in most other satyrids. Two series are recognized in the Satyrini, as follows: The Satyrus-series, sensu stricto, distinguished by the extremely short midtibia and the presence of a heavy spine on the dorsal, distal end of the midtibia. The Ocneis-series, distinguished by the longer midtibia lacking the spine at the dorsal, distal end. The Satyrini are characterized as follows: The eyes are naked. The antennae are more than two-fifths, but less than half, the length of the forewing costa. The antennal club is well developed, occupying the distal one-fourth to one-fifth of the antenna and three to more than five times as thick as the shaft (Fig. 297). The third segment of the palpus is short, generally one-fourth to one-fifth as long as the second segment. The hairs of the second palpal segment are long, two and a half to five times as long as the segment is wide. The male foreleg is greatly reduced, the tibia and femur are of about the same length and the tarsus is mono- to trimerous without ° spines (Figs. 299, 306). The female foreleg is also greatly reduced, as much so as that of the male in several genera (Fig. 295), with LEE D. MILLER 121 Figs. 296-303. Satyrinae: Satyrini. 296. Satyrus actaea (Linné), ¢ venation. 297. S. actaea, antennal club. 298. S. actaea, palpus. 299. S. actaea, 6 foreleg. 300. S. actaea, 2 foretarsus. 301. S. actaea, midleg. 302. Brintesia circe (Fabricius), ¢ venation. 303. Neominois ridingsii (Edwards), é venation. the tarsus unspined, or only weakly spined, and comprised of one to four subsegments. The hindleg is significantly longer than the mid- leg (Fig. 295). In most genera the midtibia is less than one and one-fourth times the length of the proximal midtarsal subsegment, but in Davidina and the Oeneis-series the midtibia is at least half again as long as the first midtarsal subsegment. The midtibia is very spiny dorsad, the spines being particularly heavy in this tribe, and the tibial spurs are present and well developed. There is a charac- teristic heavy spine at the dorsal, distal end of the midtibia in the MEM. AMER. ENT. SOC., 24 122 THE SATYRIDAE Satyrus-series (Fig. 301), which is absent in the Oeneis-series. The forewing cell is square-cut to slightly excavate and half to almost three-fifths the length of the forewing costa. The forewing 306 f ; 305 a0 7 8 Figs. 304-308. Satyrinae: Satyrini, all Oeneis norna (Thunberg). 304. é venation. 305. palpus. 306. é foreleg. 307. 2 foretarsus. 308. midleg. radial veins arise from the cell in three branches, and veins Rs and Mi arise separate, but they may be approximate. Vein Mb arises nearer M; than Ms, and Cui arises midway between M3; and Cup, or slightly nearer the latter. Forewing vein Sc is inflated basad, but the other veins are little, if at all, thickened. The hindwing cell is produced by the distad displacement of me-ms and half to three-fifths (slightly more in Oeneis) the length of the wing measured to the end of Ms. Veins Sc+Ri and 3A are of about the same length, and Ms and Cu: are well separated at their origins. Vein Mp» arises nearer Ms; than Mi. The pattern is relatively uniform. The upper surface is shaded black, brown and/or orange and white with forewing ocelli in spaces M:-Ms and Cus-2A. The under surface pattern repeats the forewing ocelli, and the hindwings are cryptically marked. Davidina is white or off-white with dark brown to black veinal and interveinal lines. Illustrations of many of the species are given by Moore, 1893(1890- 1893). LEE D. MILLER 123 Genera Included in the Satyrini Satyrus-series, sensu stricto Arethusana de Lesse, 1951: 40. Type-species: Papilio arethusa Esper, by original designation. Aulocera Butler, 1867b: 121. Type-species: Satyrus brahminus Blanchard, designated by Butler, 1868a: 194. Berberia de Lesse, 1951: 41. Type-species: Satyrus abdelkader Pierret, by original designation. Brintesia Fruhstorfer, 1912(1912-1915): 307. Type-species: Papilio circe Fabricius, by original designation. Chazara Moore, 1893(1890-1893): 21. Type-species: Papilio briseis Linné, by original designation. Davidina Oberthiir, 1879: 19. Type-species: Davidina armandi Oberthir, by monotypy. Eumenis Hiibner, [1819](1816-1826): 58. Type-species: Papilio autonoe Esper, designated by Grote, 1873: 62. Hipparchia Fabricius, 1807: 281. Type-species: Papilio fagi Scopoli, desig- nated by Butler, 1868a: 194. =Nytha Billberg, 1820: 77. Type-species: Papilio hermione Linné (=Papilio fagi Scopoli), designated by Scudder, 1875a: 231. =Melania Sodoffsky, 1837: 81. Type-species: Papilio fagi Scopoli, by original designation. Preoccupied by Melania Lamarck, 1799. Proposed to replace Hipparchia Fabricius, wrongly believed to be preoccupied. Kanetisa Moore, 1893(1890-1893): 42. Type-species: Hipparchia digna Marshall, by original designation. Karanasa Moore, 1893(1890-1893): 38. Type-species: Satyrus huebneri Feld- er, by original designation. Minois Hiibner, [1819](1816-1826): 57. Type-species: Papilio phaedra Linné (=Papilio dryas Scopoli), designated by Butler, 1868a: 194. Neohipparchia de Lesse, 1951: 40. Type-species: Papilio statilinus Hufnagel, by original designation. Neominois Scudder, 1875b: 241. Type-species: Satyrus ridingsii Edwards, by original designation. Paroeneis Moore, 1893(1890-1893): 36. Type-species: Chionobas pumilus Felder, by original designation. Philareta Moore, 1893(1890-1893): 23. Type-species: Papilio hanifa Nord- mann (=Papilio anthe Ochsenheimer), by original designation. Pseudochazara de Lesse, 1951: 42. Type-species: Satyrus pelopea Klug, by original designation. Pseudotergumia Agenjo, 1947: unnumbered page at end. Type-species: Pa- pilio phidia Linné, by original designation. Satyrus Latreille, 1810: 355. Type-species: Papilio actaea Linné, designated MEM. AMER. ENT. SOC., 24 124 THE SATYRIDAE by the action of the International Commission on Zoological Nomen- clature, 1943, Opinion 142. Oeneis-series Chionobas Boisduval, [1833](1832-1843): 182. Type-species: Papilio aello Hubner (=Papilio glacialis Moll), designated by Blanchard, 1840: 457. Oeneis Hiibner, [1819](1816-1826): 58. Type-species: Papilio norna Thun- berg, designated by Butler, 1868a: 196. Tribe MELANARGIINI Verity, 1920 (Figures 309-314) Melanargiinae Verity, 1920: So. =Agapetinae Verity, 1953: 3, 46. This tribe contains some of the most distinctive of all satyrids, the pattern being unmistakable. In the shape of the hindwing cell, morphology of the legs and general facies (excluding the white ground O2xML, FL ML HL Ix ML, Melanargia Ledargia Argeformia Fig. 309. Satyrinae: Melanargiini. Relative lengths of the femur + tibia + tarsus of the forelegs (FL, diagonal lines), midlegs (ML, light stip- pling) and hindlegs (HL, heavy stippling) of selected genera. In all instances the ML value is unity. The top bar for each genus represents the measure- ments obtained from males, the bottom bar those from females. color), these butterflies closely resemble the Dirini and probably developed from a common stock with them. The spiny legs, reduc- tion of the forelegs and other structures suggest a strong affinity of the melanargiines and the Satyrini. There is little to relate this tribe closely either to the Hypocystini or to the pronophilines. The pattern of these butterflies sets them apart from all relatives —white with black marbling and well developed sets of ocelli on both wings, particularly the hindwings. All members of the Melanargiini are found in the Palearctic from LEE D. MILLER UZ Europe to China and Manchuria. There are more species in Asia than in Europe. The Melanargiini are characterized as follows: The eyes are naked. The antennae vary little from half the length of the forewing costa. The antennal club is rather weakly developed, less than two to less than three times the thickness of the shaft. The third segment of the palpus is one-third to one-fourth the length of the second segment. The hairs of the second segment of the palpus are about four times as long as the segment is wide. The male foreleg is greatly reduced, with a single, unspined tarsal subsegment; the femur is longer than the tibia (Fig. 312). The female foreleg is as reduced as that of the male with a monomerous, unspined tarsus (Fig. 313). The relative reduction of the fore-, mid- and hindlegs is shown in Fig. 309. The midlegs are usually shorter than the hindlegs (Fig. 309). The midtibia ranges from just under to just over one and a half times as long as the proximal midtarsal 312 @ 313 311 Figs. 310-314. Satyrinae: Melanargiini, all Melanargia galathea (Linné). 310. 6 venation. 311. palpus. 312. 6 foreleg. 313. 2 foretarsus. 314. midleg. 314 aa, subsegment and is very spiny dorsad. The tibial spurs are present and well developed; there is no heavy spine at the dorsal, distal end of the midtibia. The forewing cell is square-cut and approximately half the length of the forewing costa. The forewing radial veins arise in three branches from the cell, and veins Rs and M: are separate, but may MEM. AMER. ENT. SOC., 24 126 THE SATYRIDAE be approximate, at their origins. Vein Me arises much nearer Mi than Ms, and Cu: arises nearer Mz than Cue. If any forewing veins are inflated, only Sc would be, and that not extensively. The hindwing cell is produced at the origin of Ms and half to three-fifths the length of the wing measured to the end of M3. Veins Sc+R:i and 3A are of about the same length, and Ms and Cu: are well separated at their origins. Vein Mb arises slightly nearer Mi than Ms. The pattern is quite distinctive: all species are white or cream- colored marbled with black and with the ocelli well developed. Many genera have been named, but it seems that one genus would suffice biologically for all the species. Genera Included in the Melanargiini Argeformia Verity, 1953: 47. Type-species: Papilio arge Sulzer, by original designation. Proposed as a subgenus of Agapetes Billberg (=Mela- nargia Meigen). Epimede Houlbert, 1922: 132, 142, 160. Type-species: Arge halimede Méné- triés, designated by Hemming, 1934a: 143. Halimede Oberthtir and Houlbert, 1922: 192. Type-species: Halimede asiatica Oberthtir and Houlbert, by monotypy. Lachesis Oberthiir and Houlbert, 1922: 192. Type-species: Lachesis rusci- nonensis Oberthiir and Houlbert (=Arge lachesis Hitibner), by original designation. Ledargia Houlbert, 1922: 157. Type-species: Melanargia leda Leech, by original designation. Melanargia Meigen, 1829: 97. Type-species: Papilio galathea Linné, desig- nated by Kirby, 1894: 41. =Arge Hubner, [1819](1816-1826): 60. Type-species: Papilio psyche Hiibner (=Papilio occitanica Esper), designated by Butler, 1868a: 196. Preoccupied by Arge Schrank, 1802. This name was suppressed in favor of Melanargia Meigen by the International Commission on Zoological Nomenclature, 1956, Opinion 400. =A gapetes Billberg, 1820: 78. Type-species: Papilio galathea Linné, designated by Scudder, 1875a: 104. This name was suppressed in favor of Melanargia Meigen by the International Commission on Zoological Nomenclature, 1956, Opinion 400. Parce Oberthiir and Houlbert, 1922: 193. Type-species: Parce fergana Ober- | thiir and Houlbert (=Melanargia parce Staudinger), by original des- ignation. LEE D. MILLER 127 GENERA OF UNCERTAIN POSITION Genus Pamperis Heimlich, 1959 (Fig. 315) Pamperis Heimlich, 1959: 177. Type-species: Pamperis poaoeneis Heimlich, by original designation. Fig. 315. Venation of Pamperis poaoeneis Heimlich (after Heimlich, 1959). Note the aberrant venation pattern discussed in the text. Having seen no specimens of the type-species of this Neotropical monotypic genus, I can only evaluate its position in terms of the original description, which leaves many questions unanswered. Heim- lich’s photographs of the types suggest he was correct, at least superficially, in placing this genus near Cosmosatyrus Felder and Felder, and as such, it certainly belongs in the Lymanopoda-series of the satyrine tribe Pronophilini. However, the venational drawing given represents either an extraordinary aberration—the veins in the apical portion of the forewing are arranged like no known satyrid, and the hindwing lacks a major vein (Fig. 315)—or represent draft- MEM. AMER. ENT. SOC., 24 128 THE SATYRIDAE ing errors. Until these structures, as well as others not considered in the original description, have been studied, it will be quite im- possible to assign this genus to its proper position among the Satyridae. Genus Setodocis Billberg, 1820 Setodocis Billberg, 1820: 78. Type-species: Papilio periboea Fabricius, de- signated by Scudder, 1875a: 268. I have no idea to what this genus refers, and neither does anyone else; periboea is a “lost” species. The best opinion is that of Hem- ming (in press), “Include at the end of the subfamily Mycalesinae as a genus dubium.” The name is in the literature, so it must be men- tioned here, but it would seem advisable to suppress it. THE EVOLUTION AND ZOOGEOGRAPHY OF THE SATYRIDAE The twin subjects of evolution and biogeography have provided biologists since the days of Charles Darwin and Alfred Russel Wal- lace with the raw material for sundry mental gymnastics and manipu- lations of the earth’s flora, fauna, history and even crust through space and time. Such exercises, while based on the best available evidence, are limited by that evidence, and what are proposed as facts must be considered in actuality only as more or less educated guesses. The preceding statement is particularly true when one is dealing with such insects as butterflies, the fossil records of which are almost non- existent. In drawing evolutionary, and especially zoogeographic, in- ferences it is necessary to guess the past history of butterflies by com- parison with fossil histories of vertebrates and to interpret tendencies, such as the simplification of structures through phylogenetic lines, by the interpretations given such trends in groups with reasonably docu- mented fossil records. Zoogeography, in particular, has been a constant source of controversy, with several opposing schools of thought attempting to analyze distributional data in accordance with conflicting theories. At present there are two major camps of biogeographers which may be called the Wegener and the Matthew schools, although each theory has been refined far beyond its originator’s proposals. “Nothing in zoogeography has brought forth more argument or . more demands for an open mind—the other man’s mind—than the idea of continental drift.” (Darlington, 1957: 606). This theory, LEE D. MILLER 129 first put forth by Wegener, states briefly that the continents were a single landmass which broke apart, and the pieces drifted across the relatively plastic mantle, chiefly during the Mesozoic and into the Tertiary, to their present positions. By rotating the present-day continents and fitting them together it is possible to arrive at a con- vincingly completed “jig-saw puzzle”; indeed, it was this sort of piecing together of the landmasses of the world which provided the basis for the theory in the first place. Caster (1952) summarized the stratigraphic evidence supporting connections between South America and Africa, particularly during the Mesozoic, and further cited the similarity between the shallow water invertebrates of the two continents, but Dunbar (1952: 154) demonstrated by the simi- larity of shallow water molluscs of Kwajalein, Bikini and Guam that many shallow water invertebrates can cross deep water “barriers”, probably as free-swimming larvae. Darlington (1957: 607-608) showed another of the problems with the acceptance of the Wegener hypothesis as an explanation of the present-day animal distributions, when he stated, “. . . if a group of animals now occurs only in Africa and Brazil, it is said to date from and be evidence of a hypo- thetical African-Brazilian continent. This method depends on an assumption which Wegenerians usually do not put into words: that animal distributions are more permanent than land, that animals move less than continents.” This argument more or less summarizes my Own objections to continental drift as an explanation for the dis- tributions of present-day organisms. Most American zoogeographers today subscribe to the general theory of animal dispersal proposed by Matthew (1915) and elab- orated upon by Hesse, Allee and Schmidt (1951) and Darlington (1957). Matthew set forth his thesis in the beginning of his paper GUSIS 73). While the details of Matthew's work have been refined, the general skeleton has remained intact. Geologic evidence, such as that presented by Schuchert (1935) for the history of the Neotropics, has strengthened the Matthew theory. Biogeographers have been better able to devote their attention to the more knotty problems of how, when and why organisms arrived where they are. Since Schuchert (1935) showed the existence of several Ter- tiary “water gaps” between North and South America and roughly MEM. AMER. ENT. SOC., 24 130 THE SATYRIDAE dated them, many authors have considered it an absolute necessity to close these gaps physically so that organisms might cross them. This practice, however, has recently come under scrutiny from sev- eral authors, notably Simpson (1952), who demonstrated mathe- matically that given a length of time of sufficient duration and a sufficiently large “feeder” population even the most improbable dis- persal of a terrestrial organism over water could be a virtual certainty. While this mathematical exposition does not serve as evidence of a population of an area via waif dispersal, it shows a method by which such dispersal could be accomplished. It is of great importance to the understanding of the aerial dispersal of such insects as butterflies because it shows that landmasses need never be physically connected to exchange faunal elements across barriers. Such studies indicate that reappraisals are in order concerning the necessity of erecting land bridges across ocean deeps. Certainly physical barriers work less hardship on such strong flying butterfly genera as Danaus (Da- naidae), Precis and Vanessa (Nymphalidae), Libythea (Libytheidae ) and Leptotes and Lampides (Lycaenidae) than on the more sedentary Satyridae and Ithomiidae. I wish here to call attention to the second paragraph of Matthew’s thesis (1915: 173). Nearly all the principal routes of migration pass through either the eastern Palearctic or the Indo-Malayan region. If a group is in a rather generalized condition at the time of its expansion, it seems illogical that such a group would not leave some more or less primitive members along its primary path of dispersal. In the case of biotic interchange between the Palearctic and the Nearctic, the Paleotropics and the Neotropics, Asia and Australia, Africa and Australia, these waves of dispersal would have extended through eastern Asia. Eastern Asia also provides the complete spec- trum of biotopes, from Arctic tundra to tropical rainforest to the most arid desert—a prospective “home” for almost any group. No other region on earth is fed by so many major routes of dispersal, hence, no other region should be expected to have organisms derived from so many different sources. This diversity has led many students (e.g., Darlington, 1957) to consider eastern Asia the major center of dispersal, a veritable “Garden of Eden”. In the following pages I shall attempt to show that at least the Satyridae need not have arisen in this region, although eastern Asia certainly played an LEE D. MILLER 131 important role in the subsequent evolution of the family. ORIGIN OF THE SATYRIDAE Consideration of the origin and subsequent evolution of the Satyridae must be prefaced by a discussion of their relatives. Two nymphaloid families, the Morphidae (including the Amathusiidae ) and Ithomuidae, are closely allied to the family under consideration here and have been characterized at the beginning of the systematic revision. The evolution of all three families must be considered together to lay the groundwork for that which follows. The living nymphaloid families divide basically into two major groups on larval foodplant requirements. With very few exceptions (Ehrlich and Raven, 1965) the “true” nymphalids feed on dicoty- ledonous plants, whereas the larvae of the satyrids utilize monocoty- ledons exclusively. Forbes (1932) has assumed that the modern lepidopterous fami- lies had already differentiated by the Jurassic, and Carpenter (1930) felt that the Lepidoptera as an order could be traced only as far back as the Jurassic. Present evidence (Just, 1948) indicates that the proliferation and differentiation of the angiosperms took place in the late Mesozoic. The differentiation of the nymphaloid stock must have been somewhat synchronous with this proliferation of the flowering plants, but there is some question about this point. Fox (in litt.) feels that there is no reason to assume that morphological differentiation may not have preceded plant evolution, i.e., present monocotyledon feeders may have begun as dicotyledon feeders, or vice versa. Whether plant or butterfly evolution came first becomes a moot point, but the potential new niche provided by the evolution of the two basic stocks of plants must have provided some impetus for the subsequent, if not the initial, evolution of the butterflies. In any event, the butterflies are a relatively recent group compared with the Orthoptera and the Odonata, which were well developed by the Carboniferous. The Ithomiidae feed as larvae almost exclusively on the So- lanaceae (Fox, 1956: 13) and belong on this basis to the “true” nymphalids, although they are not far removed from the satyrid complex. These butterflies have reached their highest development MEM. AMER. ENT. SOC., 24 is2 THE SATYRIDAE in the Neotropics, although a single tribe, the Tellervini, is found in Australia and New Guinea. The Satyridae, sensu stricto, overwhelmingly feed as larvae on Gramineae and Cyperaceae, and all known satyrids feed on mono- cotyledons. Some members of the family feed on Palmaceae (Elym- nias and relatives). All known brassolines feed on monocots, generally utilizing the primitive ones, but a few also feed on bamboo; none are known to use dicots. The brassolines are exclusively Neo- tropical, none living farther north than Mexico. The foodplant requirements of the other primitive Neotropical satyrids (Haeterinae and Biinae) are unknown, but since they are restricted to the deep forest they probably do not use grasses, other than possibly bamboo. There are many other monocotyledons in the rainforest, and these are probably the food plants. Structurally, the Indo-Australian Amathusia, Faunis, Taenaris, etc. belong with the American Morpho in the family Morphidae. All the known Indo-Australian Morphidae feed on monocotyledons as larvae, many using Gramineae. In the New World, however, the morphids (represented only by the nominate genus) predominantly feed on dicotyledonous plants of the family Canellaceae, Myrtaceae, Menispermiaceae, Rhamnaceae, Sapindaceae and Leguminoseae. One species, however, Morpho aega (Fabricius), feeds on bamboo (Gramineae). Morpho is one of the few nymphaloid genera known which utilizes both monocotyledons and dicotyledons as larval food; indeed, the Morphidae is the only family commonly showing such habits. It seems evolutionarily significant that the genus Morpho is limited to the New World tropics. Furthermore, excluding the brightly colored and highly modified upper surface of these butter- flies, the morphids seem to typify the “primitive nymphalid pattern” of Schwanwitsch (1924): most of the elements of the pattern are present in the primitive species. In summary, many of the most primitive members of various nymphaloid families occur only in the Neotropics. The Neotropical tribe Tithoreini of the Ithomiidae is as primitive as the Indo- Australian Tellervini, and the two might well be combined were it not for their geographic isolation and a few morphologic characters (Fox, 1956: 20). As shown above, the Neotropical genus Morpho is the most primitive of the morphids. The most primitive satyrids are the Neotropical Haeterinae, which preserve the forewing vein 3A. LEE D. MILLER 133 Palearctic Nearctic (\ Biinae to Indo-Malayan Indo- Malayan Ethiopian Neotropics —> Haeterini ace Brassolini Madagascar Antirrhini Biini Australian / Fig. 316. The early distributional history of the Haeterinae, Brassolinae and Biinae in the New World. All movement illustrated in this figure was during the Cretaceous and earliest Tertiary (open arrows). In succeeding figures stippled arrows indicate mid-Tertiary movement, whereas solid black arrows indicate late Tertiary and Quaternary migration. Major areas of the world are indicated on this map for reference, and no attempt is made to present exact continental outlines at any given period of geologic time. The next most primitive groups of satyrids are the biines and the brassolines and the majority of these, including the most primitive ones, are New World species. In addition to these satyrids, the most primitive danaids (Clothilda and the Lycoreinae) are found in Central America and the Antilles. The remaining danaids are de- rived from Paleotropical groups which arrived in the Neotropics much later. In view of the evidence compiled in the preceding paragraphs, a feasible explanation of the evolution of the nymphaloid butterflies seems to require serious consideration of tropical America as some- thing other than a receptacle for pre-existing Holarctic groups. Prob- ably a relatively undifferentiated nymphaloid stock either entered the Neotropics as part of the extensive Holarctic invasion during the MEM. AMER. ENT. SOC., 24 134 THE SATYRIDAE Cretaceous (Darlington, 1957: 561-564), or this stock was already in tropical America prior to the connection of South America with the Holarctic. Since there are so many highly primitive groups of the Satyridae, Morphidae, Ithomiidae, and Danaidae in the Neo- tropics, it further appears that it was in tropical America that the fundamental bifurcation of the nymphaloid stock occurred. At least these four families appear to have arisen in the Neotropics. Later all four reinvaded the Holarctic, probably during the Cretaceous, but only two of them (Satyridae and Danaidae) reinvaded the American tropics during the Tertiary. The Tertiary history of the danaids probably parallels that of the satyrids, which will be discussed later. Endemic, primitive, though often highly specialized, stocks of all four families may be still found in the Neotropics. It seems most likely, therefore, that these families all arose in the Cretaceous (pos- sibly earlier) rainforests of tropical America. THE LATER HISTORY OF THE SATYRIDAE The most primitive satyrids, the Haeterinae, are now restricted to the Neotropical rainforest, and there is no indication that they ever were elsewhere, and all evolution in this group has occurred within the American tropical forests (Fig. 316). No members en- tered the Old World, as did members of the Ithomiidae. It is also doubtful that these insects ever exploited the grasslands that resulted from progressive drying during the Tertiary. In fact, these insects are singularly well adapted to the deep forest—the hyaline wings of many species and the ghost-like fluttering flight render them virtually invisible except when they pass through patches of sunlight. Masters (in litt.) says that Haetera always restricts itself to the deepest forest, never entering sunlit areas, but I have seen species of Pierella and Callitaera sunning themselves early in the morning, but not later in the day. Figs. 317, 318. Fig. 317. The origin and subsequent distributional his- tory of the Melanitini (Biinae) and the origin of the Elymniinae from the Melanitini. For details, see text. Fig. 318. The origin and proliferation of the Elymniinae into the tribes Lethini (see Fig. 319), Elymniini (Fig. 320), Mycalesini (Fig. 321) and Zetherini. The latter tribe did not significantly © leave the Indo-Malayan region. The probable origin of the Eritinae is also shown on this map. For details, see text. LEE D. MILLER 135 te World biines Elymniinae 317 0 Lethini (Fig. 319) ___ \Elymniini (Fig. 320) Elymniinae \ Mycalesini (Fig. 321) Satyrinae \|Zetherint \ Eritinae Sn ee 318 if MEM. AMER. ENT. SOC., 24 136 THE SATYRIDAE The history of the Brassolinae probably closely parallels that of the Haeterinae (Fig. 316). These butterflies, however, are strong fliers and have evolved a “flash coloration pattern” on the upper surface and a cryptic under surface. They do not depend so much on camouflage as on the strength of their flight. Many species are crepuscular, particularly those occurring in more open country, but most species are more or less restricted to the rainforest. The Biinae did move into the Holarctic and apparently provided the basis for the evolution of the rest of the family. Those biines which did not leave the Neotropics (Fig. 316) have remained rain- forest butterflies with patterns and flight characteristics similar to those of the brassolines. Whether the Neotropical members of any of the preceding sub- families ever invaded the grasslands is questionable. If they did, they were wiped out, probably in competition with the better adapted Satyrinae which later came into the New World. It seems more likely, however, that these butterflies never left the rainforest of their origin and evolved into such ecologically successful species that they easily resisted competition from the satyrines; perhaps these pri- mitive groups actually forced the satyrines into more open country. Those Biinae which did move into the Holarctic were obviously much more like Antirrhini than Bia, but there are now no New World antirrhines that have extremely close relationships with the Old World species. The migrating biines were forced into the Paleotropics at a fairly early time, where they apparently evolved into the present- day tribe Melanitini (Fig. 317). Several of the melanitines have become very widespread, apparently relatively recently, and one, Melanitis leda (Linné), is now found from West Africa to Mada- gascar and east into Australia and Japan. Most of the species are found in the Indo-Malayan region and in tropical Africa (Table 7). The Melanitini responded to the increase in aridity of the early Tertiary by invading open woodlands and even more open country, being thus the first satyrids, historically and evolutionarily, to leave the deep forest. These butterflies are more or less crepuscular, like the Brassolinae. A possible explanation for the crepuscular flight ~ Figs. 319, 320. Fig. 319. The distributional history of the Lethini. See text for details. Fig. 320. The distributional history of the Elymniini. See text for details. LEE D. MILLER 137 LI 2 — B Pararge-Series 6 > lethine stock tJ Lethe-series Aeropetes-series J 320 L MEM. AMER. ENT. SOC., 24 138 THE SATYRIDAE hairy-eyed mycalesines VA i | \\ Tertiary extermination on mainland i Fig. 321. The distributional history of the Mycalesini. See text for details. habit is that such species are actually approximating the conditions of the deep forest as regards light intensity. In short, such species are basically forest species that are “carrying the forest environment with them”. The successful invasion of the open country provided the impetus for the great adaptive radiation and phylogenetic diversi- fication of the satyrids in the Old World. The Neotropical Antirrhini, on the other hand, never became successfully adapted to open country, and without leaving the forests evolved into the present tribe with fewer than twenty species. In the Indo-Australian region the Melani- tini were probably giving rise, meanwhile, to the Elymniinae through steps resembling the present-day genera Parantirrhoea (Melanitini), Ptychandra and Samanta (both Lethini), shown in Figs. 317, 318. The Elymniinae proliferated rapidly during late Cretaceous and early Tertiary into the four tribes, most of which stayed in the wood- lands and scrublands: some (a number of Mycalesini) invaded the savannas, and others (the Neorina-series) re-entered the deep forests. - The pattern of early and later migration of the subfamily rather closely parallels that of many vertebrate groups and seems to have been LEE D. MILLER 139 limited northward by the Tethys Sea. Little movement from the Paleotropics seems to have been accomplished before the closing of this “water gap” between tne Paleotropics and the Palearctic (Fig. 318). Interrelationships within the Elymniinae can be better drawn than those within any other subfamily. The Lethini grade toward the Zetherini, on the one hand, and the Mycalesini (through such genera as Mandarinia and Orsotriaena), on the other, as brought out in the systematic revision. The Mycalesini connect well with the Elym- niini, as do the zetherines. The elymniines are “throwbacks” in one respect: all known members feed as larvae on Palmaceae, hence the common name “palmflies”. All tribes of the Elymniinae but the Zetherini are widely distributed in the Old World at present, although only one—the Pararge-section of the Lethini—has managed signifi- cantly to penetrate the Palearctic. The distributional histories of the tribes are shown in Figs. 319 (Lethini), 320 (Elymniini), 321 (Mycalesini); the actual patterns are discussed in the analyses of the individual faunae of the world. Apparently within a short time—probably still in the Cretaceous —the Elymniinae gave rise to the Satyrinae (Figs. 318, 322), pre- sumably through either the Mycalesini or (more probably) the Lethini and the Hypocystini of the Satyrinae. It is also probable that the Eritinae arose from a basic elymniine stock (Fig. 318) and pos- sible that the Ragadiinae did (Fig. 322), but the origin of the latter subfamily is among the most obscure in the family Satyridae. The hypocystine genus Lamprolenis, while a fairly typical member of that tribe, retains some very primitive characteristics similar to those of the Mycalesini (Fig. 148). There is a typical Mycalesis- like hair tuft on the hindwing, and the general facies embody a com- bination of mycalesine and hypocystine characteristics. The other case of similarity between the Elymniinae and Satyrinae is to be found in South Africa, where members of the A eropetes-series of the Lethini seem to grade into the Dirini (Figs. 67-72 and 248-259, respectively). It would be tempting to ascribe great evolutionary significance to the similarity, but the Aeropetes-series contains primi- tive lethines, whereas the Dirini appear to be fairly advanced Sa- tyrinae—particularly as regards the reduction of the forelegs of both sexes. Therefore, this similarity is ascribed to convergence, not to phyletic proximity. MEM. AMER. ENT. SOC., 24 140 THE SATYRIDAE The strongest link between the Elymniinae and the satyrines occurs through the subfamily Eritinae. Eritines have many charac- teristics in common with the Lethini, as discussed in the systematic revision, and also share characters with the Hypocystini. This can- not be merely convergence, and it appears that the Satyrinae arose in the Indo-Australian region from an elymniine stock not far re- moved from the lethine stock (Figs. 318, 322). In any event, the Satyrinae—probably from a Hypocystini-like stock—-soon split into two basic groups, the Ypthimini- and Satyrini- sections (Fig. 322). This dichotomy was probably brought about by the increasing aridity and proliferation of savannas during the early Tertiary. The Ypthimini-section generally occupies more open country than do the Satyrini-section butterflies. During the early to mid-Tertiary the various tribes of the Satyri- nae, except the Pronophilini and Euptychiini, probably evolved in the Old World (Figs. 323-326). Upon the reopening of the Cen- tral American land connection between North and South America— it was probably sufficiently opened in Miocene, perhaps Oligocene, times—the forerunners of the principal South American tribes of the Satyrinae, the Pronophilini (derived from the Satyrini-section, Fig. 325) and the Euptychiini (from the Ypthimini-section, Fig. 323) reentered South America and rapidly assumed dominance over the “old” South American fauna. At least part of this dominance may be ascribed to the ecological diversity shown by both tribes, as explained in the discussions of the tribes in the systematic revision. A glance at the numbers of species involved in both the Pronophilini and Euptychiini shows the dramatic radiation that has taken place in the Neotropics. It is doubtful that the radiation of the Euptychiini and Pronophi- lini resulted in a wholesale extinction of the Haeterinae, Brassolinae and Biinae, since the new arrivals probably did not compete ecologi- cally with the already established primitive ones. Both the Pro- Figs. 322, 323. Fig. 322. The origin of the Satyrinae from the Elymniinae and the division of the satyrines into the Hypocystini, the Ypthimini-section and the Satyrini-section. The probable origin of the Ragadiinae from the Elym- niinae is also shown in this figure. For details, see text. Fig. 323. The origin . and distributional history of the Ypthimini (Old World) and Euptychiini (New World). See text for details. LEE D. MILLER 141 Ypthimint-sect ini- "Sat rinae Satyrini-sect.¢ 24ly! = \ elymniine ancestor \, \ ve Hy pocystini Ragadlinae O \\ a Ny 322 Callerebia series == radiation MEM. AMER. ENT. SOc., 24 142 THE SATYRIDAE ~ Maniolini Coenymphini Erebiini i Fig. 324. The origin and distributional history of the Coenonymphini, Maniolini and Erebiini. See text for details. nophilini and Euptychiini feed as larvae predominantly on Grami- neae, which are probably not used extensively by the primitive species. Just before and during the Pleistocene there was an exchange of faunal elements between the Nearctic and the Palearctic, resulting in such Holarctic distributions as those of Coenonympha (Fig. 324), Oeneis (Fig. 326) and Erebia (Fig. 324), although these genera probably differentiated much earlier, and some of the Holarctic mem- bers of the Maniolini (Fig. 324) and Satyrini (Fig. 326). A proposed phylogeny for the Satyridae is presented in Fig. 327, embodying the information given above. DERIVATION OF THE SATYRID FAUNAE OF THE WORLD Comparisons of the faunal compositions of various parts of the world often prove valuable in determining the derivation of the elements of some areas. With this assumption in mind Table 7 was drawn to show the composition by tribes of the Satyridae in major faunal regions of the world. LEE D. MILLER 143 \\ Wn ee a \ EE Satyrini-sect. \A LEE Aa ol Pronophilini radiation O 325 [] =e Oeneis-Series 4} Satyrus-series Sa SSIES sy Melanareifi 1 Satyrini-sect. Figs. 325, 326. Fig. 325. The origin and distributional history of the Dirini and Pronophilini. See text for details. Fig. 326. The origin and dis- tributional history of the Satyrini and Melanargiini. See text for details. MEM. AMER. ENT. SOC., 24 144 TIME — approx. Cretaceous } early Tertiary calle | THE SATYRIDAE TAXON Tribe Subfamily Melanargiini . Satyrini & Pronophilini YS Dirini SS Hypocystini “ UD SATYRINAE Erebiini a 7 Maniolini y Coenonymphini ; Be Ypthimini Bo Euptychiini xe Ragadiinigeesoeee senna ane eee RAGADIINAE ear ERITINAE Mycalesini-. Elymniini ae ELYMNIINAE Zetherini Ss Lethini Melanitini J Brassolini-------------------------- BRAS SOLINAE Haeterini---------------------------- HAETERINAE Fig. 327. A proposed phylogeny of the Satyridae showing the relation- ships of the tribes. The time scales are very approximate and conform to the hypotheses proposed in the section on evolution and zoogeography. For details, see text. LEE D. MILLER 145 The Neotropics——The Neotropics, as broadly and somewhat incor- rectly defined to include Antarctic, temperate and tropical regions of South America, show a satyrid fauna composed of six tribes which are members of four subfamilies. Five of these tribes are endemic, and four of them—Haeterini, Brassolini, Antirrhini and Biini—are the most primitive in the family. The fifth endemic tribe, the Pro- nophilini, is a member of the Satyrinae and most closely related to the Holarctic Satyrini. The pronophilines probably reached, during the mid-Tertiary, South America and proliferated there. The Eup- tychiini are exclusively American satyrids most closely allied to the Ypthimini. They were probably derived from a common stock with the Ypthimini, introduced into the Neotropics and proliferated there contemporaneous with the Pronophilini. The euptychiines, while entirely American and overwhelmingly Neotropical, cannot be con- sidered an endemic tribe because seven species are found in the Nearctic, as far north as southern Canada. In summary the Neotropical satyrid fauna appears to have been derived from two sources. The Haeterinae, Brassolinae, Antirrhini and Biini evolved in tropical America from the basic satyrid stock which either arose in tropical America from a basic nymphaloid stock that was already there or evolved from such a stock that invaded tropical America during the Cretaceous. The Neotropics were then reinvaded, probably in the Oligocene, perhaps even in the Eocene, by two basically Paleotropical groups, the Pronophilini (derived from a Satyrini-section stock) and the Euptychiini (from a Ypthimini- section stock). There has been no extensive immigration in recent times from the Nearctic; indeed, most recent movement in the Amer- icas of the Satyridae has been northward from the Neotropics. The Nearctic—The Nearctic fauna has been derived primarily from the Neotropics and the Palearctic. Seven tribes are represented in North America, none endemic, and most species are very recent arrivals, many during the Pleistocene. By far the majority of the species in the Nearctic are assignable to Holarctic tribes or genera. A few of these, such as Neominois ridinsti, Enodia portlandia and creola, Satyrodes eurydice, Coenonympha haydenii and Cercyonis species, appear to be pre-Pleistocene arrivals; these butterflies prob- ably arrived in North America in the Miocene or Pliocene, perhaps even earlier. The remainder of the genera arriving from the Palearctic —Oeneis, most Coenonympha, Erebia—belong to a basically arctic- MEM. AMER. ENT. SOC., 24 146 THE SATYRIDAE TABLE 7 The taxonomic composition of the world satyrid fauna. a= single species; b = 2 to 5 species; c = 5 to 10 species; d = more than 10 species). () Pah 5 acy =I fi] be 13) & 8 8 28. oS, 23) ns 2. eS. Ssh ee ©) ie, Be Non A 7 Shy ae ei Taxon ZZ igh SS Haeterinae d Brassolinae d Biinae d a d a c b b Antirrhini d Biini a Melanitini a d a c b b Manataria b Elymniinae b d c d d d d Lethini b d d a b Zetherini a b Elymniini d b b b a Mycalesini C d d d c d Eritinae b Ragadiinae b Cc Satyrinae d d d d d d d d a Hypocystini d Y pthimini d b d b d d a Palaeonympha a Euptychiini d c Coenonymphini b d d Maniolini b d d Erebiini c d d Dirini c Pronophilini d a Satyrini d d d b Melanargiini d d Endemic tribes 5 0) l 2 1 0 1 0 alpine fauna and probably date from the Pleistocene: Coenonympha mixturata found in the bogs of Alaska and the Yukon is virtually inseparable from the Siberian population and must be a post-Pleisto- cene immigrant. The tribes derived from the Palearctic are the Lethini, Coenonymphini, Maniolini, Erebiini and Satyrini. Two LEE D. MILLER 147 tribes, the Pronophilini and Euptychiini, have entered the Nearctic from the Neotropics. The first of these tribes is represented by a single species, Gyrocheilus patrobas, which reaches southern Arizona. This invasion is probably of rather recent origin, perhaps post- Pleistocene, though it is difficult to assign dates to an invasion of a single species. The Euptychiini have left several species in North America, and one group—the Megisto cymela complex—has become rather distinct from its Neotropical progenitors. The cymela-group probably arrived well before the Pleistocene, whereas the other species in the Euptychiini are closely allied to tropical members and may have arrived during an interglacial. The Palearctic—The importance of the Palearctic, while it was not the place of the origin of the Satyridae, nevertheless cannot be overstressed with respect to subsequent evolution. Eleven tribes, representing four subfamilies, are found in this region, although the “true” Palearctic fauna is composed only of the Elymniinae and the Satyrinae. The other two subfamilies, the Biinae and the Ragadiinae, enter the Palearctic only in its southeastern part, were derived from the Indo-Malayan region and need be mentioned only in passing. The Elymniinae are represented by the tribes Lethini, Zetherini and Mycalesini—the last two being found just in the southeastern part of the area and also properly are part of the Indo-Malayan fauna. The tribe Lethini is represented chiefly by the Pararge-series—more boreal in its distribution than other lethine groups—which probably evolved in the mid-Tertiary and became cold-adapted. The inter- esting link between the Lethini and Mycalesini, Mandarinia regalis, is found in temperate China, suggesting that the latter tribe may once have been found much farther north than it is today. The Palearctic is the metropolis of most of the tribes of the Satyrinae, with six represented. Five of these tribes—Satyrini, Me- lanargiini, Coenonymphini, Maniolini and Erebiini—probably arose in the Palearctic itself in response to progressive drying, with the increase of grasslands, during the early Tertiary. Only the Melanar- giini are truly endemic, all other tribes having expanded into the Nearctic during the later Tertiary or Quaternary. The sixth satyrine tribe, the Ypthimini, is primarily Paleotropical, and the Palearctic ypthimines must have been derived from the tribes. One group within the ypthimines, the Callerebia-series, is exclusively Palearctic and probably arose from the ypthimine stem about the same time as the MEM. AMER. ENT. SOC., 24 148 THE SATYRIDAE other tribes mentioned above. All members of the Callerebia-series now are found under the stringent ecological conditions of the central Asian mountains; none has invaded the main part of the region. In sharp contrast with the Nearctic, the Palearctic has been very important in the later evolution of the Satyridae. Just why the Pale- arctic should have played such a role in the evolution of the Satyrinae is a puzzle: there were certainly vast savannas in the Nearctic during the Tertiary (Darlington, 1957), which proved to be the route of dispersal of the forerunners of the Pronophilini and Euptychiini and thus must have been occupied during the Tertiary. The primitive forms of the five Holarctic satyrine tribes are all Palearctic, not Nearctic. The evidence for these tribes having arisen in the Old World is overwhelming. The only tribe which is a candidate for arising in part in the Nearctic 1s the Satyrini: Neominois ridingsii is a rather primitive species, most nearly related to the Asian Karanasa. The great proliferation of the Satyrus-series (Satyrini) in central Asia argues strongly for its being considered the “birthplace” of the tribe. In all fairness, however, it must be pointed out that North America was the place of the greatest Tertiary development of the horses and camels (Darlington, 1957: 352-354), yet no native species of either group is presently found in North America. Such an evolutionary pattern may explain the satyrids—evolution of some stocks in the New World, spread of these stocks into the Old World, extinction of the Nearctic fauna and reinvasion and repopulation of the Nearctic by Palearctic groups. In the absence of a fossil record, I must accept the neontological evidence that the satyrine tribes evolved in the areas of their present greatest density and diversity. Parallels exist: Simpson (1953: 51-52) mentions the New World oreodonts and pronghorn “antelopes” and the Old World giraffes. None of these groups left the hemisphere of their origin. The Indo-Malayan region—This area, bounded roughly by India, the Philippines and Wallace’s line, has been likened to the “Garden of Eden” by generations of zoogeographers. Its satyrid fauna is strong only in moderately primitive groups. In all, nine tribes of five subfamilies are represented in the Indo-Malayan fauna. This is the area of the best development of the Ragadiinae and Eritinae, and these are to be considered truly endemic to the region. The melani- tines (Biinae) are best represented in the Indo-Malayan fauna and probably arose there or arrived at an early date. LEE D. MILLER 149 The Elymniinae have reached their highest development in this region, where the four tribes are best represented; all probably arose there. Those genera tending to connect tribes, with the exception of Mandarinia, are found on the Malay Peninsula. The bifurcation of the Mycalesini into naked and hairy eyed species took place in this region, and both groups are still found throughout the Indo-Malayan region. These have spread and provide some rather interesting data in the analysis of the African-Malgache fauna. Certain considera- tions, outlined in the systematic revision, led to the conclusion that the Lethini are the most primitive elymniines. Most lethines have hairy eyes, and for this reason the hairy-eyed condition is considered primitive in the Mycalesini (see also the discussion under the Mal- gache fauna, which follows). The Satyrinae are represented by two tribes, the Satyrini and Ypthimini. The former is a secondary immigrant from the Palearctic, only entering the Indo-Malayan region along its northern end. The Ypthimini, however, probably arose in this region and apparently replaced its immediate relatives—forms closer to the present-day Hypocysta-series (Hypocystini), which is now restricted to the Aus- tralian region east of Wallace’s line. The ypthimines spread during the early Tertiary into Africa and the New World, giving rise to the Euptychiini of the latter region. In summary, due to the prevalence of the Melanitini, Elymniinae and intermediate genera in this region, the Indo-Malayan region is considered to have given rise to the elymniines. The Eritinae and Ragadiinae also arose in this area. The Indo-Malayan region did not play an important role in the later evolution of the Satyrinae: the focus of that subfamily shifted to the Palearctic. The Australian Region (including the Pacific Islands).—Faunistically the Australian region, including Australia, New Guinea, New Zea- land and the nearby islands, is one of the most interesting. It is distantly related to the Indo-Malayan region, but substantial inter- change of faunal elements between these regions occurred only in the distant past. Simpson (1961b: 443) has dated the extensive immi- gration of the marsupials into Australia as Cretaceous or Paleocene, and it was probably at that time that much of the regional butterfly fauna arrived. Elsewhere Simpson (1961b: 435-436) shows that several waves of rodent groups arrived in the Australian region from MEM. AMER. ENT. SOC., 24 150 THE SATYRIDAE the Indo-Malayan from Miocene time to the present. All the tribes represented in the Australian region have also penetrated the Pacific islands, over water, hence the islands are considered along with the continent. Just one tribe is endemic in this region, the Hypocystini. The metropolis of the Xenica-series is Australia, with some aberrant spe- cies in New Zealand. All the satyrids in New Zealand were derived long ago from the Australian ones. The Hypocysta-series is best developed in New Guinea and has spread into the Pacific islands. The other satyrine tribe represented in this region is the Ypthimini, the members of which are highly modified, particularly the Pacific island genus Xois. The Ypthimini, as shown by their expansion into Africa and the Americas, was a fairly plastic group. A single Melanitis is found throughout the region in question, M. leda, a widespread species distributed west to the westernmost part of Africa. The same may be said about those Elymniini which occur on these islands; all are widespread species. The arrival of Melanitis and Elymnias must have been a recent event since they are little differentiated from their continental progenitors. The Myca- lesini of the region reflect the probably early spread of this tribe, since it also reached Madagascar. In summary, the satyrid fauna of the Australian region was derived from the Indo-Malayan region, and this derivation very early, resulting in a highly distinctive fauna. If the time schedule for the marsupials is valid for the satyrids, most of the groups reached the region in late Cretaceous to earliest Tertiary times. This indi- cates that the Hypocystini of the most evolved subfamily, Satyrinae, were in existence by the earliest Tertiary. A few strong flying species, such as M. leda, probably arrived over water very recently. The Ethiopian Region.—This region includes both the tropical part of Africa and temperate South Africa. These are faunistically very different areas, but they are so intimately related that they must be considered together. There is a single endemic African satyrid tribe, the Dirini. This tribe, which has morphological affinities with both the Elymniinae and the Satyrini-section of the Satyrinae, is restricted to South Africa. Several interpretations are possible for the Dirini: (1) the Satyrinae arose in Africa, (2) the Dirini were exterminated in all places but LEE D. MILLER 151 South Africa or (3) the Dirini stock was isolated in South Africa with the expansion of the tropical rainforest belt into eastern Africa, and it evolved there. The first alternative is highly improbable, as I have shown in the discussion of the Hypocystini in the systematic revision, but either (2) or (3) might be possible. In the absence of a fossil record we shall never really know. The only other tribe of the Satyrinae represented in the African fauna is the Ypthimini. The ypthimines of tropical Africa are closely related to the Indo-Malayan ones, whereas most of the South African ypthimines belong to the more or less endemic Melampias-series, one member of which is also found on Madagascar. It seems likely that the Melampias-series were early invaders of Africa—contemporaneous with the invasion of the Dirini—which differentiated and were then forced south and were replaced in tropical Africa by the ypthimines that arrived later. The Lethini show a similar pattern. In southern Africa two very primitive lethines, the Aeropetes-series, are found which prob- ably are remnants of an early fauna. The East and South African genus Aphysoneura is much more like the Indo-Malayan lethines, and it was probably a later faunal addition. The Mycalesini have made a significant invasion of Africa. All the species on the conti- nent have naked eyes, unlike those on Madagascar, and probably represent a later immigration, contemporaneous with the arrival of the “conventional” ypthimines and Aphysoneura. The mycalesines will be considered in further detail in the discussion of the fauna of Madagascar. The elymniines and the melanitines of Africa are rep- resentatives of the most widespread groups and almost certainly arrived from the Indo-Malayan region, probably fairly recently: frequently the African members are conspecific with their Indo- Malayan counterparts. In summary, Africa appears to have been populated more than once from the Indo-Malayan region. The tropical African satyrid fauna looks more like a depauperate Indo-Malayan fauna than any- thing else, but that of temperate Africa has evolved significantly and is quite distinct. Probably the South African endemic groups rep- resent the “old” African fauna of the early Tertiary, whereas that of tropical Africa is younger, perhaps dating from the Oligocene or even the Miocene (Darlington, 1957: 590). Such species as Mela- nitis leda probably arrived in tropical Africa much more recently. MEM. AMER. ENT. SOC., 24 [52 THE SATYRIDAE Madagascar.—There can be little doubt that the colonization of Madagascar was from Africa, but mostly so long ago that the species bear little resemblance to species now found on continental Africa. The Melanitini and the Elymniini of this island, on the other hand, are conspecific with their mainland counterparts, as is the only mem- ber of the Satyrinae, a species of Ypthimini: all these butterflies are probably recent introductions, no doubt by waif dispersal. The most interesting satyrids in Madagascar, from a zoogeo- graphic standpoint, are the Mycalesini—the dominant tribe on the island. As noted before, both naked- and hairy-eyed mycalesines occur in the Indo-Malayan regions, whereas those from continental Africa have naked eyes. The species found in Madagascar, without exception, have hairy eyes. One would be tempted at first glance to postulate that the Malgache mycalesines were derived directly from the Indo-Malayan ones. This is precisely why the mythical continent of “Lemuria” was postulated: a land bridge to account for the similar Malgache and Indo-Malayan lemurs; the Madagascar genera are less like continental African genera than any of the continental genera are like each other. In the mycalesines, dissimilarities indicate a long isolation of the Malgache fauna. For this reason it seems more likely that the mycalesines migrated into Africa very early (Creta- ceous or Paleocene?), and that stock which made the first immigra- tion was hairy-eyed, as is characteristic of primitive elymniines. Fur- thermore, the Malgache mycalesines have a less clubbed female foretarsus than do the continental species. These mycalesines were contemporaneous with the Dirini, Melampias-series of the Ypthimini and the Aeropetes-series of the Lethini and formed part of the “old” African satyrid fauna. Madagascar received its mycalesine fauna from this very early invasion. Just when this occurred is problematic: Matthew (1915: 204-205) thought that Madagascar and Africa were never physically connected, and Darlington (1957: 535) be- lieved that the Malgache fauna may have begun to accumulate in the Oligocene. In any event, the Malgache mycalesine fauna is probably of early Tertiary age. Climatic changes or pressure of competition by the later mycalesine immigrants—which did not reach Madagascar— replaced the hairy-eyed mycalesines on mainland Africa, leaving the morphologically primitive Malgache fauna completely isolated geo- LEE D. MILLER 153 graphically from their nearest relatives in the Indo-Malayan region. Such invasion and subsequent extermination of geographically inter- mediate populations is well known among fossil vertebrates (Dar- linetonts 957) 102212 1-15 9=172-7206-230)): BIBLIOGRAPHY AGENJO, R. 1947. Catalogo ordenador de los lepiddpteros de Espana. Gra- ellsia, 5(3): unnumbered pages at end. DALMEIDA, R. F. 1951. 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Eine neue Gattung und zwei neue Arten aus der Familie der Satyriden. Ent. Zeits. Stuttgart 23: 163-165. 1910-1912. Satyridae. in Seitz, A., Die Grossschmetterlinge der Erde, vol. 5 (Die Amerikanische Tagfalter). Stuttgart, Alfred Ker- nan Verlag: pp. 173-283; ill. Woop-Mason, J. 1880. Description of Parantirrhoea Marshallii, the type of a new genus and species of rhopalocerous Lepidoptera from South India. Jour. Asiatic Soc. Bengal 49(2): 248-250. ZELLER, P.C. 1850. Zwei neue Tagfalter. Stettiner Ent. Zeitg. 11: 308-313. MEM. AMER. ENT. SOC., 24 164 THE SATYRIDAE INDEX TO TAXA The numbers in bold face type indicate that the taxon is figured on that page. names. abdelkader Pierret, 123 abeona Donovan, 81 abretia Capronnier, 92 achanta Donovan, 79, 81 acraeida Butler, 118 Acrophtalmia Felder & Felder, 67, 68, 69 “Acrophthalmia’ Felder & Felder, 69 Acropolis Hemming, 69 actaea Linné, 121, 123 actoriaena Linné, 32-34 Admiratio Hemming, 58, 62, 63 adolphei Guérin, 63 adoptiva Weymer, 31, 32 aega Fabricius, 132 aegeria Linné, 48, 50 aello Hiibner, 124 Aeropetes Billberg, 43, 44, 46, 47, 48, 52 Aeropetes-series, 40, 41, 42, 43, 47, 72, 137, 139, 151, 152 aethiops Esper, 105 Agapetes Billberg, 126 Agapetinae, 124 agondas Boisduval, 56, 57 Agrusia Moore, 56 albinotata Butler, 94 albonotata Godman, 117 albopunctata Weymer, 117 alini Bang-Haas, 103, 104 alope Fabricius, 101 Altopedaliodes Forster, 114 Amathusia Fabricius, 132 Amathusiidae, 131 ambiorix Wallengren, 117 Amecera Butler, 50 Amechania Hewitson, 51, 52, 53 Amphidecta Butler, 110, 111, 113, 114 amphirhoe Hubner, 27 Anadebis Butler, 50 anaxias Hewitson, 63 anaxioides Marshall & deNicéville, 63 Anchiphlebia Butler, 31 andromacha Hubner, 47 “andromeda Fabricius’, 22 anthe Ochsenheimer, 123 Those names preceded by a + mark are preoccupied antipodum Doubleday, 79, 80 Antirrhea Hiibner, 9, 30, 31, 32 Antirrhini, 26, 28, 29-32, 34, 35, 36, 37, 133, 136, 138, 144, 145, 146 antonia Staudinger, 114 Antopedaliodes Forster, 114 Apaturina Herrich-Schaffer, 19 Aphantopus Wallengren, 95, 96, 97, 98 Aphysoneura Karsch, 43, 47, 48, 151 aramis Hewitson, 62 Aranda Fruhstorfer, 49 arcesilaus Cramer, 31 archaea Hiibner, 31, 32 Archeuptychia Forster, 92 Archondesa Moore, 47 areolatus Smith & Abbot, 94 arethusa Esper, 123 Arethusana deLesse, 123 tArge Hiibner, 126 Arge Schrank, 126 arge Sulzer, 126 Argeformia Verity, 124, 126 argenteus Blanchard, 110, 112, 117 Argestina Riley, 87 Argus Bohadsch, 49 t Argus Scopoli, 49 Argynnina Butler, 75, 77, 80 Argyreuptychia Forster, 93 Argyronympha Mathew, 77, 80 Argyrophenga Doubleday, 74, 75, 78, 79, 80 Argyrophorus Blanchard, 109, 110, 112, 113, 117 armandi Oberthiir, 123 armilla Butler, 94 Arpidea Duncan, 31 artemis Felder & Felder, 68, 69 ashna Hewitson, 94 asiatica Oberthiir & Houlbert, 126 asochis Hewitson, 63 Atercoloratus Bang-Haas, 102, 103 atreus Kollar, 24 Auca Hayward, 110, 117 Aulocera Butler, 123 LEE D. MILLER “aurora Felder’, 22 automedon Cramer, 27 autonoe Cramer, 94 autonoe Esper, 123 baladeva Moore, 47 baldus Fabricius, 85 banghaasi Weymer, 94 bathseba Fabricius, 101 bega Westwood, 112, 117 Berberia deLesse, 123 bhadra Moore, 49 Bia Hiibner, 29, 30, 32-34, 136 Bicyclus Kirby, 58, 62, 63 Biina, 3, 4, 28, 32 Biinae, 3, 4, 8, 9, 11, 12, 13, 16, 17, 18, 19, 23, 28-36, 39, 72, 132, 133, 135, 136, 140, 144, 146, 147, 148 Biini, 10, 28, 29, 30, 32-34, 133, 144, 145, 146 Blanaida Kirby, 49 blanda Moschler, 93 Bletogona Felder & Felder, 40, 57, 58, 59, 60, 61 Boeberia Prout, 86, 87 bolanica Marshall, 85 Bolboneura Godman & Salvin, 16 brahminus Blanchard, 123 Brassolidae, 4, 23 Brassolides, 2, 23 Brassolina, 3, 4 Brassolinae, 2, 3, 4, 8, 9, 11, 13, 16, 17, 18, 19, 23-28, 29, 72, 133, 136, 140, 144, 145, 146 Brassolini, 23-28, 133, 144, 145 Brassolis Fabricius, 24, 25, 26, 27 Brassolis-series, 25, 26, 27 Brintesia Fruhstorfer, 121, 123 briseis Linné, 123 Bruasa Moore, 54, 56 butleri Fereday, 79, 80 byses Godart, 93 Caerois Hiibner, 8, 30, 31, 32 caerulea Butler, 92 Caeruleuptychia Forster, 92 Caliginae, 3, 4, 23 Caligo Hiibner, 23, 24, 25, 26, 27 MEM. AMER. ENT. SOC., 24 Caligo-series, 25, 27 Calisto Hiibner, 113, 114 Callarge Leech, 51, 52, 53 Callerebia Butler, 82, 86, 87 Callerebia-series, 83, 87, 102, 141, 147, 148 Callicore Hiibner, 16 Callinaga Moore, 19 callipteris Butler, 47 Callitaera Butler, 22, 134 Callyphthima Butler, 86 Calysisme Moore, 61 Capronnieria Forster, 92 cassiae Linné, 24 cassina Butler, 108 Cassionympha vanSon, 86 cassiope Cramer, 27 cassiope Fabricius, 105 cassius Godart, 86 Cassus vanSon, 106, 107, 108 cassus Linné, 107, 108 Catargynnis Rober, 112, 114 Catoblepia Stichel, 25, 27 cecilia Vallantin, 101 Celebina Fruhstorfer, 61 Cepheuptychia Forster, 92 cephus Butler, 92 Cercyonis Scudder, 99, 100, 101, 145 ceto Hiibner, 104 Charma Doherty, 47 Chazara Moore, 123 Cheimas Thieme, 112, 114 chenui Guérin, 85 “chiliensis C. et R. Felder’, 118 chiliensis Guérin, 118 Chillanella Herrera, 118 Chionobas Boisduyal, 120, 124 Chloreuptychia Forster, 92 chloris Cramer, 92 Chonala Moore, 49 Choranesa Moore, 47 chorinaeus Fabricius, 31, 32 Chortobius [Dunning & Pickard], 96, 97, 98 christophi Leech, 49 circe Fabricius, 121, 123 cirta Felder & Felder, 116 Cissia Doubleday, 91, 92 120, 166 Cithaerias Hibner, 20, 21, 22 clarissa Cramer, 92 clio Weymer, 94 Clothilda Blanchard, 133 cluena Drury, 92 clyte Hubner, 108 clytus Linné, 107, 108 Coelites Westwood & Hewitson, 64, 65, 66, 67, 72 Coenonympha Hibner, 96, 97, 98, 142, 145 Coenonymphini, 9, 10, 11, 70, 73, 74, 75, 81, 89, 95-98, 99, 142, 144, 146, 147 Coenyropsis vanSon, 86 Coeruleotaygetis Forster, 93 Colaenis Hiibner, 19 constantia Cramer, 35, 36 Corades Doubleday, 112, 114 corderoi Dognin, 114 Corderopedaliodes Forster, 114 Cosmosatyrus Felder & Felder, 118, 127 Crebeta Moore, 44, 48, 150 creola Skinner, 145 crisia Geyer, 68, 69 Culapa Moore, 59, 60, 61 cyclopina Staudinger, 115, 118 cyllastros Westwood & Hewitson, 24, 28 Cyllo Boisduval, 36 Cyllogenes Butler, 30, 35, 36 Cyllopsis R. Felder, 90, 91, 93 cymela Cramer, 87, 90, 91, 94, 147 Daedalma Hewitson, 109, 112, 116 Dalapa Moore, 58, 59, 61 Dallacha Moore, 82, 83, 85 damaris Doubleday, 50 Danaidae, 15, 19, 55, 130, 134 Danaus Latreille, 130 t Dasyomma Felder & Felder, 61 Dasyomma Macquart, 61 Dasyophthalma Westwood, 24, 25, 26, 27 Dasyophthalma-series, 25, 27 Davidina Oberthiir, 119, 120, 121, 122, 123 Debis Doubleday & Hewitson, 47 decorata Butler, 117 decorata Felder & Felder, 116 deidamia Eversmann, 48, 50 dendrophilus Trimen, 47, 48 dexamenus Hewitson, 61 THE SATYRIDAE diademoides Moore, 50 Dichothyris Karsch, 58, 62, 63 Didonis Hiibner, 56 digna Marshall, 123 Dingana vanSon, 106, 108 dingana Trimen, 108 dinias Hewitson, 112, 116 Dionana Moore, 47 Dioriste Thieme, 111, 116 Dira Hubner, 106, 107, 108 Dirini, 9) 105 12; 415 71, 725 7385.02" 105-108, 119, 124, 139, 143, 144, 146, 150-151, 152 Dodonidia Butler, 74, 75, 78, 79, 89 doraete Hewitson, 116 dorus Esper, 98 drepana Westwood, 62, 63 Drucina Butler, 110, 111, 116 drusia Cramer, 61 drusillodes Oberthiir, 61 Drusillopsis Fruhstorfer, 61 dryas Scopoli, 123 Dubierebia Muschamp, 98 Dulcedo d’Almeida, 20, 22 dumetorum Oberthiir, 48, 50 duplex Butler, 87 duponcheli Guérin, 63 Dyctis Boisduval, 55, 56, 57 Dynastor Westwood & Hewitson, 25, 26, 27 dynsate Hewitson, 47 Elina Blanchard, 109, 115, 117 Elina-series, 108, 110, 117 Elymnias Hiibner, 54, 56, 57, 132, 150 Elymniina, 3, 4, 38, 54 Elymniinae, 3, 4, 8, 9, 12, 13, 16, 17, 18, 19, 23, 26, 29, 38-63, 64, 67, 72, 135, 138, 139, 140, 144, 146, 147, 149, 150 Elymniini, 10, 11, 12, 13, 17, 18, 23, 26, 39, 40, 41, 42, 52, 54-57, 135, 137, 139, 144, 146, 150, 152 Elymniopsis Fruhstorfer, 54, 55, 56, 57 emma Staudinger, 116 Enodia Hiibner, 40, 41, 43, 46, 47 Enodiinae, 3, 4, 38, 40, 41 Enodiini, 41 t Enope Moore, 49 Enope Walker, 49 LEE DS MILLER enyo Hewitson, 112, 114 Epigea Hubner, 104 Epimede Houlbert, 126 epimenides Ménétriés, 50 Epinephele Hubner, 101, 109 “Epinephila” auctt., 101 “Epinephile” auctt., 101 epiphron Knoch, 105 epistygne Hubner, 103, 105 Erebia Dalman, 74, 78, 80, 102, 104, 105, 108, 114, 142, 145 Erebiinae, 101 Erebiini, 9, 71, 73, 81, 95, 98, 101-105, 108, 119, 142, 144, 146, 147 erebioides Felder & Felder, 115, 118 Erebiola Fereday, 74, 78, 79, 80 £ Erebomorpha Elwes, 87 Erebomorpha Walker, 87 Eretris Thieme, 116 erichtho Butler, 93 Erichthodes Forster, 93 Erites Westwood, 64, 65, 66 EBritinae; 4; 85 9, 12; 116, 17, 19) 39; 64- 66, 67, 72, 135, 139, 140, 144, 146, 148, 149 Eritini, 64-66, 144 Erycinidia Rothschild & Jordan, 75, 76, 80 Eryphanis Boisduval, 25, 26, 27 esaca Westwood & Hewitson, 56 Etcheverrius Herrera, 118 Eteona Westwood, 109, 111, 116 Ethope Moore, 44, 50 Eumenis Hibner, 123 euphemia Westwood & Hewitson, 76, 80 Euploea Fabricius, 55 Euploeamima Holland, 50 Euptychia Hiibner, 90, 91, 93 Euptychiini, 9, 70, 73, 74, 81, 87, 88, 89-95, 98, 99, 140, 141, 142, 144, 145, 146, 147, 148, 149 Euptychoides Forster, 93 euripides Weymer, 117 europa Fabricius, 45, 48 eurydice Linné, 46, 49, 145 eurylochus Hiibner, 24, 27 Eurytelidae, 38, 54 eurytus Fabricius, 94 evadne Cramer, 61 103, MEM. AMER. ENT. SOC., 24 167 | fagi Scopoli, 123 Faunis Hubner, 132 Faunula Felder & Felder, 118 fergana Oberthtir & Houlbert, 126 francisca Stoll, 60, 61 fumata Butler, 95 fuscum Felder & Felder, 60, 61 galathea Linné, 125, 126 £ Gallienia Oberthir, 63 Gareris Moore, 61 “Geirocheilus” Holland, 116 Geitoneura Butler, 78, 79, 80 germainii Felder & Felder, 118 geticus Esper, 98 glacialis Moll, 124 Gnophodes Westwood, 36 Godartiana Forster, 93 Gorgo Hubner, 102, 104 gotama Moore, 63 gracilis Rothschild & Jordan, 76, 80 Gyrocheilus Butler, 109, 110, 111, 11 113, 116 2 “s gyrtone Berg, 117 Haetera Fabricius, 20, 21, 22, 134 Haeternnaenss:4aa/asse 9 LON Tee 2 2113: 1S, 1G, 17, 1G, 102), D3, POs We, isv, 133, 134, 136, 140, 144, 145, 146 Haeterini, 19, 20-22, 133, 144, 145 Halimede Oberthiir & Houlbert, 126 halimede Ménétriés, 126 Hallelesis Condamin, 63 Hamadryopsis Oberthir, 61 “Hanifa” auctt., 47 hanifa Nordmann, 123 Hanipha Moore, 47 Harima Moore, 47 Harjesia Forster, 93 Harsiesis Fruhstorfer, 80 haydenii Edwards, 145 Haywardella Herrera, 116 Haywardina Aczel, 93 t Haywardina Forster, 93 hedemanni R. Felder, 91, 93 helmsii Butler, 79, 80 Hemadera Moore, 85 Henotesia Butler, 63 hercyna Hubner, 36, 37 168 hermes Fabricius, 93 Hermeuptychia Forster, 93 Hermianax Fruhstorfer, 50 Hermias Fruhstorfer, 42, 44, 47 hermione Linné, 123 herse Cramer, 93 hesione Cramer, 62 hesione Sulzer, 94 Hetaerina, 3, 4, 19, 20 Heteronympha Wallengren, 75, 77, 81 Heteropsis Westwood, 58, 62, 63 hewitsonii Doumet, 62, 63 hilda Westwood, 50, 51 himachala Moore, 50 Hipio Hubner, 30, 35, 36 Hipparchia Fabricius, 119, 123 Hipparchiadae, 15, 69, 119 Hipparchioides Butler, 81 hippia Cramer, 87 hobartia Westwood & Hewitson, 80 Homoeonympha Felder & Felder, 109, 118 Houlbertia Oberthiir, 63 huebneri Felder, 123 huebneri Kirby, 85 humilis Felder & Felder, 118 hyagriva Moore, 85 Hyalodia Jordan, 75, 80 hygeia Hewitson, 80 hypaesia Hewitson, 22 “hy peranthus” auctt., 98 hyperantus Linné, 97, 98 hyperbius Linné, 86, 87 hypermnestra Linné, 56, 57 Hypocysta Westwood & Hewitson, 75, 76, 80 Hypocysta-series, 12, 74, 76-77, 78, 80, 81, 149, 150 Hypocystini, 7, 9, 11, 12, 57, 70, 72, 73, 74-81, 102, 105, 108, 110, 119, 124, 139, 140, 141, 144, 146, 149, 150, 151 Hyponephele Muschamp, 101 Idata deLesse, 101 Idiomorphus Chaudoir, 63 t Idiomorphus Doumet, 63 t Idioneura Felder & Felder, 118 Idioneura Selys, 118 THE SATYRIDAE Idioneurula Strand, 110, 113, 115, 118 incerta Hewitson, 51, 53 Indalasa Moore, 61 inga Fruhstorfer, 61 inica Hewitson, 85 Ithomiidae, 9, 12, 131-132, 134 itonia Hewitson, 85 14-15, 16, 19, 130, jalaurida deNicéville, 49 janardana Moore, 61 janira Linné, 101 Jatana Moore, 61 jurtina Linné, 100, 101 jynx Hubner, 57 Kabanda Moore, 61 kalinda Moore, 87 Kallima Doubleday, 19 Kanetisa Moore, 123 Karanasa Moore, 119, 123, 148 Kerrata Moore, 47 Kirinia Moore, 43, 50 Kirrodesa Moore, 47 klugii Guérin, 79, 80 Kolasa Moore, 82, 85 Lachesis Oberthiir & Houlbert, 126 lachesis Hiibner, 126 lais Cramer, 57 lais Fabricius, 57 Lampides Hiibner, 130 Lamprolenis Godman & Salvin, 72, 74, 75, 76, 77, 80, 139 lanaris Butler, 47 languida Butler, 94 Lasiommata Westwood, 50 Lasiophila Felder & Felder, 110, 116 lathionella Westwood, 81 latipicta Fruhstorfer, 50 leaena Hewitson, 118 leda Gerstacker, 87 leda Leech, 126 leda Linné, 35, 36, 136, 150, 152 Ledargia Houlbert, 124, 126 lemur Schrank, 101 leonata Butler, 111, 116 leprea Hewitson, 81 Leptoneura Wallengren, 108 LEE D. MILLER leptoneuroides Felder & Felder, 118 Lepitotes Scudder, 130 Lethe Hiibner, 16, 41, 43, 44, 45, 47, 48 Lethe-series, 42, 43, 47-49, 137 Lethinae, 3, 4, 38, 40, 41 Lethini, 28, 34, 37, 38, 39, 40-51, 52, 54, 57, 4 Wp TES OSs webs, ils}757 alei3s 140, 144, 146, 147, 149, 151, 152 leucoglene Felder & Felder, 118 libye Linné, 94 Libythea Fabricius, 130 Libytheidae, 130 ligea Linné, 103, 104 Limenitinae, 52 lise Hemming, 56, 57 Loesa Moore, 61 Lohana Moore, 85 Lohora Moore, 61 Lopinga Moore, 44, 48, 50 lorquinii Felder & Felder, 46, 49 lowi Doubleday & Hewitson, 50 Loxerebia Watkins, 82, 87 Lycaena Fabricius, 74 Lycaenidae, 130 lycaon Rottemburg, 101 Lycoreinae, 133 Lyela Swinhoe, 95, 96, 98 Lymanopoda Westwood, 109, 118 Lymanopoda-series, 108, 110, 113, 114, 118, 127 lyrnessa Hewitson, 81 macmahoni Swinhoe, 98 madura Horsefield, 66 Magneuptychia Forster, 94 t Magula Fruhstorfer, 44, 49 Magula Scudder, 49 maianeas Hewitson, 63 malsara Moore, 49 malsarida Butler, 61 Manataria Kirby, 10, 18, 34, 36, 37, 38, 41, 146 Mandarinia Leech, 41, 43, 46, 50, 51, 12, 714, 1139, 149 Mandarinia-series, 42, 43, 50 Manerebia Staudinger, 110, 118 Maniola Schrank 2, 99, 100, 101, 109 Maniolidi, 98 Hilit, Ibe MEM. AMER. ENT. SOC., 24 169 Maniolinae, 69, 98 Maniolini, 9, 71, 73, 74, 81, 89, 98- 101, 109, 120, 142, 144, 146, 147 margaretae Elwes, 47 Marica Hubner, 104 marshallii Wood-Mason, 36 Martanda Moore, 58, 59, 61 Mashuna vanSon, 83, 84, 85 mashuna Trimen, 84, 85 masoni Elwes, 49 Masoura Hemming, 58, 63 masoura Hewitson, 59, 63 medus Fabricius, 60, 62 medus Schiffermiller, 104 Medusia Verity, 102, 104 megera Linné, 50 Megeuptychia Forster, 94 Megisto Hubner, 91, 94 Melampias Hibner, 82, 86, 87 Melampias-series, 83, 84, 86-87, 141, Sit, Sy Melanargia Meigen, 4, 11, 16, 124, 125, 126 Melanargiinae, 124 Melanargiini, 5, 9, 10, 71, 73, 105, 119, 124-126, 143, 144, 146, 147 Melania Lamarck, 123 t Melania Sodoffsky, 123 Melanitini, 10, 18, 28, 29, 30, 34-36, 37, 39, 41, 135, 136, 138, 144, 146, 149, Sy Melanitis Fabricius, 16, 30, 35, 36, 150 Melynias Moore, 54, 55, 57 menander Drury, 22 Meneris Westwood, 47 mermeria Cramer, 91, 95 merope Fabricius, 81 Mestra Hiibner, 11, 16 mestra Hewitson, 62 Mimadelias Moore, 54, 57 minerva Fabricius, 49 Minois Hubner, 99, 119, 120, 123 Mintha vanSon, 108 mintha Geyer, 107, 108 mirabilis Staudinger, 118 mixturata Alpheraky, 95, 146 mnasicles Hewitson, 60, 61 mollina Hubner, 91, 92, 93 monachus Blanchard, 117 170 Moneuptychia Forster, 94 Monotrichtus Hampson, 61 monticolens Butler, 118 moorei Felder, 61 Morphidae, 2, 3, 14-15, 16, 19, 23, 131, 132, 134 Morpho Fabricius, 15, 132 Muscopedaliodes Forster, 116 muscosa Thieme, 116 Mycalesini, 11, 12, 13, 33, 39, 40, 52, 54, 57-63, 72, 74, 75, 128, 135, 138, 139, 144, 146, 147, 149, 150, 151, 152- 153 Mycalesis Hiibner, 42, 46, 58, 60, 61, 63, 139 Mycalesis group, 67 mycalesis Felder & Felder, 61 mycalesoides Felder, 94 Mydosama Moore, 60, 61 Mygona Westwood, 116 mynois Hewitson, 61 myops Staudinger, 98 Myrtilus deNicéville, 61 mystes deNicéville, 61 Nadiria Moore, 85 napoleon Westwood & Hewitson, 27 narasingha Moore, 85 nareda Kollar, 85 Narope Westwood & Hewitson, 23, 24, 25, 26, 28 Narope-series, 25, 28 Nasapa Moore, 62 natalii Boisduval, 86 Nebdara Moore, 62 necys Godart, 93 Neita vanSon, 87 neita Wallengren, 87 Nelia Hayward, 109, 117 Nemetis Moore, 49 nemyroides Blanchard, 117 Neocoenyra Butler, 87 Neohipparchia deLesse, 123 Neomaenas Wallengren, 109, 117 Neomaniola Hayward, 117 Neominois Scudder, 119, 120, 121, 123 Neonympha Hibner, 69, 87, 91, 94 Neope Butler, 49 Neorina Westwood, 43, 50, 51 Neorina-series, 37, 39, 40, 42, 43, 50, 52, THE SATYRIDAE 138 Neosatyrus Wallengren, 117 nereis Drury, 21, 22 nesaea Linné, 57 Nesoxenica Waterhouse & Lyell, 77, 81 Ninguta Moore, 49 Nissanga Moore, 62 nitida Godman & Salvin, 76, 80 norna Thunberg, 122, 124 nothis Westwood & Hewitson, 66 Nymphalidae, 52, 130 Nymphaloidea, 7, 8, 19 Nytha Billberg, 123 obsoleta Westwood, 118 Ocalis Westwood, 94 occitanica Esper, 126 Odonata, 131 oedipe Hiibner, 98 oedippus Fabricius, 97, 98 Oeneis Hiibner, 16, 119, 120, 122, 124, 142, 145 Oeneis-series, 73, 143 opalina Butler, 87, 88 opalinus Staudinger, 114 Opoptera Aurivillius, 25, 27 Opsiphanes Westwood & Hewitson, 24, 25, 26, 27 Oreas Hubner, 22 Oreina Westwood, 105 Oreixenica Waterhouse & Lyell, 81 oreseis Hewitson, 63 Oressinoma Westwood, 99, 92, 93, 94 Orinoma Gray, 43, 50 ornata Rothschild & Jordan, 80 oroatis Hewitson, 61 Orsotriaena Wallengren, 40, 57, 58, 59, 60, 62, 139 Orthoptera, 131 Oxeoschistus Butler, 116 120, 121, 122, 124, Pachama Moore, 62 Palaeonympha Butler, 74, 87-89, 146 pales Philippi, 117 Palmaris Herrera, 118 Pampasatyrus Hayward, 117 Pamperis Heimlich, 127-128 pamphanis Westwood & Hewitson, 27 pamphilus Linné, 97, 98 LEE D. MILLER 17a Panarche Thieme, 116 panda Boisduval, 84, 87 Pandima Moore, 83, 85 pandoea Hopffer, 63 Panyapedaliodes Forster, 116 panyasis Hewitson, 116 Papilio Linné, 2 Papilionoidea, 3 paradoxa Mabille, 62, 63 “Parage” auctt., 50 Paralasa Moore, 82, 83, 87 Paralethe vanSon, 9, 43, 44, 47, 48 Paramecera Butler, 90, 93, 94 Parantirrhoea Wood-Mason, 30, 34, 35, 36, 138 Parapedaliodes Forster, 116 “Pararga” auctt., 50 Pararge Hiibner, 42, 43, 45, 48, 50 Pararge-series, 39, 42, 43, 44, 46, 49-50, 137, 139, 147 Parataygetis Forster, 94 Paratisiphone Watkins, 81 Parce Oberthiir & Houlbert, 126 parce Staudinger, 126 parepa Hewitson, 116 Pareuptychia Forster, 94 parmenio Boeber, 86, 87 parmeno Westwood, 36 Paroeneis Moore, 123 Paryphthimoides Forster, 94 Pasiphana deLesse, 101 passandava Ward, 63 Patala Moore, 49 patnia Moore, 62 patrobas Hewitson, 113, 116, 147 Pedaliodes Butler, 111, 113, 116 pegala Fabricius, 100, 101 pelopea Klug, 123 penanga Westwood & Hewitson, 56 penelea Cramer, 94 penelope Fabricius, 91, 93 Penetes Westwood & Hewitson, 25, 27 Penrosada Brown, 113, 118 Percnodaimon Butler, 74, 78, 79, 80, 81 periboea Fabricius, 128 periboea Godman & Salvin, 93 Periplysia Gerstacker, 83, 87 phaedra Linné, 123 phanias Hewitson, 116 MEM. AMER. ENT. SOC., 24 phantoma Fassl, 22 phares Godart, 94 t Pharia Fruhstorfer, 69 Pharia Gray, 69 Pharneuptychia Forster, 94 t phegea Fabricius, 57 Pherepedaliodes Forster, 116 pheretiades Smith & Kirby, 116 phidia Linné, 123 Philareta Moore, 123 philomela Linné, 84, 85 pholoe Staudinger, 112, 114 Phorcis Hiibner, 102, 103, 104, 105 phoronea Doubleday, 117 t Phryne Herrich-Schaffer, 98 Phryne Meigen, 98 phryne Pallas, 97, 98 Physcaeneura Wallengren, 83, 84, 87 physcoa Hewitson, 116 Physcon deNicéville, 63 Physcopedaliodes Forster, 116 piera Linné, 21, 22 Pierella Herrich-Schaffer, 4, 20, 21, 22, 134 Pierellinae, 3, 4, 19, 20 Pieridopsis Rothschild & Jordan, 75, 76, 78, 80 pigmentaria Karsch, 47, 48 pignerator Butler, 111, 114 pimplea Erichson, 51, 52, 54 Pindis R. Felder, 90, 94 pireta Cramer, 21, 22 Placilla Moore, 49 Platypthima Rothschild & Jordan, 75, 77, 80 pluto Fereday, 79, 81 poaoeneis Heimlich, 127 poesia Hewitson, 111, 116 polita Hewitson, 22 poltys Prittwitz, 94 Polymastus Thieme, 116 polyxo Godman & Salvin, 117 portlandia Fabricius, 46, 47, 145 Posteuptychia Forster, 94 Posttaygetis Forster, 94 Praefaunula Forster, 94 Praepedaliodes Forster, 116 Praepronophila Forster, 116 pratorum Oberthtr, 87 172 Precis Hubner, 130 pringlei Sharpe, 47 Proboscis Thieme, 113, 116 pronoe Esper, 105 Pronophila Doubleday & Hewitson, 109, TUS eas ali) Pronophila-series, 10, 110, 114-117 pronophila Felder & Felder, 116-117 Pronophilinae, 3, 4, 108 Pronophilini, 7, 9, 10, 71, 73, 102, 105, 108-118, 119, 127, 140, 143, 144, 145, 146, 147, 148 propylea Hewitson, 113, 116 Pseudeuptychia Forster, 94 Pseudochazara deLesse, 123 Pseudodebis Forster, 94 Pseudohaetera Brown, 20, 22 t Pseudomaniola Rober, 114 Pseudomaniola Weymer, 114, 117 Pseudonympha Wallengren, 82, 87 Pseudosteroma Weymer, 116-117 Pseudotergumia Agenjo, 123 Psyche Hiibner, 126 Ptychandra Felder & Felder, 34, 39, 41, 43, 44, 45, 46, 49, 138 puerta Hewitson, 116 pulchra Mathew, 80 pumilus Felder, 123 Punapedaliodes Forster, 117 pusilla Felder & Felder, 118 Putlia Moore, 47 Pyronia Hubner, 100, 101 Quilaphoethosus Herrera, 117 Ragadia Westwood, 9, 67, 68, 69 Ragadiina, 3, 4, 67 Ragadiinae, 3, 4, 8, 9, 12, 13, 15, 16, 17, 18, 39, 67-69, 72, 139, 141, 144, 146, 147, 148, 149 Ragadiini, 67-69, 144 Rangbia Moore, 45, 49 Rareuptychia Forster, 94 regalis Leech, 50, 51, 57, 147 Rhaphicera Butler, 45, 46, 50 Rhaphiceropsis Sharpe, 47 ridingsii Edwards, 121, 123, 145, 148 ruscinonensis Oberthiir & Houlbert, 126 rusina Godart, 24, 27 THE SATYRIDAE Sabatoga Staudinger, 118 Sadarga Moore, 63 safitza Hewitson, 61 sagitta Leech, 51, 53 saitis Hewitson, 77, 80 sallei Westwood & Hewitson, 27 Samanta Moore, 10, 18, 34, 39, 41, 49, 138 sambulos Hewitson, 62, 63 samio Doubleday & Hewitson, 47 samius Westwood, 110, 115, 118 Samundra Moore, 63 sanatana Moore, 61 Sarromia Westwood, 109, 118 Satoa Moore, 58, 63 satricus Westwood & Hewitson, 45, 50 saturnus Butler, 93 Satyrides, 2, 6, 15, 69, 119 Satyrina, 3, 4 satyrina Bates, 94 satyrina Butler, 45, 49 Satyrinae, 3, 4, 6, 7, 8, 9, 10, 12, 13, 16, 17, 18, 39, 41, 42, 57, 64, 67, 69- 126, 136, 139, 140-142, 144, 145, 146, 147, 148, 149, 150, 151, 152 Satyrini, 6, 7, 9, 10, 71, 73, 99, 105, 106, 108, 110, 119-124, 142, 143, 144, 145, 146, 147, 148, 149 Satyrini-section, 12, 73, 74, 75, 101, 105, 108, 110, 140, 141, 143, 145, 150 Satyrodes Scudder, 43, 44, 46, 49 Satyrotaygetis Forster, 94 Satyrus Latreille, 2, 119, 121, 123 Satyrus-series, 10, 120, 122, 123, 148 scanda Kollar, 86, 87 scanda Moore, 45, 49 schrenkii Ménétriés, 49 Selenophanes Staudinger, 25, 27 servilaea Wallengren, 117 sesara Hewitson, 84, 85 Setodocis Billberg, 128 Sevanda Moore, 63 Sicca Verity, 98 sicelis Hewitson, 47 sidonis Hewitson, 49 sihala Moore, 47 similis Butler, 95 Simplica Verity, 105 Sinarista Weymer, 29, 30, 31, 32 143, LEE D. MILLER Sinchula Moore, 49 t Smithia Mabille, 63 Smithia Milne-Edwards & Haime, 63 sophorae Linné, 24, 26, 27 soter Butler, 94 Spinantenna Hayward, 110, 114, 115, 117 Splendeuptychia Forster, 94 squamistriga R. Felder, 94 statilinus Hufnagel, 123 stelligera Butler, 118 Steremnia Thieme, 117 Steroma Westwood, 109, 112, 113, 117 Steromapedaliodes Forster, 117 Stibomorpha Butler, 117 Strabena Mabille, 82, 83, 87 Stuardosatyrus Herrera & Etcheverry, 117 Stygionympha vanSon, 87 stygne Ochsenheimer, 104 Stygnolepis Strand, 118 + Stygnus Felder & Felder, 118 Stygnus Perty, 118 sudra Felder, 61 sura Westwood, 49 suradeva Moore, 36 Suralaya Moore, 63 syme Hubner, 27 Syngea Hiibner, 105 114, Taenaris Hiibner, 19, 132 tagala Felder, 62 tamatavae Boisduval, 87 Tanaoptera Billberg, 48 Tansima Moore, 42, 45, 49 Tarsocera Butler, 106, 108 Tatinga Moore, 50 tauropolis Westwood, 116 Taygetina Forster, 94 Taygetis Hiibner, 90, 91, 92, 95 Telinga Moore, 63 Tellervini, 132 tena Hewitson, 114 tenuisquamosa Joicey & Talbot, 80 Tetraphlebia Felder & Felder, 118 thalia Leech, 69 t thelebe Doubleday, 116 thelebe Doubleday & Hewitson, 111, 116 Theope Doubleday & Hewitson, 50 MEM. AMER. ENT. SOC., 24 173 t Theope Moore, 50 thibetanus Oberthir, 50 Thiemeia Weymer, 117 thione Berg, 117 Thympia Moore, 82, 85 tircis Stoll, 98 Tisiphone Hubner, 81 tisiphone Boisduval, 116 tithone Hiibner, 101 tithonus Linné, 100, Tithoreini, 132 tomasia Butler, 31 Torynesis Butler, 10, 106, 107, 108 Triariia Verity, 105 triarius dePrunner, 105 101 | tricordata Hewitson, 116 trimacula Leech, 47 Triphysa Zeller, 95, 96, 97, 98 tristigmata Elwes, 47 tristis Guérin, 115, 117 Triteleuta Strand, 30, 31 Trophonina Rober, 118 Truncaecfalcia Verity, 105 | tulbaghia Linné, 47, 48 typhla Westwood, 92, 93, 94 valentina Cramer, 94 Vanessa Fabricius, 130 vanessoides Blanchard, 115, 117 Vareuptychia Forster, 95 vasudeva Moore, 57 verma Kollar, 47 vigilans Trimen, 87 Virapa Moore, 63 virgilia Cramer, 95 virgo Rothschild & Jordan, 76, 80 viridicans Weymer, 95 | vitellia Cramer, 56 waltoni Elwes, 87 | wardii Butler, Callyphthima, 86 wardii Butler, Henotesia, 63 | weidemeyeri Edwards, 52 Weymerana Forster, 95 williamsianus Butler, 117 | Xenica Westwood, 75, 78, 79, 81 Xenica-series, 74, 76, 77, 78, 80-81, 108 150 5) 174 THE SATYRIDAE Xeniconympha Novicky, 81 Ypthimini-section, 12, 73, 74, 75, 81, 90, xicaque, Reakirt, 93, 94 95, 99, 102, 140, 141, 145 Xois Hewitson, 82, 84, 85, 150 Ypthimomorpha vanSon, 85 Ypthimorpha Overlaet, 85 yama Moore, 49 “Yphthima”’ auctt., 85 Zabirnia Hewitson, 118 yphthima Felder, 95 zangris Hiibner, 114 Yphthimoides Forster, 95 Zethera C. Felder, 51, 52, 53, 54 Ypthima Hiibner, 82, 84, 85 Zetherini, 10, 39, 40, 41, 52-54, 135, 139, Ypthima group, 67 144, 146, 147 Ypthima-series, 82, 83, 84, 141 zigomala Hewitson, 118 Ypthimini, 9, 10, 70, 73, 74, 75, 81-87, | Zipaetis Hewitson, 75, 77, 78, 80 88, 89, 95, 99, 101, 102, 141, 144, 145, | Zischkaia Forster, 95 146, 147, 149, 150, 151, 152 Zophoessa Westwood, 43, 49 MEMOIRS OF THE AMERICAN ENTCMOLOGICAL SOCIETY An irregular serial of monographic papers by students of authority in their re- spective subjects. t a 4. The Cresson Types of Hymenoptera. Ezra T. Cresson. 1916. 46.00. A Venationai Study of the Suborder Zygoptera (Odonata), with Keys for the Identification of Genera. Philip A. Munz. 1919, $2.00, The Blattidae of Panama. Morgan Hebard. 1920. $3.00. The Types of Hymenoptera in the Academy of Natural Sciences of Philadelphia other than those of Ezra T. Cresson. Ezra T. Cresson. 1928. $2.00. Revision of the Rhipiphoridae of North and Central America (Coleoptera). Ezekiel Rivnay. 1929. $2.00. A Revision of the Dipterous Family Rhagionidae (Leptidae) in the United States and Canada. Mortimer D. Leonard. 1930. $4.50. The Eumastacinae of Southern Mexico and Central America. James A. G. Rehn and John W. H. Rehn. 1934. $2.50. The Generic Names of the Sphecoid Wasps and their type species. V. S. L. Pate. i937. $2.50. A Revision of the North American species belonging to the genus Pegomyia. H. C. Huckett. 1941, $3.00. Catalogue and reclassification of the Nearctic Ichneumonidae. Henry K. Townes, Jr. 1944. $15.00. ao Biology and Identification of Trypetid Larvae. Venia Tarris Phillips. 1946. 5.00. reas of North America (Microlepidoptera). Annette F. Braun. 1948. 50. Classification of the Blattaria as Indicated by their Wings (Orthoptera). John W. H. Rehn. 1951. $5.00. The Neotropical Species of the “Subgenus Aeschna” sensu Selysii 1883 (Odo- nata). Philip P. Calvert. 1956. $10.00. A Taxonomic Study of the North American Licinini with Notes on the Old World Species of the Genus Diplocheila Bruilé (Coleoptera). George E. Ball. 1959. $10.00. A Taxonomic Study of the Milliped Family Spirobclidae (Diplopoda: Spiro- bolida). William T. Keeton. 1960. $5.50. The Genus Bucculatrix in America North of Mexico (Microlepidoptera). Ann- ette F. Braun. 1963. $8.50. The Butterflies of Liberia. Richard M. Fox, Arthur W. Lindsey, Jr., Harry K. Clench, and Lee D. Miller. 1965, $12.50. A Revision of the Mexican and Central American Spider Wasps of the Sub- family Pompilinae (Hymenoptera: Pompilidae). Howard E. Evans. 1966. $12.50. . A Taxonomic and Zoogeographic Survey of the Scarabaeinae of the Antilles (Coleoptera: Scarabaeidae). Eric G. Matthews. 1966. $4.00. A Monograph of the Ithomiidae (Lepidoptera) Part III. The Tribe Mechanitini Fox. Richard M. Fox. 1967. $9.00. . A List of New North American Spiders, 1940-1966. Beatrice R. Vogel. 1967. $9.00. The Higher Classification, Phylogeny and Zoogeography of the Satyridae (Lepi- doptera), Lee D. Miller. 1968. $7.00. In making inquiries relative to publications, address: THE AMERICAN ENTOMOLOGICAL SOCIETY 1900 Race Street, Philadelphia, Pa. 19103