Systematic (1977) 2, 161-197

Geographical patterns of evolution in Neotropid : differentiation of the of MeZimea and ( ,Ithomiinae)”

KEITH S. BROWN, Jr Departamento de Zoologia, Instituto de Biologia, Universidade Estadual de Campinas, Sao Paulo,

ABSTRACT. Biosystematic analysis incorporating abundant new field data from many parts of the Neotropics has led to an ordered revision of the mimetic ithomiine genera and Mechanitis. The various polytypic species of these genera probably served as prime movers for the differentiation of other mimetic in Quaternary forest refuges. The revisions are presented in the form of supplements to the works of fichard M. Fox on these genera, with analyses based on his divisions. Seven species (or monophyletic species-groups) and sixty- three well-differentiated geographic subspecies (six of these described here for the first time) are recognized in Melinaea. Specimens are illustrated which demonstrate intergradation between refuge-derived subspecies. Five species and fifty-two differentiated subspecies are recognized in Mechanitis, whose members are more abundant and gregarious, more plastic, and apparently more vagile than those of Melinaea, resulting in fewer clear-cut mimetic associations, more exten- sive blurring of differentiation patterns, and apparently fewer incipient biological species in this genus than in Melinaea.

Introduction very difficult groups without ever having studied any of the species alive in nature. The mimetic tropical species in the Unfortunately, it is not always practical to genera Melinaea and Mechanitis (Nymphalidae, effect a reliable analysis of the relationships Ithomiinae) have a confused histoi y of syste- between named forms in these genera without matic arrangements, due primarily to their extensive field work in many different apparent plasticity of colour-pattern and regions. The plasticity of the patterns observed monotony of morphology (d’Almeida, 1951; in these species is probably in part due to Fox, 1949). The speciation patterns in these extensive recent recombination of genes two genera were analysed by Forbes (1924, arising from past differentiations in isolated 1927, 1948), d’Almeida (1951) and Fox forest refuges (Brown et al., 1974; Brown, (1960, 1965, 1967); all three studied a limited 1976a). Large samples must thus be obtained number of specimens. Fox’s genius is amply from a wide variety of localities before any demonstrated by the fact that he succeeded in consistent differentiation patterns become reaching a profound understanding of these apparent. Fortunately, most of the members * Contribution number 12 from the Prograrna de of the genera are common Mullerian models Ecologia, Instituto de Biologia, Universidade Estadual for rings. Thus, it is possible to de Campinas. obtain long series in many areas, showing the Correspondence: Dr Keith S. Brown, Jr, Departa- mento de Zoologia, Instituto de Biologia, Universi- limits of local polymorphic variation and dade Estadual de Campinas, C.P. 1170, Campinas, intergradation between named taxa; and at Sao Paulo 13,100, Brazil. least a few specimens may be captured almost

161 162 Keith S. Brown. Jr anywhere, helping to suggest the levels of gene in their systematics. Melinaea and Mecharzitis flow between imperfectly isolated geographical species are notable for highly localized occur- races. rence and dramatic fluctuations in population As these widespread and common species levels. Many populations of Melinaea seem to have recently been implicated as prime movers be absent during parts of the year, but reach in the differentiation of other mimetic Lepi- high levels under certain select conditions. In doptera (Brown & Benson, 1974; see also a 1 ha ‘ithomiine pocket’ recently studied in Kaye, 1907), it is especially important that southern Brazil (Brown & Vasconcellos Neto, there exist a complete and detailed under- 1976), mark-recapture experiments indicated standing of their geographical variation, in- that the population level of Mechanitis polym- corporating as much data as possible from the nia casabranca varied over several powers of field and from museum collections. The ten in a period of but 3 months. In the same present analysis attempts to bring up to date study it was shown that this species can move the systematic ordering of the groups presen- rapidly over reasonable distances (at least ted by Fox (1 960, 1965, 1967), in the light of 2 km). Many similar investigations support a recent field data from many areas, museum picture of general demographic instability specimens not examined by Fox, analyses (Brown & Benson, 1974), and reasonably made by d’Almeida, and a modern under- great dispersion ability between sharply standing of the bases for differentiation physically bounded populations, for the more in the Neotropics. It is not claimed that this strongly coloured, mimetic ithomiines. Thus, represents a final taxonomic revision, as much it is not surprising that many polymorphic field, insectary, genetic, cytological, physio- populations are known in these genera, with logical, and biochemical work must still be traits appearing in local colonies which undertaken before a complete and definitive apparently have immigrated from regions as comprehension of these groups can be much as thousands of kilometres away. The achieved. Rather, the interpretations are pre- narrow ecological tolerances of ithomiines sented in the light of the patterns discovered have undoubtedly contributed to rapid dif- in the previous analyses of probable Quaternary ferentiation during dryer periods, when the refuges in the Neotropics (Brown et al., 1974; colonies would be widely separated by large Brown, 1976a), which are then used to and essentially impassable areas of dryer vege- suggest solutions for some especially difficult tation, and also to the maintenance of these cases. differentiations in the present day. Moreover, A discussion of the philosophy used in sub- the dispersal abilities of many of the mimetic species designation in relatively sedentary species, especially evident at dawn and dusk polytypic continental species is presented in and on cloudy or very humid days, would the previous paper in this series (Brown, permit a mixing of complex colour-pattern 1976a). That paper also includes a map of elements of the evolved races with consequent the probable Neotropical refuges operative in production of confusingly variable local the last Quaternary dry cycle (and in some populations. These factors are probably the cases, probably still operative today), in the source of much of the difficulty in the syste- differentiation of heliconians and ithomiines; matics of these groups. This problem will be a somewhat revised map (incorporating exten- examined more carefully in some of the cases sive data collected in 1975-76) accompanies discussed in the following sections. this paper. Methodology for species analysis in difficult mimetic groups has been presented Differentiation of the species of Melirzaea with relation to the silvaniform (Brown, 1976b). The following analysis of species and races is The demography and ethology of the based on the papers of Fox (1960, 1965), to Ithomiinae have been discussed by many which this constitutes a supplementary revi- authors (see especially Kaye, 1907; Collenette sion. Names within the section headings refer & Talbot, 1928; Fox, 1967; Gilbert, 1969; to the separate species recognized by Fox, in Brown & Benson, 1974;Brown& Vasconcellos the order established by him, except that the Neto, 1976), and are probably very important first name represents the oldest, to be applied Differentiation of Melinaea and Mechanitis 163 to the species as redefined in this analysis. The comma Forbes, romualdo Fox, lucifer Bates, papers of Fox should be read in parallel with lutzi Fox, and eryx d’Almeida is represented the present discussion, as here only important by so few specimens in toto (only comma and new information will be specifically men- Zucifer are moderately frequent in collections) tioned, and only relevant specimens illustrated that it is not strange that the assemblage was (many additional Melinaea have been figured regarded as three separate species by Fox. The by Brown & Benson, 1974). A complete list Andean subspecies (comma and romualdo) of the new ordering of the taxa, with refer- lack the characteristic series of submarginal ences to Fox’s revisions, is given in the Appen- spots present in all the others. The west dix. Amazonian (lucifer) and east Amazonian The species are presented here in proposed (mnasias) types were not known to intergrade; evolutionary order, though the present data Fox remarked that ‘on distributional grounds on this sequence are admittedly sparse. The there would be little objection to combining known entities are united into seven groups, the two as a single species, but the pattern in each of which appears, from present inforrna- lucifer is strongly modified’ (1 960 : 165). Re- tion, to represent a single polytypic species or cently discovered specimens, however, which monophyletic species-group (insectary test include three new subspecies and three impor- crosses have not yet been possible inMelinaea). tant transitional elements, permit a fuller There are four widespread, well-differentiated picture to be drawn of this very widespread Amazonian species; one of these (mnasias) is but rarely encountered, presumably relict very rare and little known, but well integrated, Melinaea. In size and colour-pattern, mnasias and includes a splinter subspecies in southern looks more like a member of the genus Hypo- Brazil (thera). Each of the other three species thyris, where indeed some early authors placed (maenius, marsueus and menophilus) represents it. It can be separated from other Melinaea, a tentative union of a lowland complex with a and from Hypothyris as well, by the shape of pair of premontane subspecies; with maenius the forewing comma-mark which extends is also placed a recently discovered isolate basad from the margin along vein Cul. It is from the Guianas (mediatrix). There is also regarded as a proto-Melinaea, well separated one poorly-differentiated Amazon-Guiana from all other members of the genus, though species (mneme). In the two widespread some important colour-pattern affinities with species which enter (ethra the Melinaea ethra complex can be seen and and ludovica), the northern and southern are mentioned below. populations have still imperfectly defined The south Brazilian subspecies thera (Fig. continuity through the associated Amazonian 1) has been captured in southern Espirito subspecies. Santo (Negro and the ltabapoana area) and Recent karyological work (Brown, Emmel western Sao Paulo (Aracatuba). Only six & Suomalainen, in preparation) suggests that specimens are known to this author, who has reproductive barriers may be found to exist never seen the subspecies in 10 years’ inten- between the lowland and premontane sub- sive field work in southern Brazil, including a units of marsaeus as defined here, and number of trips directed specifically at re- between the northern, Amazonian, and locating it as well as the sympatric and similarly southern subunits of ethra and ludovica. How- ‘lost’ ithomiine Hyafyrzs fiametta (Hewitson j. ever, the evolutionary unity of these com- Its close resemblance to the abundant and co- plexes is affirmed here, and in the present occurring Hypothyris ninonia daeta (Boisdu- work they are treated as species in the phenetic Val) may cause it to be overlooked by collec- and phyletic, if perhaps not the biological tors. No transitions between thera and adjacent sense. subspecies are known, but its affinities with mnasias are evident. The strange disjunct distri- (1) Melinaea mnasias (Hewitson) (including bution and extreme rarity of thera today M.comma Forbes and M.lucifer Bates) strongly suggest that it differentiated in the The polytypic species composed of the Rio de Janeiro refuge in the past (probably known taxa mnasias Hewitson, thera Felder & not in the most recent dry cycle, but in a pre- Felder, terra Haensch, eratosthenes Hall, vious one), and has been displaced from the 164 Keith S. Brown, Jr area in modern times by more aggressive hindwing much like that of comma and a fore- species, occurring locally only in peripheral wing only differing from that of this Andean habitats. Similar disjunct distributions are subspecies by the retention of the Amazon known in other rare and/or relict Lepidoptera Basin submarginal spot series, is in the Museu (Fox, 1949; Brown & Benson, 1975), and Nacional from ‘San Felippe’ = EirunepC on may be useful to define refuge limits. A corre- the middle Jurua (Fig. 9). The transitional sponding race, possibly derived from the series terminates with comma (Fig. lo), prob- Bahia refuge farther north on the Brazilian ably derived in the Chanchamayo refuge. coast, has yet to be discovered if indeed it Three new subspecies of mnasias have now exists at all. been discovered occupying the northwest, The east Amazonian m. mnasias (Fig. 2) has north-central, and south-central peripheries of turned up fairly frequently in recent collec- the known range of the species, and are des- tions from Para and western Maranhao, and is cribed here. probably a product of the BelCm refuge. On the north coast of occurs a somewhat modified, less yellow-transparent Melinaea mnasias abitagua subspn. phenotype, which possibly differentiated in the Roraima refuge and bears the subspecific Female (Fig. 11) name tecra. The highlands in the interior of FW 40 mm. Similar to M.m.comma (Fig. Amapa and the Guianas, and south-central lo), but possessing a much narrower yellow , are inhabited by the mimetic postmedian band and united discocellular phenotype described as eratosthenes (Fig. 3), spots on the forewing as in lucifer and lutzi, with a black hindwing typical of many species reduced black submarginal elements on the associated with the Manaus/Guiana and hindwing, and a darker ground-colour. Ventuari refuges. The very few known speci- mens of recta and eratosthenes suggest that Male they may be partly sympatric today in western , though they probably originally Unknown, but probably identical in colour- differentiated in allopatry. pattern. Intergradation southwestward from mnasias Holotype 9, : Oriente (Napo- to the last of the three new subspecies des- Pastaza), Km 43 of the Puyo-Nap0 road, near cribed below is evident in a recta-like specimen Santa Clara (lo 16‘S, 77’52’W). 600m, from Itaituba, on the Rio Tapajos, in the col- 9.xii.71 (K. S. Brown, Jr), donated to the lection of R. Diringshofen of Sao Paulo (Fig. Museu Nacional, Rio de Janeiro. 4); this may eventually merit a name as a This subspecies almost certainly evolved in Tapajos-derived subspecies. Westward, a clear the Abitagua refuge, though it may eventually intergrade between the characters of these be found further east in the Putumayo and two and lucifer (which probably evolved in Nap0 areas also. the Loreto refuge) can be seen in a specimen from Tefe, deposited in the Museu Nacional in Rio de Janeiro (Fig. 5), which may also merit Melinaea mnasias neblinae subsp.n. a name if and when a series becomes available. Male (Fig. 12) Transition between lucifer (Fig. 6) and the north Peruvian lutzi (Ucayali refuge?) can be FW 38 mm. Similar to M.m.tecta but more seen in a specimen from southeast of Benjamin heavily marked, the black spots on the FW Constant in western Brazil (Fig. 7, from the larger and the hindwing median and marginal Rio Javari). While eryx, from the Alto Rio black areas broad, barely separated by a nar- Jurua farther south, still preserves a dark hind- row orange stripe. Single intervenal yellow wing (Fig. 8), the forewing pattern is much submarginal dots prominent on forewing and modified toward that of romualdo, derived in on hindwing to space Cul-Cu2. Yellow band the Inambari refuge; eryx may be a locally on forewing not extending discally into cell or derived race, or a mere transitional form. A space Cpl-Cu2. Ventral surface essentially much more interesting specimen, with a dorsal identical in colour-pattern to dorsal. Differentiation of Melinaea and Mechanitis 165

FIGS. 1-13. Melinaea mnasias. All 0.9Xlife size, black, yellow and orange. (1) M.m.thera Felder & Felder, male, dorsal, Arafatuba, Sao Paulo (Museu Nacional, Rio de Janeiro,no. 13/850). (2) M.m.mnasias (Hewitson), male, dorsal, Km 64 Estrada Para-Maranhiio, Mun. Ourhm, Para, 17.ifi.64 (MN-Rio, 0. Mielke leg.). (3) M.m.erQfosfhenesHall, male, dorsal, Serra do Navio (ICOMI), Amapi, Brazil, 7 .iii.63 (MN-Rio, Roppa & Mielke leg.). (4) M.m.subsp., male, dorsal, Itaituba, Rio Tapajds, Par& i.62 (R. Diringshofen). (5) M.m.(mnusius X lucifer), male, dorsal, Tef6, Amazonas, Brazil (MN-Rio, no. 16/624, coll. Arp - ‘flavosignata hicetas’). (6) M.m.lucifer Bates, male, dorsal, Benjamin Constant, Amazonas, V111/947 (MN-Rio, Parko leg.). (7) M.m.(lucifer X lutzi Fox), male, dorsal, Rio Javari above Benjamin Constant, Brazil/ border, 11-16.ix .40 (MN-Rio, Parko leg.). (8) M.m.eryx d’Almeida, holotype male, dorsal, Alto Rio Jurui, Acre (MN-Rio, no. 14/001). (9) M.m.(lucifer X comma), male, dorsal, ‘San Felippe, Acre’ = Eirunepi, Amazonas, middle Rio Jurui (MN-Rio, no. 13/767 - ‘M.hiCefas - d’Almeida det.’). (MI) M.m.comma Forbes, female, dorsal (right) and ventral (left), La Merced, Chanchamayo, Peru, v.63 (KB, L. & G. Harris leg.). (1 1) M.m.QbitQgUQnov., holotype female, dorsal (right) and ventral (left) (left side has a large tear in forewing cell M3-Cul), near Santa Clara, Km 43 of Puyo-Napo Road, east Ecuador, 9.xii.71 (MN-Rio, KB leg.). (12) M.m.neblinae nov., holotype male, dorsal, Cerro Neblina, norte Brasil, 1500m, 25.iv.64 (Facultad de Agronomia, Maracay, Venezuela, J. & B. Bechyne leg.). (1 3) M.m.rondonia nov., holotype male, dorsal (left) and ventral (right), near Riozinho, Rondhnia, 7.vu.72 (MN-Rio, KBleg.). 166 Keith S. Brown, Jr

Female which it was captured, along with twenty Unknown, but probably identical in colour- other species of Ithomiinae, have since been pattern. destroyed. Holotype d, BRAZIL: Amazonas, Pic0 Further races of mnasias may be predicted Neblina, 1500 m (as ‘Cerro Neblina’) (0’ 46’ N, to occur in areas marginal to the Putumayo, 66’04’W), 25.iv.64 (J. & B. Bechyne),Facul- Napo, and Huallaga refuges, and some or all of tad de Agronomia, U.C.V., Maracay, Vene- those mentioned above may prove to be good zuela. subspecies rather than transitional forms, asso- Paratypes. Oned, same data as holotype, ciated with the Tapajos, TefC, and Madeira and one d, same data except 24.iv.64, in the refuges. The species seems to be everywhere same collection. mimetic and very rare, and though it obviously This subspecies probably evolved in the has a very extensive distribution, many gaps Imeri refuge, and may be looked for elsewhere remain to be filled. Most of the known speci- in northwestern Brazil. The high altitude of mens have been captured in localities peri- the capture is exceptional, possibly represent- pheral to supposed refuges, flying with ing an altitudinal variation of the marginal- transitional or hybridized populations of more ization phenomenon general in this species. common species, which strongly suggests that mnasias has been subjected to strong, prob- ably competitive dislocatory phenomena by its relatives, in many areas (see Fox, 1949 Melinaea mnasias rondonia subspn. and Brown & Benson, 1975).

Male (Fig. 13) (2) Melinaea ethra (Latreille) (including FW 36-38 mm. Similar to mmnasias but Melinaea lilis Doubleday, M.scylax Salvin, M. somewhat larger, with the forewing strongly mnemopsis Berg, M.maelus (Hewitson), orange, possessing a narrower yellow post- M.rnaeonis Hewitson, and M.maenius (Hewit- median band, a somewhat reduced comma- son) (sensu Fox, part)). mark, and much anteapical orange coloration The three allopatric species lilis, scylax, ventrally. Hindwing median and marginal and ethra were recognized by Fox ( 1960: 1 18) black areas wider; no yellow on hindwing. A as closely related, forming a superspecies within full series of single intervenal submarginal Melinaea. As all of the pattern characteristics yellow spots present on both wings, dorsally of scylax (Fig. 14), except for its character- and ventrally. istic unmarked hindwing, are represented in the adjacent and allopatric subspecies related to Iilis, imitata Bates and parallelis Butler Female (these two intergrade in west-central ; Unknown, but probably identical in colour- an excellent transitional series from Cerro pattern. Campana exists in the collection of Gordon Holotype d, BRAZIL: Rondhia, near Small, Fig. 15); and as the three form a Central Riozinho, in woods north of Km489 of the American triad assignable to differentiation Porto Velho-Vilhena Highway (1 1” 31’S, 61” from a single stock, under mimetic pressure, 20’W), 300 m, 7.vii.72 (K. S. Brown, Jr), in the Quaternary refuges designated Chiriqui, donated to the Museu Nacional, Rio de Guatemala, and DariCn, respectively (the J aneiro . Mexican ftavicans Hoffmann, Fig. 16, is Paratypes. One d, same data as holotype, apparently a north-peripheral isolate), it seems retained in the collection of the author; one best to regard scylax as a subspecies of the d, same locality, 5 .vii.72, in the collection of widespread and polytypic lilis-group, in spite W. W. Benson (Campinas, Sao Paulo). of the morphological differences reported This subspecies probably evolved in the (Fox, 1960). Rond6nia refuge, on whose southeastern The pattern elements in the southern margins it was captured, and may be expected M.ethra (Fig. 17) are considerably different to appear rarely in various parts of the territory from those of lilis-group races. This butterfly today. The scrubby woods (‘cerradao’) in has recently been found far into the interior Differentiation of Melinaea and Mechanitis 167 of southern Brazil (upper Rios Sao Francisco 1965); such mislabelling with generalized and Grande) and may be very widespread if localities was quite common in early material rather sparse over extra-Amazonian Brazil. from South America, and often introduced In view of the wide geographic separation great confusion into nomenclature until the between ethra and lilis (all of the Amazon respective taxa could be relocated in their Basin, at least 3000 km), it becomes necessary proper ranges. to identify associated races in the intervening Melinaea lilis lirnitata Hall (Fig. 20) was area in order to propose that the two may be correctly synocymized by Fox (1960) with regarded as conspecific. The logical candidates sola Kaye from Trinidad and represents an are the members of the M.maelus complex, endemic Sucre/Trinidad subspecies. The types which are closely related and allopatric to, of limitata are from the Cumana area in and possess many pattern elements in extreme northeastern Venezuela, and recent common with lilis and ethra. specimens are available from Caripe. M.I. Colour-pattern elements of rnaelus and lateapicalis Hall (Fig. 2 1) is entirely different ethra as well as of the lilis-group races are in appearance and represents a good geogra- conspicuously present on both fore- and hind- phically defined subspecies from northwestern wings of M.mnemopsis from northern Venezuela. It has not been relocated in recent (Fig. 18), which should thus form part of this collecting, which has produced an ezra (des- polytypic species also. cribed from the Santa Marta range in extreme Melinaea dodona Hopffer was named from northern ) from the Sierra Perija an apparently unique specimen from ‘Bolivia’. (Fig. 22). Moreover a single transitional speci- Its description strongly suggests that it is a men (Fig. 23) between lilis (northcentral member of the lilis-group, where it was placed Venezuela, Rancho Grande refuge) and late- by Fox (1960). Indeed, its colour-pattern sug- apicalis has been taken in Barinitas, where the gests that it should occur in Colombia and not probably conspecific zamora (see below) also Bolivia, as it combines characters of rnessatis occurs very rarely. More specimens are needed (Hewitson) with those of erica Bargmann or from this region to define the transition ezra Fox, both of which should intergrade at between Venezuelan races of ethra. Pure the peripheries of the central Colombian lateapicalis should be sought in the southern valleys with messatis. The author captured in part of the Catatumbo refuge, west of MBrida December 1971 a male apparently very near on the northern slopes of the Andes. dodona on the Rio Anchicay6 above Buena- An additional undescribed and mimetic ventura, on the west coast of Colombia. Four subspecies ofthe lilis-group has been discovered recent specimens from the same area, in the in the collection of the British Museum Facultad de Agronomia, Maracay, Venezuela, (Natural History): include two essentially identical to dodona (Fig. 19), with the forewing postmedian band broken into separate spots. These considera- Melimea ethra kayei subspn. tions suggest that dodona, rather than erica, is the typical Choc6 race of M.ethra, identical to The prominent black-hindwing mimicry ring west Colombian phenotypes; erica would then of the Potaro area of (formerly British) Guyana be a transitional form to messatis in the central (Kaye, 1907) includes four known species in Colombian valleys. It seems extremely unlikely the genus Melinaea: M,rnneme mneme (Fig. that an isolated subspecies of the lilis-group 33), M.maenius mediatrix (Fig. 35; see below), with typical Colombian colour-pattern charac- M.ludovica crameri (Fig. 43; long confused ters, should occur in Bolivia more than with mnerne due to Cramer’s erroneous figure 4000 km to the south, where no similar of this species), and M. mnasias eratosthenes specimens have been captured in the last (Fig. 3; very rare, mixed with m.tecta). In hundred years, and which is inhabited by ano- view of this diversity already present, it was a ther conspecific race. It is noteworthy that great surprise to discover mixed among the one of the five specimens in Hewitson’s type mneme from Guyana in the British Museum series of messatis, essentially identical with (Natural History) three examples of a fifth the others, is also labelled ‘Bolivia’ (Fox, mimetic species of Melinaea. These were, once 168 Keith S. Brown, Jr

FIGS. 14-24. Melinaea ethra. lilis-group and southernmost races. All life size except 15 (0.65 X), black, yellow (or white, in 15 upper left), and orange. (14) M.e.scylux Salvin, male, dorsal, Potrerillos, Chiriqui, Panama, 31 .xii.65 (KB, G. B. Small leg.). (15) M.e. (irnirutu X parallelis), transitional series (upper left, female near parallelis Butler, others males, lower right near imirara Bates), dorsal, Cerro Campana, Panama (C. B. Small). (16) M.e.flauicuns Hoffmann, female, dorsal, Santa Rosa Cornitin, Chiapas, MBxico, vi.66 (KB, T. Escalante leg.). (17) M.e. erhru (Latreille), male, dorsal, Baixo Guandu, Espirito Santo, Brazil (KB, P. C. Elias leg.). (18) M.e.rnnemopsis Berg, male, dorsal, Colonia Hardeman, norte de Differentiation of Melinaea and Mechanitis 169 recognized, easily distinguished from all other M.m.mneme and M.maenius mediatrix sympatric ithomiines by the great extent and from the same locality (Quonga) are illustrated shape of the black area on the hindwing. Minor in Figs. 33 and 35. This represents a most un- elements of colour-pattern, especially the expected geographical and mimetic expansion fused discocellular bar on the forewing (shared of the lilis-group of M.ethra. only with M.ethra sola, Fig. 20, and some Fox’s species maelus and maeonis show races of M.mnasias), suggest that this new race many related aspects of colour and pattern, is closely allied to the lilis-group, representing and are essentially allopatric except for zamora the farthest southeastern extension of this Haensch, the unique type of which bears a fundamentally northwestern complex. generalized locality label (Ucayali). The south- western Brazilian entity brunnea Riley (Fig. 25) represents in fact an intergrading series Male (Fig. 24) between maelus cydon Godman and Salvin FW 37-42mm. Similar to M.m.mneme and maeonis patterns, specifically a new (Fig. 33), but immediately distinguishable maeonis-group subspecies described below. from this species and from M.maenius media- Other intergrading patterns, closer to cydon, trix (Fig. 35) by the extensive black suffusion are represented by the specimens in Fig. 26. on the hindwing, reaching the costal margin in Zamora suggests by its name a possible the median region and cutting off a narrow origin in southern Ecuador; however, it has orange basicostal dagger both dorsally and never been re-collected in any part of Peru or ventrally. Distinguished from sympatric M. Ecuador, and its forewing pattern does not ludovica crameri (Fig. 43) by the shape of this correspond with that of Ecuadorian maeonis, orange dagger (in crameri, a basal block) and though the overall colour-pattern might fit in by the full dark anal bar and fused discocel- with a predominant mimetic group in the lular spots on the forewing; and from upper Rio Maranon. On the other hand, the sympatric M.mnasias eratosthenes (Fig. 3) by type of zamora, in the Berlin Museum (Zoo- the lack of single yellow intervenal sub- logisches Museum der Humboldt-Universitat) marginal spots on the hindwing. Forewing (Fig. 27), is essentially identical to south- discocellular spots united into a short, western Venezuelan specimens of borealis irregular, narrow bar, and a narrow black stripe Hall (Apure refuge), opening the possibility under the forewing cubitus, both of these that the specimen is mislabelled. Forbes (1 948) characters occurring only in other subspecies considered the two as synonyms; in this case, of ethra (lilis-group) and mnasias (Figs. 1-1 3, zamora is the oldest name for the Venezuelan 20-22). Paired submarginal white spots well populations, and will be so regarded until developed ventrally. specimens of this phenotype are found else- where. Of the other subspecies mentioned by Fox Female under maelus, flavomacula Weymer, 1894: FW 43 mm. Very similar, without the dark 322 (=madeira Moulton) (Fig. 28) isa resident streak under the forewing cubitus. of the Madeira refuge area; ‘purusana’ is a Holotype d, GUYANA: Quonga (probably transitional form between flavomacula and about 6’ 30‘N, 59’W) (H. Whitely), in the maelus (TefC refuge); and ‘mayi’ is treated BMNH (Rothschild Bequest, 1939-1). below under M.marsaeus. Paratypes: one d and one 9, same data and It is also evident that taruporensis Haensch collection as holotype. (Fig. 29), placed by Fox in M.maenius, is in

Santisteban, Santa Cruz, Bolivia, 16.ii.71 (KB). (19) M.e.dodona Hopffer, male, dorsal, ‘Cali, Colombia’ (generalized), 5 .viii .70 (FA-Maracay, L. Denhez leg.). (20) M.e.solu Kaye (syntype of M.limitutu Hall), male, dorsal, Campo Alegre, Cumanh, Sucre, Venezuela (British Museum (Natural History), London). (2 1) M.e.luteapicn1is Hall, holotype male, dorsal, Mbrida, Venezuela (BM-London). (22) M.e.ezra Fox, male, dorsal, El Tucuco, Sierra Perijh, Zulia, Venezuela, 21-27 .v.71 (FA-Maracay). (23) M.e.(lilis Doubleday X lateapicalis), female, dorsal, El Mijao, Barinitas, Barinas, Venezuela, 1 .iii. 65 (FA-Maracay). (24) M.e.kayei nov., holotype male, dorsal (left) and ventral (right), Quonga, Guyana (BM-London, H. Whitely leg., Rothschild Bequest). 170 Keith S. Brown, Jr

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FIGS. 25 --32. Meliflaea erhm. maelus-maeonis complex. All life size, black, yellow, and/or orange to brown. (25) M.c.(marlus (Hewitson) or cydnn Godman & Salvin X lamasi nov.) (brZ4nneQ Riley), holo- type female (left), dorsal, Boca do Acre, upper Rio Purhs, Amazonas, Brazil (BM-London, Rothschild Bequest), and a male and a female (centre and right), Alto Rio JuruB, Acre (MN-Rio, nos. 13/831 and 13/832). (26) M.e.cydofl, varieties transitional to lamasi, three males, dorsal, Alto Rio Jurua, Acre (MN- Kio). (27) M.e.zamora Haensch, holotype male, dorsal, ‘Ucayali-Zamora’ (Zoologisches Museum der Differentiation of Melinaea and Mechanitis 17 1 fact a subspecies of the otherwise allopatric by the heavy forewing median markings, the maeluslmaeonis complex of ethra; it shows discocellular bar being characteristically close affinity with only maelus in the pattern toothed on its distal (lower) extremity. elements of the forewing median area and hindwing margin, and is associated with and Female (Fig. 3 2) nearly identical to the local race of Heliconius pardalinus (H.p.sergestus),exactly as are other Essentially identical to male, though lack- subspecies of the maelus group at lower eleva- ing the silvery hair-pencil on the costal border tions in Peru and Brazil. A transitional speci- of the hindwing, having this replaced by a black men between tarapotensis and cydon (Fig. 30) bar, as in the majority of Melinaea and Ithomi- was captured by the author near Shapaja, inae in general. 20 km southeast of Tarapoto, in December Holotype d, PERU: Madre de Dick, Iberia 1975. (1 1’22‘S, 69” 35‘W), 200 m, 8.ix.75 (Jose All maelus and maeonis related forms M. Schunke), in the Museo de Historia Natural show, like many lilis-group subspecies, a heavy ‘Javier Prado’, Universidade Nacional Mayor fusion of the end-cell spots on the forewing, de San Marcos, Lima, Peru. usually toothed on the distal border (this is Paratypes: one 0, BRAZIL: Acre, Xapuri also present in many mnasias subspecies; see (10” 38’S, 68”32’W), I90 m (Guinle & Oiti- Figs. 1-13), and a heavy, strongly angled cica), in the d’Almeida collection (no. 2979), comma-mark, which usually serve to distin- now in the Departamento de Zoologia, Uni- guish them from sympatric races of M.mar- versidade Federal do Parana, Curitiba; one d, saeus and M.maenius. By these characters, a BRAZIL: Acre, Alto Rio Jurua (about 8” 30’ new subspecies of ethra can be recognized and S, 73’ 50’W), 190m, in the Museu Nacional, separated from M.marsaeus Clara, with which Rio de Janeiro (ex coll. E. May), no. 13/830 it co-occurs in southeastern Peru and south- (Melinaea Clara, det. d’Almeida). western Brazil, and has been confused in the This subspecies, which is dedicated to Dr past: Gerard0 Lamas Muller of the Museo ‘Javier Prado’, was surely derived in the Inambari refuge, and represents a combination of Melinaea ethra lamasi subsp.n. characters from tarapotensis and cydon to the north (the last cocnected by ‘brunnea’ pheno- Melinaea Clara, ‘female allotype’ (no syste- types) and mnemopsis to the south of its matic validity as divorced from original des- range. cription), d’Almeida (1960: 9, Plate 10, Fig. 1 ). (3) Melinaea mneme (LinnC) The black-hindwing M.m.mneme (Fig. 33) is a typical member of the Guyana mimetic (Fig. 3 1 ) Male complex, as mentioned above. The lighter- FW 40 mm. Completely orange and black; hindwing M.m.mauensis Weymer is here re- hindwing clear orange with at most faint black garded as a good subspecies, originating south markings marginally; forewing apex completely of the Amazon River, though presently found black; a narrow, jagged orange postdiscal band widely in the predominantly black-hindwing on the forewing. Otherwise patterned as in northern m.mneme populations, probably in- e.maeonis, though median spots on forewing vading through the Roraima area. The sub- not fused. Readily distinguished from the species with a large orange anteapical area superficially similar marsaeus Clara (Fig. 5 1) (Fig. 34), now also widespread in mneme

Humboldt-Universitat, Berlin). (28) M.e.llavomacuIa Weymer, male, dorsal, Manicorb, Amazonas, Brazil, X-41 (MN-Rio, no. 1/036, Parko leg.). (29) M.e.tarapotensis Haensch, female, dorsal, near Shapaja, Peru, 12.xii.75 (KB). (30) M.e.(tarapotensis Xcydon), female, dorsal, near Shapaja, Peru, 12.xii.75 (KB). (31) M.e.Iamasi nov., holotype male, dorsal, Iberia, Madre de Dick, Peru, 8.ix.75 (Museu Javier Prado, Lima, J. M. Schunke leg.). (32) M.e.Iamasi nov., paratype female, dorsal, Xapuri, Acre, Brazil (Departa- mento de Zoologia, Universidade Federal do Parana, d’Almeida collection, no. 2979, Guinle & Oiticica leg. - ‘allotypr female of Melinara clara’). 172 Keith S. Brown, Jr

FIGS. 33-39. Melinaea mneme, M.maenius mediatrix, and M.menophilus aberration. All life size, black, yellow, and orange. (33) M.m.mneme (Linnh), male, dorsal (left) and ventral (right), Quonga, Guyana (BM-London, H. Whitely leg., Rothschild Bequest). (34) M.m. ca. mauensis Weymer (new subsp.?), male, ventral, Jaru, RondBnia, 5 .viii.75 (KB). (35) M.maenius mediatrix Weymer, male, dorsal (left) and ventral (right), Quonga, Guyana (BM-London, H. Whitely leg., Rothschild Bequest). (36) M.maenius mediatrix form ‘anina’ Haensch, holotype male, dorsal (upper) and ventral (lower), British Guiana (ZM- Differentiation of Melinaea and Mechanitis 173 populations, may have originated in south- Obidos, Pari, in the Museu Nacional (Rio de western Brazil where it is presently quite uni- Janeiro) (Fig. 38), is regarded as an aberration form, in populations which, being fairly well of mneme which has lost essentially all yellow differentiated from those farther to the north, markings through invasion of black pigment in may merit a name. the apical areas. A similarly apicomelanic On the basis of careful analysis of chromo- specimen of M.menophilus menophilus was some numbers, de Lesse (1970) separated captured by the author in Jaru, RondBnia, in populations formerly united under Melinaea southwestern Brazil, in August 1975 (Fig. 39). mneme in (French) Guyane into two species. One, with N= 16-17, corresponded to the (4) Melinaea ludovica (Cramer) (= egina true h4.mneme of LinnB. The other, with N = Cramer, 1777) (including M.satevis (Double- 22, was identified by de Lesse with the name day) and M.idae (Felder & Felder)). mediatrix Weymer, through designation of an The name Papilio egina Cramer, 1777 is an appropriate lectotype from the type-series of invalid junior homonym of P.egina Cramer, this taxon. Specimens of mediatrix can be 1775, and the later name ludovica of the same identified superficially by inspection of the author must be applied to the common ventral hindwing (de Lesse, 1970), which bears species with a markedly proximal yellow band a broad orange median band with no yellow on the forewing (Ebert, 1965: 76). The very scaling at the anal margin (Fig. 35); in mneme, widespread 1.iudovica (Fig. 40) is common the orange median band narrows and from the base of the Colombian, Ecuadorian becomes yellow at the anal margin (Fig. 33). and north Peruvian Andes through all of Mediatrix and its minor aberration anina northern Brazil to southern Venezuela and the Haensch (Fig. 36) are treated below under Guianas, forming everywhere an effective M.maenius; they have been found in Vene- mimetic association with the ‘silvana’ morphs zuela, the three Guianas, and Para and Amapa of (see Brown & Benson, in northern Brazil, always flying together with 1974). It is replaced in eastern coastal Brazil, M.mneme. from Paraiba to Sao Paulo, by the isolated Fox (1 960: 126) mentioned that ‘the British subspecies M.l.paraiya Reakirt which forms a Museum (Natural History) has an aberrational mimetic association with Heliconius numata specimen (of mneme) from the Rio Madeira in ethra Hiibner and robigus Weymer. A further which the postmedian fascia and subapical little-known race, M.I. manuelito Tessmann, spots of the forewing are tawny rather than occurs together with the local mimetic H. yellow.’ As M.m.mauensis is common in many numata illustris Weymer in central Peru parts of the Rio Madeira area, such an aber- (Ucayali refuge area); this subspecies (Fig. 41) ration would not be unlikely to appear; the is phenotypically quite similar to paraiya. specimen in the BM(NH) bears a note hand- The mahogany-coloured, clean-hindwing written by Fox in which he affirms that it is satevis from Bolivia (Fig. 42) and the orange- but an individual variant. The specimen, how- coloured, black- or orange-suffused-hindwing ever, was separated by Mr N. D. Riley into a crameri Godman and Salvin (Fig. 43) and drawer of mediatrix after the publication of aurantia Forbes (Fig. 44) from western Guyana de Lesse’s paper. Careful inspection of the and Venezuela possess many pattern elements ventral hindwing (Fig. 37) indicates that this in common. Fox treated these as a single may have been a correct judgment. As the species, but they are separated by 2000 km of specimen lacks all yellow pigment on the the Amazon Basin in which it is difficult to wings, however, and as the area is inhabited find reasonable candidates for linking them. It by another subspecies of maenius (‘juruaensis), seems highly unlikely that they should be so it may indeed be a simple mutant of mneme. separate and still be regarded as races of a A further unusual melanic Melinaea from single species, which in the extensive tropical

Berlin). (37) M.m. ca. mauensis aberration?, male, dorsal (left) ,and ventral (right), ‘Marmore, Rio Madeira’ (BM-London). (38) Mmneme, aberration?, male, dorsal, Obidos, Par& (MN-Rio, no. 13/785). (39) M.menophilus menophilus, apicomelanic aberration, male, dorsal (left) and ventral (right), Jaru, RondBnia, 2.viii.75 (KB). 174 Keith S. Brown, Jr

FIGS. 40-47. Melinaea ludovica. All life size, black, yellow, and orange to brown. (40) M.l.ludovica (Cramer), male, dorsal, Belhm, Parh (KB). (41) M.l.manuelito Tessmann, male, dorsal, Rio Monzhn, Tingo Maria, Peru (JP-Lima). (42) M.l.satevis (Doubleday), male, dorsal (left) and ventral (right), Caranavi, La Paz, Bolivia, 6.x.74 (KB). (43) M.1.cramei-i Godman & Salvin, male, dorsal (left) and ventral (right), Curiapo, Delta Amacuro, Venezuela, 13 .iv.69 (KB, K. Negishi leg.). (44) M.l.auran?ia Forbes, male, dorsal, Caripito, Monagas, Venezuela, 17 .ix.69 (KB, K. Negishi leg.). (45) M.l.idae Differentiation of Melinaea and Mechanitis 175 forests of the present time are virtually always being closely disposed rather than widely nearly contiguous or else even extensively separated; but this is not necessarily a ‘good’ mixed along the edges of their respective character in Melinaea, being quite variable in ranges. While widely disjunct distributions are other species. Satevis and crameri have more not rare in subtropical, temperate, or highly rounded forewings than idae or ludovica, but habitat-specialized organisms in Central and this character is variable in M.ethra and M. South America, they are practically unknown menophilus. Finally, both 1.ludovica and in tropical forest species. M.ludovica occupies crameri have been reliably recorded from essentially the entire area between satevis and western Guyana, and near Km 85 of the El crameri, and shares with them an unusual Dorado-Santa Elena road in southeastern shape of the forewing midcell and discocellular Venezuela (not in the same locality nor on the spots, and lack of the forewing hindmarginal same date), indicating possibly imperfect allo- bar. These characters are present also in the patry of these entities. However, in the only closely related northwestern M.idae complex locality known in which both have been reli- (Fig. 45), which is allopatric to both satevis ably taken on the same day (Potaro Road; and ludovica-related subspecies. Union of the Kaye, 1907), they are both rather infrequent three into a single widespread and polytypic (9% and 5%, respectively, of M.mneme), species is strongly suggested. ludovica is almost absent for half the year, A number of arguments could be advanced and ‘no less than 63% of the ludovica show against the union of these species but none is some slight development towards the pattern definitive. Although the subspecies may differ of crameri on the underside’ (Kaye, 1907: in chromosome number (de Lesse, 1970; de 41 5), in the form of an expanded black patch Lesse & Brown, 1972; Brown, Emmel & on the hindwing. Additionally, a few crameri Suomalainen, in preparation), similar differ- show a divided black patch on the hindwing ences have been found between subspecies (form ‘incisa’ Kaye), further suggesting inter- and even within local populations of mimetic gradation with ludovica. In general, crameri heliconians and ithomiines (de Lesse, 1967; seems confined to eastern Venezuela and Wesley & Emmel, 1975). The races of satevis western Guyana, from Delta Amacuro and possess a yellow spot in the anal angle of the Berbice to Roraima and commoner coastally. forewing (a character regarded as fundamental It does not occur into the interior, in south- and differentiative by Fox), and this is also eastern Guyana or southwestern Venezuela, or present in idae but absent inludovica. However, in Surinam, Guyane, or northern Brazil this is clearly a variable character in M.mnasias (Amapa, Para and Amazonas), where its co- and M.ethra subspecies, and probably in M. mimics mneme, mediatrix and Mechanitis maenius as well (see below). They also have mazaeus pannifcra, in addition to many heli- the yellow band of the forewing in a more conians and other butterflies with the same apical position than in any known forms of pattern, are common. Indeed, it seems most idae or ludovica (but see Fig. 44). However, unusual that crameri has not invaded the vast the position of this band has been shown to majority of the enormous region where its co- be controlled by a single gene in the mimetic mimics occur; the presence of a different but Heliconius numata (Brown & Benson, 1974), conspecific race (ludovica) in these areas and a similar mechanism could operate in this could help to explain the unexpected absence ithomiine species. The disposition of the of crameri. paired white submarginal spots on the ventral Thus, we have at present a number of justi- surface of ludovica (Fig. 40) is different from fications for the union of crameri, ludovica that in idae or crarneri (Fig. 43) (though more and satevis, and as yet no compelling reason similar to that in satevis, Fig. 42), each pair for their separation; certainly, they appear to

(Felder & Felder), male, dorsal, Cerro Campana, Panamh, 17 .viii.64 (KB, G. Small leg.). (46) M. (ludo- uica ludouica X mneme mauensis) (holotype of agncola Hall), probably an interspecific hybrid, male, dorsal (left) and ventral (right), Tef6, Amazonas, Brazil (BM-London). (47) M. (ludovica paraiya X erhra erhra), interspecific hybrid, male, dorsal (left) and ventral (right), Baixo Guand6, Espirito Santo, Brazil (KB, P. C. Elias leg.). 12 176 Keith S. Brown, Jr

be of common phyletic stock, and essentially In Rio de Janeiro, M.ludovica paraiya allopatric. oviposits on Markea viridiflora, a Solanaceous The orange-washed, variable aurantia tree; the mature larva has not been observed only in the Sucre area of northeastern Vene- yet, however. zuela, although a stray has been captured in Trinidad (Barcant, 1970: 275, as ‘Melinaea (5) Melinaea marsaeus (Hewitson) (includ- melus madeira’); the Orinoco delta is occupied ing M.egesta Godman and Salvin (sensu Fox, by typical crameri, which probably arose in part), M.maenius (Hewitson) (sensu Fox, part), the Imataca region. and M.mothone (Hewitson)). The northwestern races carry pattern ele- The intergrading specimens of Melinaea ments present in the crameri forewing (form from the Amazon Basin in the collections of of discocellular spots), the ludovica forewing the British Museum (Natural History) and the (proximal yellow band), and the satevis hind- Museu Nacional (Rio de Janeiro) suggest that wing (absence of the median bar), and seem to only two rather than three species are present fit satisfactorily into the species also, though in Fox’s marsueus-egesta - maenius complex, reproductive compatibility has not been for which he himself offered a supplementary tested. The more northerly idae (Fig. 45) revision (1 965). No more than two entities in forms a close mimetic association with Heli- this complex (discounting obvious intergrades) conius ismenius boulleti Neustetter and H.- are found in any given area, and intermediate hecale melicerta Bates in Panama and Colom- specimens link most of these with correspond- bia, while the west Ecuadorian race vespertina ing pairs in adjacent regions. The species mar- Fox has a narrower black hindwing border saeus, whose nominate subspecies occurs near like the sympatric H.ismenius metaphorus Tef6 with the much darker nominate sub- Weymer and the rare and recently discovered species of maenius, can often be separated H.hecale australis (see Brown, 1976a). from the latter species by the more hooked There remains in this complex Melinaea (less squared) comma-mark, the tendency of agricola Hall, an unusual form known only the forewing discocellular spots to fuse into a from the type, captured in TefC, Brazil (Fig. broad bar, and the different (often more 46). It differs from the nominate subspecies, proximal) position of the last distal undulation also known from Tefe, by the presence of a in the forewing postmedian fascia, but all of cubital spot, the reduction of yellow in the these characters are variable within and be- median band. and the complete series of tween populations. Marsaeus appears to yellow postmedian spots forming a near- include the following additional subspecies: continuous band, on the forewing; there is m.vileyi Fox (mostly western Brazil and orange in the submarginal part of cell Cul- northeastern Peru, Loreto refuge; transitions Cu2, and between the median and distal to m.rnarsaeus and the following two sub- bands of the hindwing, and a trace of a yellow species are well known (Fig. 48), including anal spot on the forewing. ‘acreana’ d’Almeida (Fig. 49); ‘manga’ Haensch A rather similar and equally puzzling speci- represents a transition to m. marsaeus) ; men (Fig. 47) was captured recently by Paulo m. macaria Godman and Salvin (mostly CCsar Elias in Baixo Guandd, Espirito Santo, southern Colombia, Putumayo refuge; many near the east coast of Brazil. This specimen, specimens farther east show influence of which will be fully discussed in a general rileyi genes); paper on Ithomiinae hybrids now in prepara- m.phasiana Butler (Fig. 50) (central Peru tion, is clearly an interspecific hybrid between to southwestern Brazil, Ucayali refuge; many M.e.ethra and M.ludovica paraiya. It seems transitions to all three adjacent subspecies are probable that the unique type of agricola, known from western Acre); representing a pattern never again collected, m.clara Rosenberg and Talbot (Fig. 51) (a is also an interspecific hybrid. One of the typical mimetic form of southern Peru and parents would be M.ludovica ludovica, while southeastern Acre, probably derived in the the other is probably the only closely related Inambari refuge); and sympatric species, M.mneme mauensis; the m.pothete d’Almeida (a variable subspecies pattern of agricola suggests exactly this mix- from Rond6nia whose populations often in- ture clude individuals tending towards phasiana Differentiation of Melinaea and Mechanitis 177

FIGS. 48-56. Melinaea marsaeus. All life size, black, yellow, and/or orange. (48) M.m. (marsaeus (Hewitson) X rileyi Fox), male, dorsal, Benjamin Constant, Amazonas (MN-Rio). (49) M.m. (marsaeus X rileyi X phasiana Butler) (paratype of M.acreana d’Almeida), male, dorsal, Alto Rio Jurui, Acre (MN- Rio, no. 13/775). (50) M.m.phasiana. male, dorsal, Alto Rio Jurua, Acre (MN-Rio). (51) M.m.clara Rosenberg & Talbot, male, dorsal, Xapuri, Acre, Brazil (DZ-Parani, d’Almeida collection). (52) M.m. pothete d’Almeida, three males including transitions to phasiana. ventral, Riozinho, Rondbnia, Brazil (KB). (53) M.m.pothete with markings similar to those of ethra races, male, ventral, Jaru, RondBnia, 28.vii.75 (KB). (54) M.m.mothone (Hewitson), male, dorsal, Chanchamayo, Junin, Peru (KB, L. & G. Harris leg.). (55) M.m. (phasiana X mothone?), female, dorsal (left) and ventral (right), Shapaja, Peru, 12.xii.75 (KB). (56) M. (marsaeus rileyiX mneme or ethra cydon?) (holotype of M.mayi d’Almeida), male, dorsal, Alto Rio Jurua, Acre (MN-Rio, no. 13/768). 178 Keith S. Brown, Jr Differentiation of Melinaea and Mechanitis 179

(Fig. 52), as mentioned in the original descrip- Brown & Benson, 1974, as M.marsaeus tion (d’Almeida, 1945), and even specimens macaria, with which it flies and which it very with ethra-like markings (Fig. 53)). strongly resembles). The melanic race mothone (Fig. 54) is Fox (1960: 149) mentions a characteristic, apparently nowhere microsympatric with vestigial and unringed yellow anal spot present marsaeus subspecies, and may be regarded on the forewing of m.egesta. This spot is un- (along with its northern differentiate messenina questionably present in the holotype of Pavo- Felder & Felder) as a premontane isolate of signata Staudinger (Fig. 59), selected by Fox this species. Transitional specimens have been but later placed by him in M.menophilus;this reported by Haensch (1 903) and illustrated by name must thereby take precedence over egesta Fox (1 960: Fig. 54). An apparent phasianal for the yellow-striped subspecies of maenius mothone transition (Fig. 55) was caught by with the broken hindwing bands. the author in December 1975 near Shapaja, Furthermore, the recently separated media- 20 km SE of Tarapoto, Peru, where the parent trix Weymer, resident in the Guianas and populations are known to meet (Fox, 1960). northern Brazil (see above, and Fig. 35), Melinaea mayi d’Almeida, a very strange shows many pattern elements in common and unique specimen from the Alto Jurua (Fig. with maenius subspecies, including that by 56), is very probably an interspecific hybrid which it is separated from sympatric mneme between marsaeus and mneme or ethra cydon; (the broad, fully orange anal end of the its very aberrant colour-pattern could be supramedian orange stripe on the ventral hind- approximated, however, by a rather selective wing). Tentatively, mediatrix is regarded as a mixture of elements occurring irregularly in northeastern isolate of the widespread species marsaeus rileyi, phasiana, Clara and macaria. maenius, differing from all other subspecies It surely is not a normal ethra race, as placed by the retention of a strong yellow spot in the by Fox (note the discocellular spots), nor can anal angle of the forewing (indicated above to its relationships be analysed simply. be variable in mnasias, ethra and ludovica sub- (6) Melinaea maenius (Hewitson) (includ- species as well). The extensive range of mae- ing M.egesta (sensu Fox, part), M.isocomma nius in eastern South America has recently Forbes, and M.mediatrix Weymer). been confirmed by the capture of m.maenius- The elements of M.maenius fit together like specimens in south-central Par& east of somewhat less comfortably than those of the middle Rio XingG (reported by David marsaeus, as fewer transitions are known and Gifford, Brasilia). large variations in pattern are evident. How- The southern race juruaensis d’Almeida ever, a long series in the author’s hands from (Fig. 60) seems to fit here also. Limoncocha in eastern Ecuador (Fig. 57) The melanic pair isocomma (Colombia) shows a part of the expected intergradation and simulator Fox (Ecuador and Peru) prob- between typical and resident cocana (whose ably belong with maenius, just as their co- type locality is very nearby) and occasional mimics messenina and mothone seem to be immigrant flavosignata Staudinger (see below) close to marsaeus. d’Almeida (1951) pointed and hicetas Godman & Salvin. A further inter- out the great resemblance between isocomma grade from southern Colombia, closer toflavo- and dark m.maenius, strengthening the signata, is illustrated in Fig. 58 (mislabelled by impression of conspecificity.

-

FIGS. 57-63. Melinaea maenius and Mmenophilus. All life size, black, yellow, and orange to orange- brown. (57) Mmaenius cocanu Haensch, and varieties, four males, dorsal, Limoncocha, east Ecuador, 31 .i.71 (KB). (58) M.maenius (cocanu Xflawosignota), female, dorsal, Vista Hermosa, La Macarena, Meta, Colombia, 25 .i .72 (KB). (59) Mmaenius flavosignata Staudinger (= egesta auctorum), holotype male, dorsal, Jurimaguas, Peru (ZM-Berlin). (60) M.maenius juruaensis d’Almeida, male, dorsal, Riozinho, Rondhia (KB). (61) M.menophi2us zaneka Butler, female, dorsal, Km 25 of the Puyo-Nap0 road, east Ecuador, 10.xii.71 (KB). (62) Mmenophilus ernestoi nov., holotype male, dorsal (left) and ventral (right), 40 km above Florencia, Caquet;, Colombia, 22 .iii.67 (MN-Rio, E. W. Schmidt-Mumm leg.). (63) Mmenophilus emestoi, paratype female, dorsal (left) and ventral (right), 40 km above Florencia, 22.iii.67 (KB, E. W. Schmidt-Mumm leg.). r 180 Keith S. Brown, Jr

(7) (Hewitson). 1650m (lo 38’N, 76Oll‘W)), is close to m. Fox’s analysis of this species requires little zaneka, suggesting that ernestoi may have a modification. It is worthy of note that typical highly restricted peripheral range to zaneka m.menophilus occurs southeast of as well as phenotypes. It may be possible that ernestoi northwest of populations of m.orestes Salvin, only represents a transitional population be- and that these two subspecies may be found tween zaneka and the wide-yellow-banded in mixed populations over much of central Colombian menophilus, though the parent Peru and southwestern Brazil, as far as phenotypes are not represented in the type- Rondhia. Colombian populations of m.meno- series. philus have wider yellow postmedian bands on The subspecies is dedicated to its collector, the forewing than southern representatives. Dr Ernest0 W. Schmidt-Mumm of Bogota, The unusual race zaneka Butler (Fig. 6 1) is who has greatly helped the author in the known through eastern Ecuador to northern understanding of the complex biogeography of Peru; at its eastern (and lower) limits it may Colombian Lepidoptera. meet menophilus, with which intergradation is suspected in such forms as ‘maculosa’ The larva of M.menophilus (Figs. 64 and Haensch and ‘discurrens’ Haensch. Menophilus 65), like those of and Aeria (Guppy, and zaneka differ in important details of size, 1904; Gilbert & Ehrlich, 1970; Brown, Lamas, colour-pattern (including the comma-mark Ebert & Dias Filho, in preparation), closely and the postmedian band on the forewing), resembles a danaine caterpillar, departing and behaviour, and probably represent the dramatically from the ‘usual’ ithomiine larval same sort of lowland/premontane pair as do form and pattern (Fox, 1967; Brown & marsaeuslmothone and maeniuslisocomma; d’Almeida, 1970). The foodplant (in like these, they are regarded as conspecific for Rondbnia) is a still unidentified epiphyte, the time being. probably Forsteronia sp. (Apocynaceae : A new subspecies, close to mzaneka, can be Echitoideae). Larvae of sympatric M. maenius found at moderate elevations in southern juruaensis and M.marsaeus pothete were Colombia, above Florencia: essentially identical and fed on the same plant. This larval type would seem to confirm Fox’s placement of Melinaea as a primitive genus, Melinaea menophilus ernestoi subsp.n. near the Tithoreini. The pupa (Figs. 66- 68) is unlike either or Ithomiinae Male (Fig. 62) models, but closer to the latter, being bowed FW 46 mm. Very similar to M.m.zaneka, more than 90° in the first abdominal segments with, however, a wider yellow postmedian like in species of more advanced ithomiine band on the forewing, and appreciable black groups, and marked like the pupa of the primi- markings on the hindwing, heavier apically. tive genus Methona which seems to be pre- cursorial to these more evolved groups (Dircennini, Godyridini). Female (Fig. 63) The coincidence of appearance and food- FW 50 mm. Similar to male, without scent- plant for larvae derived from distinct females patch on hindwing. identified as M. menophilus, M.maenius and Holotype d, COLOMBIA: Caqueta, 40 km Mmarsaeus, strongly implies that the last above Florencia, l000m (lo42’N, 75’ 38’W), word on the relationships among the various 22.iii.67 (E. W. Schmidt-Mumm), donated to races of these three species has not yet been the Museu Nacional, Rio de Janeiro. written. Series from southern Colombia (Fig. Paratypes: two dd (one strongly transitional 58), eastern Ecuador (Fig. 57), and south- to m.menophilus) and one 0, same data as western Brazil (Figs. 52, 53 and 60;d’Almeida, holotype, in the collection of E. W. Schmidt- 1945: Plate 11) suggest mutually contradictory Mumm, Bogota; one 0, same data, in the col- associations of forms, which become even lection of the author. more difficult to interpret when Peruvian and A female from somewhat farther south- Acre lots are examined. The present text can west, in Huila (road from Pitalito to Mocoa, only be regarded as logical proposal, in accord Differentiation of Melinaea and Mechanitis 181

FIGS. 64-68. Melinaea menophilus menophilus. Jaru, Rondbnia, early stages, black and yellow. (64) Third instar larva, 6 X. (65) Mature (fifth instar) larva, 3.5 X. (66) Pupa, dorsofrontal view, 5 X. (67) Pupa, ventral view, 5 X. (68) Pupa, lateral view, 5 X. with the data now in hand, subject to revision been improperly demoted to infrasubspecific when further biosystematic work becomes status; and a number of predictable forms possible. which are yet to be discovered. Although the analysis of speciation patterns may require (8) Summary revision, the fundamental patterns of sub- or By the present analysis, the genus Melinaea semi-speciation seem now to be well enough is composed of seven species, some perhaps established to permit an evaluation of the representing species-groups, redivisable when effects of Quaternary refuges on the genus, more biological information becomes available. which are fully presented in the Appendix. Sixty-three well-differentiated geographic sub- species are recognized, and, in addition, there are three transitional forms of M.mnasias, Differentiation of the species of Mechanitis each known from but a single specimen, which may merit subspecific status; also transi- The monograph of Fox (1967) requires but tional forms of other species which may have one important change. This, however, involves FIGS. 69 and 70. Mechanitis . Dorsal, 0.6 x 9 black, yellow and orange. (69) M.P. (bolivarensis Fox X polymnia (Lid)),series from Boa Vista, Roraima, north Brazil (KB). (70) M.p.(angustifascia Talbot X eurydice Haensch), series from Caranavi, La Paz, Bolivia, x.74 (KB). Differentiation of Melinaea and Mechanitis 183

the reshuffling and recombination of forms To the north, extensive intergrading series assigned by him to seven of his nine species, have been captured between p.polymnia, not forming three very widespread species, one of common west of Manaus, and the darker which is highly polymorphic in most known northeastern populations (possibly meriting a populations. name) of the Peruvian p.dorissides Staudinger (1) (LinnB) (including (doryssides auctorum), found as far east as M. isthmia Bates). TefB (Fig. 7 1). The most surprising discovery which has The races associated by Fox with isthmia come to light in the reexamination of the and polymnia, often the commonest ithomi- genus Mechanitis, shared by a number of cur- ines wherever they occur and usually rent workers in the group, is that subspecies incredibly abundant in some seasons, are assigned by Fox to the northern M.isthmia essentiaHy allopatric except for the north clearly intergrade with others assigned to M. coastal South American and Central American polymnia in various areas around the periphery doryssus, labotas and solaria (placed by Fox of the Amazon Basin. Thus, Fox (1967) with polymnia), which overlap with former named isthmia bolivarensis from southern isthmia subspecies (lycidice Bates, isthmia, Venezuela, noting that it strongly resembled veritabilis Butler, kayei Fox, caucaensis polymnia. The differences between bolivarensis Haensch, chimborazona Bates, and possibly and neighbouring (Guyana) p.polymnia are werneri Hering from the Choco, which how- appreciable, but become irrelevant when ever seems to represent a series of intergrading populations occurring in Roraima (extreme populations between caucaensis and chim- northern Brazil) are examined (Fig. 69). These borazona). Fortunately, doryssus has now include individuals approximating typical been shown to intergrade cleanly with bolivarensis and others near typical polymnia, M.lysimnia macrinus wherever the two occur and intergrades showing every possible together (see below), and the three north- recombination of the different characters. The western races of Fox’s ‘polymnia’ are thus long type-series of bolivarensis in the BM(NH), transferred to M.lysimnia. re-examined in 1974, further confirms this Northwestern populations of p.polymnia intergradation; many strikingly polymnia-like often show a dark hindwing (Fig. 72), and individuals are included, possessing much less may eventually merit a name; southern yellow on the hindwing than in Fox’s illustra- Amazon populations have already been desig- tion. A further obvious intergrade, closer to nated mauensis by Forbes (Fig. 73), and are bolivarensis, is represented by Zikan’s name often appreciably different phenotypically ‘apicenotata’ (illustrated by d’Almeida, 1956: from usual Amazon forms. However, the great 3), from the upper Rio Negro; in fact, one of dispersal ability of M.polymnia probably has the paratypes of bolivarensis comes from erased much of its past or potential differentia- slightly farther north, on the upper Rio tion in continuous tropical forests, retaining Orinoco, races only where a combination of heavy The intergradation of polymnia angusti- selective pressures and well-defined physiogeo- fascia Talbot with ‘isthmia’ eurydice Haensch graphic barriers make this possible, against the can also be seen in the other corner of the pressures of gene-flow and autoselective stabil- Amazon Basin (RondBnia, Acre, and northern ization (see Brown, 1976a: 210; Mechanitis Bolivia). A long series taken by the author in polymnia, like the heliconians mentioned there, is a mass breeder which shows deficient Caranavi, La Paz, Bolivia (600 m) in October mimetic association in many parts of its 1974 shows this integration reasonably well (Fig. 70). Further intergradation is evident in range). the Acre form ‘travassosi’ d’Almeida (I 95 1: (2)Mechanitis limnaea Forbes. 11, Plate 2, Figs. 1 and 2). However, angusti- This little known and unusual ‘splinter fascia itself may be merely a series of transi- species’ was treated satisfactorily by Fox. It tional populations between southern p.casa- has been captured in numbers recently in branca Haensch and Amazonian p.polymnia, Amapa (Ebert, 1965: 5-6) and near Obidos. though the hindwing markings are character- It is sympatric with M.polymnia and istic of only these southwestern populations. M.mazaeus, but not M.lysimnia or M.menapis, 184 Keith S. Brown, Jr

FIGS. 71-73. Mechunitis polymniu. Dorsal, black, yellow, and orange to red-brown. (71) M.p. @olymniu X var. dorissides Staudinger), series from Benjamin Constant, Amazonas, western Brazil (MN- Rio), 0.6 X. (72) M.p. ca. polymnia, female, dorsal, St Laurent, French Guyane, 25.viii.70 (W. W. Benson, Campinas), 0.8 X. (73) M.p. ca. muuensis Forbes, male, dorsal, Maubs, Amazonas (DZ-Paranh, d’Almeida collection), 0.8 X. Differentiation of Melinaea and Mechanitis 185 but it does not have any obvious relationship ‘sulphurescens’ Haensch (resembling nesaea to these last two species. It seems best to but lacking the yellow postmedian spots, or retain it as a good species though it may alternatively like lysimnia with a yellow sub- eventually be shown to be an Amazonian apical spot), which occurs as far north as does isolate derived from M.lysimnia. nesaea in BelBm, Para. In fact, these two represent the expected transitions (presuming (3) (Fabricius) (includ- a two-gene difference), but they are so infre- ing M.poZymnia (sensu Fox, part)). quent in the overlap zone that it must be Fox’s discussion of this species (1967) re- assumed that either the white/yellow and quires a few additional comments and one postmedian spots present/absent genes are important addition. closely linked, producing a ‘switch’ from one The supposed subspecies ‘connectens’ subspecies to the other, or else there are Talbot was well named. It is, in fact, but a behavioural and/or chromosomal barriers to series of intergrading populations occupying a easy fertilization between lysimnia and nesaea. narrow contact zone between M.I.lysimnia The wide overlap is not difficult to accept, as (which occurs in pure populations as far west the differences between the two subspecies as eastern Mato Grosso, near Alto Garqas, and are not large, and mobility is great in the as a minor element in ‘connectens’ popula- species; some degree of autoselective stabil- tions as far west as extreme western Mato ization may also exist, as lysimnia larvae occur Grosso) and M.1.elisa (Guerin-MBneville) in groups, though not as large as those of (which is found in nearly pure populations polymnia. The gene spread to the confines of 500km west of Alto Garqas, in the Rio the species’ range is also not hard to acknow- Cabaqal area, and occurs occasionally all the ledge, since southern Brazil populations of way east to the Serra do Mar in southeastern lysimnia occasionally include elisa-like indivi- Brazil). The area in central Mato Grosso con- duals (whose pure populations are over 1200 tains no recognizable forest refuge, and indeed km to the west), presumably from similar is mostly scrub and field interspersed with gene spread. Thus, while nesaea and lysimnia occasional gallery forest and headwater woods are accepted as conspecific their overlapping where ‘connectens’ may be found, The poly- populations seem to merit more intensive morphism of ‘connectens’ populations is evi- biological study. dent in the type-series and has been illustrated M.1ysimnia acreana d’Almeida, while (Brown, 1970). The topotypic series at hand occurring sparsely in many populations includes both M.l.lysimnia and M.I.elisa. The throughout Peru, is essentially monomorphic name ‘connectens’ should thus not continue from the Madre de Dibs region as far east as to be applied in the sense of a geographic sub- central Rondbnia (Fig. 74); in this area it is species or well-defined race. mimetic of many other endemic butterfly As d’Almeida (1 95 1) noted, M.I.lysimnia species. It probably can be regarded as a geo- and M.l.nesaea Hiibner are widely sympatric graphic race associated with the Inambari in eastern Brazil, without extensive intergrada- refuge. Further differentiation of I.elisa tion. Both have been recently captured in northward into extreme southern Colombia is mixed populations, as far north as Dois easily seen. For example, ocona Drury, with a Irmaos (Pernambuco) and as far south as more lightly marked forewing, predominates Baixo Guandu (Espirito Santo); these points in the Chanchamayo area of central Peru, and are separated by almost 1200 km of near- is much less frequent than typical elisa in continuous humid forest, and in the inter- Bolivia or other parts of Peru and Ecuador. mediate localities both phenotypes may be The discrete patterns are so mixed in most equally common. In this overlap region, the known populations, however, that no simple only candidates for intermediate phenotypes association of any of the available names are ‘albescens’ Haensch (resembling lysimnia other than ocona with any past refuge can be but with additional white postmedian spots effected, with the material presently in hand. on the forewing, where nesaea bears yellow The northwestern subspecies macrinus spots), which occurs regularly as far south as Hewitson, whose conspecificity with lysimnia the limits of 1.lysimnia in Argentina, and cannot yet be regarded as certain (some 186 Keith S. Brown, Jr

FIGS. 74-81. Mechanitis lysimnia and M.menapis, life size, black, yellow and orange. (74) M.l.acreana d’Almeida, female, dorsal, Jaru, RondBnia, 5 .vui. 75 (KB). (75) M.I.(doryssus Bates X macrinus Hewitson), four specimens showing transition, dorsal, Cerro Campana, Panamh (G. B. Small). (76) M.I. (doryssus X macrinus), two transitions, dorsal, Rio Dagua (upper) and Cananche (lower), Colombia (BM-.London). (77) M.I.(doryssusX macrinus), transitional series, dorsal (left) and ventral (right), El Tucuco, Sierra Perijh, Zulia, Venezuela, iii.68 (upper) and 21-27 .v.71 (centre and lower) (FA- Maracay). (78) M.l.so2aria Fox, transitional to doryssus (upper) and typical (lower), male and female, dorsal, Caripe, Monagas, Venezuela, 12.ix.65 and 15.ix.65 (FA-Maracay). (79) M.l.labotas Distant, female, dorsal, Rincon, Osa Peninsula, Costa Rica, 26.iv.68 (WB-Campinas). (80) M.m. danensis nov., holotype male, base of Cerro Pine near El Real, Daribn, Panamh, 16.iv.76 (American Museum of Natural History, New York, G. B. Small leg.). (81) M.m.dariensis, paratype female, base of Cerro Pirre, 15.iv.76 (AMNH-New York, K. Cook leg.). Differentiation of Melinaea and Mechanitis 187 important behavioural differences can be doryssus populations; this would explain their noted), produces at all its distribution limits appearance in Venezuela, Colombia, and unusual variants apparently intermediate to central Panama. some other race with a ‘normal’ postmedian yellow band and well-developed cubital spots. (4) Hewitson. Gordon B. Small of Panama has suggested that The discussion of this northwestern species these may be hybrids with doryssus, which by Fox (1 967) is completely satisfactory. would thus represent a very atypical subspecies Only the subspecies m.menapis in central of lysimnia; this suggestion is supported by Colombia presents a problem in its polymor- the gross allopatry of doryssus and macrinus phism, produced in part by infusion of genes in the northwestern Neotropics. In fact, as from neighbouring saturata Godman, cari- mentioned above, the recognition that Central bensis Fox and occasiva Fox. The last two American isthmia races are conspecific with subspecies intergrade in the upper Rio Amazonian polymnia required transferral of Anchicaya west of Cali, Colombia; it is possible the Mexican to north Colombian and north that the Choco populations merit a subspecific Venezuelan doryssus Bates, the Chiriqui name separate from caribensis, with which labotas Distant, and the SucreITrinidad solaria they are not identical. Intermediates to the Forbes out of the subspecies of polymnia, southwesternmost mantineus Hewitson have since they are everywhere sympatric with for- not yet been seen. Gordon Small has also in- mer ‘isthmia’-related races. In Panama, formed me that Central American msaturata doryssus occurs as far east as Veraguas province show very distinct behaviour and habitat pre- and Cerro Campana, where it encounters ferences from those of menapis in general. macrinus and produces abundant intergrading Whether this differentiation will also be specimens (Fig. 79, which can also be found reflected in reproductive isolation of saturata as far west as Costa Rica. Only macrinus is from the rest of menapis, can only be ascer- found in DariCn and the central valleys of tained when the relevant biological data are in Colombia, but doryssus is known in the Cali hand; preferably, crossing should be performed area and on the north coast of Colombia, pro- in Panama with a new subspecies of menapis, ducing similar intergrades where it meets discovered by Mr Small in the eastern part of macrinus (Fig. 76). In all of Venezuela, a form the country and described here. very near doryssus is found, but macrinus occasionally occurs in the Sierra Perija on the northwest frontier (facing the Santa Marta Mechanitis menapis dariensis subs p.n. range in Colombia). That the two hybridize is indicated by the number of intermediate Male (Fig. 80) specimens which can be captured in this FW 30 mm. Partakes of pattern character- region (Fig. 77). Thus, doryssus and solaria istics of mmenapis and m.caribensis, but the (which intergrade in eastern Venezuela; Fig. over-all appearance is unique in the species 78) are here placed as subspecies of lysimnia; and in the genus. Forewing base orange out to at least, they are conspecific with macrinus. a large black midcell spot and small black The clean-hindwing race labotas (Fig. 79), spot in the inner corner of space Cul-Cu2, predominant in the Chiriqui refuge area (along then yellow (enclosing a heavy bar-shaped with frequent doryssus and accidental comma-mark filling the marginal three-fifths macrinus) and mimetic of many sympatric of space Cul-Cu2 and fused to the black species such as Melinaea ethra scylax (Fig. 14), inner angle, and a heavy, barely joined endcell clarescens Butler and bar) to a strongly sinuate outer border identi- Ithomia celemia plaginota Butler & Druce, has cal with that in other menapis subspecies; spread out genetically to produce similar apex black with a small, diffuse yellow spot. forms today in populations of the surrounding Hindwing orange with a narrow, undulate parts of Central America, where doryssus median band, uniting with the broad black usually predominates. Labotas-like phenotypes border near vein M3 to produce a very exten- may also be produced (at least in females) in sive black apical area. Ventrally similar, with a other areas where macrinus genes contaminate yellow humeral spot on the hindwing, small 188 Keith S. Brown, Jr single white intervenal spots on both wings, verified by the above list, all or almost all of and the subapical yellow streak on the fore- the supposed subspecies of wing better developed. Patagia orange, together with the typical subspecies, abdomen yellow ventrally. mazaeus mazaeus, except for elevatus and pannifera. ’ Having verified from field work and Female (Fig. 81) museum collections that mazaeus-related Similar to male, but forewing more roun- phenotypes occur in highly polymorphic ded, endcell black bar broken into two populations from the middle Amazon up to separated spots, and inner angle of forewing the base of the Andes, without however find- may have an orange spot; lacking silvery hair- ing any clear support for either d’Almeida’s or pencil on hindwing costa. , Fox’s separation of Forbes’ mazaeus into Holoiype d, PANAMA: DariCn, near El several species, I resolved to try to discover a Real, base of Cerro Pirre (8O 04’N, 77” 44’W), new basis for possible separation of these taxa near sea level, 16.iv. 76 (G. Small), donated to into valid biological species. This came the American Museum of Natural History, through the application of the refuge model, New York. developed initially from analysis of the more Paratypes: One 0, same locality as holo- cleanly differentiated Heliconiini and Ithomi- type, 15.iv.76 (Kerry Cook), donated to the inae. In several heliconian species analysed, it AMNH; one 0,DariCn, Rio Tuquesa (8” 27’ N, became apparent that races originally generated 77” 43’W), 100 m, 11 .vii.75, in the collection by mimetic pressure in isolated refuges had, of G. B. Small, Balboa, Canal Zone; one 9, because of appreciable mobility and the possi- Panama province, Rio Bayano at mouth of bility of intersystem crossovers for mimetic Rio MajC, near high bridge of Pan-American association, spread out to appear together Highway (9’ 08’ N, 78” 46’ W), 20. x. 74, in with other phenotypes in polymorphic popula- the G. B. Small collection. tions. This was especially evident in ithomiine mimics such as isabella and Heliconius Mechanitis menapis seems in some ways hecale and numata (Brown & Benson, 1974; very close to M.mazaeus, and the two are Brown, 197613); morphs of the last species had presently sympatric only in extreme north- been placed in three different species by the eastern Venezuela (menapis caribensislmazaeus principal recent reviser of Heliconius (Emsley, beebei) ; they probably differentiated from a 1965) and in two species by myself (Brown & single widespread stock, but do not seem to Mielke, 1972), based on the sympatry of these be conspecific today. entities over large areas. (5) Mechanitis mazaeus Hewitson (includ- Once a reasonable map of the probable ing M.lanei Fox, M.messenoides Felder & refugia had been drawn by analysis of the Felder, M.egaensis Bates, and M.polymnia Heliconiini and the better-known and less (sensu Fox, part)). polymorphic Ithomiinae (including Melinaea), The confusion surrounding this poly- and once local polymorphic populations were morphic species was well discussed, if not accepted as possible in Mullerian mimics, it resolved, by d’Almeida (1951), who divided it was an easy step to determine if any taxon in into six species, and Fox (1 967), who separated the Mechanitis mazaeus complex could not be it into five; Forbes (1948) seemed to have a clearly assigned to a well-defined refuge more mature grasp of the complex unity of (through association with a principal mimetic the species. Both of the former authors based complex apparently derived therein), or else their analyses on the extensive sympatry of obviously considered as a recombinant of such related entities at a number of given collecting principal phenotypes. As indeed no such points. Said Fox, ‘only by separating them taxon could be found, and, more importantly, (mazaeus, egaensis, and messenoides) can the as no single refuge produced more than one geographic confusion . . . be resolved.’ Simi- principal and mimetic phenotype in the larly, after examining abundant material in mazaeus complex (even in the cases where this the Museu Nacional (Rio de Janeiro), refuge evidently produced a number of dis- d’Almeida remarked (in translation) ‘as can be tinct mimicry rings), it seemed probable that Differentiation of Melinaea and Mechanitis 189 190 Keith S. Brown, Jr

FIG. 84. Mechaniris mazaeus (virendu X fallux X pothetoides d’Almeida X eguensis Bates X muzaeus X nigroapicalis X lucifera Haensch), dorsal, series from Alto Rio Jurub, Acre, southwestern Brazil (MN-Rio), 0.6 X, black, yellow, and/or orange to brown. Differentiation ofMelinaea and Mechanitis 1 9 1 all the taxa belonged to a single polytypic, The named phenotypes of mazaeus include: polymorphic, and interbreeding species, beebei Forbes (Fig. 86);bipuncta Forbes (Fig. mazaeus. 88; associated with mazaeus by ventral HW This conclusion was strongly supported by pattern, though may be a race of lysimnia or, the complete and stepwise intergradations more likely, of limnaea); pannifera Butler seen between phenotypes assigned by Fox to (Fig. 87);visenda Butler (82AB, 83CD, 84CFI; mazaeus, egaensis and messenoides, or by occurs phenotypically in three isolated regions, d’Almeida to mazaeus, fallax and messenoides, in the northeast: south, and northwest of the in populations occurring in low-elevation species’ range); fallax Butler (82E, 83IM, areas near the Andes of Ecuador (Fig. 82, 84B); mazaeus (82RS, 84MNO) and its minor Limoncocha) and Colombia (Fig. 83, La variety ‘phasianita’ Haensch (84PQ); nigro- Macarena). Further series in the Museu Nacio- apicalis Haensch (82TU, 83J, 84STUVW); nal (Rio de Janeiro) from the middle and lucifera Haensch (Fig. 85); ‘williamsi’ Fox upper Amazon (Fig. 84, Alto Rio Jurua) con- (82CDM, 83A, 84E) = visenda (Napo form) x firmed the complete mixing of characters mazaeus; pothetoides d’Almeida (84GH); between these taxa, and also showed inter- ‘foxi’ d’Almeida (84R) = pothetoides X gradation to phenotypes regarded by mazaeus; egaensis (84K); ‘obscura’ Butler = d’Almeida as pannifera, egaensis, foxi, and egaensis; elevata Riley (82IJN); ‘contracta’ ‘dorissides’ (not Staudinger). Those who Riley = egaensis x elevata; ‘obumbrata’ would wish to pursue a study of this unusual d’Almeida (825) = egaensis x mazaeus; mes- polymorphic species can find in the Appendix senoides (82GH, 83NPR); deceptus Butler a summary of the probable refugial origins of (82XY) 4 ‘simplex’ Bryk; ‘holmgreni’ Bryk = the better-defined mimetic morphs of deceptus X ballucatus; ballucatus Fox (824) mazaeus, and in the following paragraph a list which is also produced by visenda (Napo)x of the known taxa in the Amazon/Orinoco deceptus. area; the numbers and letters in parentheses The recently named lanei Fox is sympatric refer to specimens illustrated in Figs. 82-84. with M.polymnia and M.1ysimnia in the region

FIGS. 85-88. Mechanitis mazaeus, dorsal, 0.75 X, black, yellow and orange. (85) M.m.lucifera, female, Benjamin Constant, western Brazil (MN-Rio). (86) M.m.beebei Forbes, female, Caripito, Monagas, Venezuela (FA-Maracay). (87) M.m.pannifera Butler, male, road from El Dorado to Santa Elena, Bolivar, Venezuela (FA-Maracay). (88) M.m.bipuncta, paratype (?) male, Surukum Basin, alto Caroni, Venezuela (FA-Maracay). 13 192 Keith S. Brown, Jr of Belkm, Para and southeastward. Although tion and lodging in eastern Ecuador; access it is quite rare, a number of recent specimens to collecting areas in northern Bolivia was are known. As mazaeus has not been captured facilitated by Madre Raymunda Maureen in the same region, it seems very likely that (Hermanas de Maryknoll). Work in the collec- the very elevata-like lanei should be the tion of the Facultad de Agronomia, Maracay, easternmost (BelBm-refuge-derived) race of and much information on Venezuelan Lepi- this highly variable species. doptera was offered by Francisco Fernandez YBpez, and Koroku Negishi and S. S. Tillett (6) Summary provided opportunities for field work in many Thus, by the present analysis, the number parts of Venezuela. H. E. Hannemann kindly of species in Mechanitis is reduced from nine sent three key Melinaea types from the to five, with a total of fifty-two recognized Zoologisches Museum der Humboldt-Univer- differentiated subspecies apparently associated sitat. Information on Peruvian Lepidoptera, with past refuges (Appendix). While the and access to important collecting localities in joining of these forms into polytypic species this country, were provided by Gerard0 Lamas may yet be modified through further field and Muller. Key specimens were sent to me also insectary experiments, the basic patterns of by L. W. Harris (Ped), Tarsicio Escalante differentiation seem to be sufficiently clear at (MBxico) and Jorge Kesselring (from many this time to permit an evaluation of the evolu- parts of South America). Without the assist- tionary forces at work in the genus during ance of the many people mentioned above, past climatic cycles; this is indicated in the this paper could not have been carried Appendix. forward to its present position. Financial assistance is also gratefully acknowledged, from the Conselho Nacional de Pesquisas (in- Acknowledgments cluding a travel grant through a collaborative The author is especially grateful to the programme with the Royal Society), the trustees of the British Museum (Natural Nacional Science Foundation (Grant GB 5398 History) and to Messrs R. I. Vane-Wright, P. R. XI), the Banco Nacional do Desenvolvimento Ackery and R. L. Smiles of the Butterfly Sec- Econ6mico (FUNTEC 47 and 101), the Minis- tion, Entomology Department of that institu- tkrio do Planejamento (FINEP/FNDCT, tion, for much assistance during work in the contrato 140/CT), and the Conselho de collection in July 1974; to A. R. do Rego Pesquisas e Ensino para Graduados of the Barros of the Museu Nacional, Rio de Janeiro, Universidade Federal do Rio de Janeiro, and Olaf H. H. Mielke of the Departamento de especially while the author was working in the Zoologia, Universidade Federal do Parana, Centro de Pesquisas de Produtos Naturais (to Curitiba, for free access to the large collec- late 1973). Photographs are by the author; tions under their care. H. Ebert, R. I. Vane- many of the enlargements were prepared with Wright, H. Real and G. Lamas M. read and the assistance of Jorge H. Lea0 and Ismael criticized the manuscript; the last contributed. Gioia. considerable taxonomic information. Boyce A. Drummond I11 offered valuable informa- tion on the species of eastern Ecuador, and References greatly contributed to the organization of the d’Almeida, R.F. (1943) Dois novos Mechanitididae manuscript. Claudionor and Paulo Char Elias do Brasil (Lep. Mechanitididae). Pap& avulsos provided key specimens from Esphito Santo; Zool. S. Paulo, 3, 165-166. Dr E. W. Schmidt-Mumrn of Bogota helped in d’Almeida, R.F. (1945) Novos Ithomiidae da fauna Brasileira (Lepidoptera Rhopalocera). Bol. Mus. access to collecting areas in Colombia and Nac., Rio de J. Zoologia 39, 1-13. donation of specimens; Gordon Small of d’Almeida, R.F. (1 950) Ligeiras notas s8bre Ithomi- Balboa, Canal Zone, likewise gave many ideas idae da America do Sul. An. Inst. Biol. Univ.MPw. and information on Panamanian species, as 20,393-397. well as specimens, photographs, and visits to a d’Almeida, R.F. (1951) Algumas consideraq6es s8bre 0s gdneros Mechanitis Fabr. e Melinaea Huebn. number of key localities. The Instituto (Lep. Ithomiidae). Bol. Mus. Nac., Rio de J. Linguistic0 del Verano provided transporta- Zoologia, 100, 1-21. Differentiation of Melinaea and Mechanitis 193 d’Almeida, R.F. (1956) Notas sinonimicas sBbre Forbes, W.T.M. (1927) The genus Melinaea Hiibner, Ithomiidae. Bol. Mus. Nac., Rio de J. Zoologia, with a description of a new species. Jl N. Y. ent. 143.1-18. SOC. 35.23-36. d’Almeida, R.F. (1 960) Estudos sobre algumas esp6- Forbes, W.T.M. (1948) A second review of Melinaea cies da familia Ithomiidae (Lepidoptera Rhopalo- and Mechanitis (Lepidoptera, Ithomiinae). Ji cera). BOl. iMus. Nac., Rio de J. Zoologia, 215, N. Y. ent. SOC. 56, 1-24. 1-31. Fox, R.M. (1949) The evolution and systematics of Barcant, M. (1970) Butterflies of Trinidad and the Ithomiidae (Lepidoptera). Univ. Pittsburgh Tobago. Collins, London. Bull. 45.1-12. Brown, K.S., Jr (1970) Proposta: Uma reserva bio- Fox, R.M. (1960) A monograph of the Ithomiidae 16gica na Chapada de Guimarzes, Mato Grosso. (Lepidoptera). Part 11. The tribe Melinaeini Clark. Brad Florestal, 4, 17-29. Duns. Am. ent. Soc. 86, 109-171. Brown, K.S., Jr (1976a) Geographical patterns of Fox, R.M. (1965) Additional notes on Melinaea evolution in Neotropical Lepidoptera. Systema- Hiibner (Lepidoptera: Ithomiidae). Proc. R. ent. tics and derivation of known and new Heliconi- SOC.Lond. B, 34, 77-82. ini (Nymphalidae, Nymphalinae). J. En?. B, 44 Fox, R.M. (1967) A monograph of the Ithomiidae (1975), 201-242. (Lepidoptera). Part IZI. The tribe Mechanitini Brown, K.S., Jr (1976b) An illustrated key to the Fox. Mem. Am. ent. SOC.22, 1-190. silvaniform Heliconius (Lepidoptera :Nymphali- Gilbert, L.E. (1969) Some aspects of the ecology dae), with descriptions of new subspecies. Trans. and community structure of ithomid butterflies Am. ent. Soc. 102, 373-484. in Costa Rica. Organization for Tropical Studies, Brown, K.S., Jr & d’Almeida, R.F. (1970) The Advanced Population Biology, individual research Ithomiinae of Brazil (Lepidoptera: Nymphalidae). reports, July-August, Ciudad Universitihia, San 11. A new genus and species of Ithomiinae with Jos6, Costa Rica, pp. 69-93 (Mimeographed). comments on the tribe Dircennini d’Almeida. Gilbert, L.E. & Ehrlich, P.R. (1970) The affinities of Trans. Am. ent. SOC. 96, 1-17. the Ithomiinae and the Satyrinae (Nymphalidae). Brown, K.S., Jr & Benson, W.W. (1974) Adaptive J. Lepid. SOC.24,297-300. polymorphism associated with multiple Miil- Guppy, I. (1904) Notes on the habits and early lerian mimicry in Heliconius numata (Lepid. stages of some Trinidad butterflies. Trans. ent. Nymph.). Biotropica, 6, 205-228. SOC.Lond. 1904,225-228. Brown, K.S., Jr & Benson, W.W. (1975) The Heli- Haensch, R. (1903) Kurzer Bericht uber des entomo- conians of Brazil (Lepidoptera :Nymphalidae). log. Ergebnisse meiner Ecuador-Reise. Die Ithomi- Part VI. Aspects of the biology and ecology of inen (Neotropiden) meiner Ecuador-Reise, mit Heliconius demeter, with description of four new Beschreibungen neuer Formen und biologischen subspecies. Bull. Allyn Mus. 26, 1-19. Notizen. Berl. Ent. Z. 48, 149-214. Brown, K.S., Jr & Mielke, O.H.H. (1972) The Heli- Haensch, R. (1909) Familie: Danaidae. In: Seitz, A. conians of Brazil (Lepidoptera :Nymphalidae). Die Gross-schmetterlinge der Erde, Stuttgart Part 11. Introduction and general comments, with (Kernen), vol. V, pp. 113-71. a supplementary revision of the tribe. Zoologica, Hewitson, W.C. (1852-76) Illustrations of New N. Y. 57, 1-40. Species of Exotic Lepidoptera, 5 vols. Privately Brown, K.S., Jr, Sheppard, P.M. & Turner, J.R.G. printed, London. (1 974) Quaternary refugia in tropical America: Kaye, W.J. (1907) Notes on the dominant Mullerian evidence from race formation in Heliconius group of butterflies from the Potaro district of butterflies. Proc. R. SOC.B, 187, 369-378. British Guiana. Trans. ent. SOC.Lond. 54, 41 1 - Brown, K.S., Jr & Vasconcellos Neto, I. (1976) Pre- 439. dation on aposematic ithomiine butterflies by de Lesse, H. (1967) Les nombres de chromosomes tanagers (Pipraeidea welanonota). Biotropica, 8, chez les LBpidoptdres Rhopalocdres n6otropicaux. 136-141. Annls SOC.ent. France N.S. 3.67-136. Collenette, C.L. & Talbot, G. (1928) Observations de Lesse, H. (1970) Formules chromosomiques de on the binomics of the Lepidoptera of Matto quelques LCpidoptdres Rhopalocdres de Guyane. Grosso, Brazil. Trans. ent. SOC.Lond. 76, 391- Annls SOC.ent. Fr. N.S. 6, 849-855. 416. de Lesse, H. & Brown, K.S., Jr (1972) Formules Ebert, H. (1965) Uma coleqlo de borboletas (Lepid. chromosomiques de Gpidoptdres Rhopalocdres Rhopal.) do Rio .4mapari (Territbrio do Amapi) du Br6sil. Bull. SOC.ent. Fr. 76, 131-137. com anotaq6es taxon6micas sabre Rhopalmera Wesley, D.J. & Emmel. T.C. (1975) The chromo- do Brasil. Pap& avulsos Zool, S. Paulo, 18, 65- somes of neotropical butterflies from Trinidad 85. and Tobago. Biotropica, I, 24-31. Emsley, M.G. (1965) Speciation in Heliconius (Lep., Weymer, G. (1894) Revision der ersten Gruppe der Nymphalidae): Morphology and geographic distri- Gattung Heliconius. Dt. ent. 2. Iris, 6,281-345. bution. Zoologica, N. Y. 50, 191 -254. Forbes, W.T.M. (1924) The genus Mechanitis Fabr., (Lepidoptera, Ithomiinae). Jl N. Y. ent. SOC. 32, 146-158. Received 25 August 1976 194 Keith S. Brown, Jr Differentiation of Melinaea and Mechanitis 195

Appendix is indicated by a question mark after the refuge name. Species within each genus are placed in a Assignment of and to Melinaea Mechanitis hypothetical evolutionary order, which differs postulated refuges from that of Fox (1960, 1967) only in details, except that Melinaea mnasias is judged to be a AS most Melinaea and Mechanitis are common relict species, the most primitive of its genus, where found, and as they are almost always and Mechanitis lysimnia is regarded as more taken by entomologists, abundant material is primitive than the polymorphic M.mazaeus. available in collections to permit evaluation of The differentiated taxa are placed in order of areas of monomorphism, polymorphism, and refuges as described in Brown (1976a). secondary hybridization. Some species are Full and up-to-date distribution maps for poorly differentiated, but the vast majority of these species will be presented in a future taxa are clearly assignable to single refuges; publication along with a more complete dis- often, this assignment is supported by mimetic cussion of centres of evolution and probable resemblance with predominant complexes pre- refuges in the Neotropics. A map of proposed sently endemic to the refuge areas. refuges, as presently understood, is included In certain cases where post-refugial dispersal here, to help clarify the following analysis. has apparently led to the blurring of sub- With each subspecies recognized in the pre- specific boundaries and to polymorphism over sent analysis is included the position given to wide areas, and the entities have been assigned this taxon by Fox (1960, 1965, 1967) with primarily on the basis of membership in the page(s) on which it was mentioned and refuge-associated mimicry rings, an asterisk is analysed and on which a distribution map for placed after the butterfly’s name in the follow- the taxon was presented by that author; also ing list. Non-asterisked names are judged to be the Figure number for the taxon in Fox (1 960, valid geographic subspecies. 1967) or in other places where it is illustrated, Uncertain assignment of a race to a refuge including in this work (‘here’).

Fox (1960, 1965) Melinaea Subspecies Presumed species or form In species Map Plate (Fig.) refuge (pages) (page) or where figured mnasias abitagua nov. - - here Abitagua lu ni lucifer (1 65) (160) V(74) Ucayali(?) comma comma (163; 81) (1 60) V(70), here Chanchamayo romualdo comma (-; 80) - - Inarnbari evx lucifer (166) (160) d’Alm. 1951;here (W Acre, Brazil) tecra mnasias (-; 82) - Haensch 1909, Roraima 33d; Kaye 1907 eratosthenes them (164) (160) V(72), here Manaus/Guina, mnasias (82) Ventuari neblinae nov. - - here Imeri mnasias mnasias (-; 81) - here Bel6m newsubsp.? - - here Tapaj6s rondonia nov. - - here Rondania newsubsp. ? - - here Tef6 lucifer lucifer (1 65) (160) V(73), here Loreto thera fhera (1 64) (160) V(71), here Rio de Janeiro mnasias (82) ethra flavicans liiis ( 12 8) (134) here (E-C M6xico) imirata lilis (129) (134) II(31), here Guatemala scylax scylax (1 3 5) (134) II(38), here Chiriqui parallelis lilis (1 30) (134) 11(32), here DariCn 196 Keith S. Brown, Jr

Fox (1960, 1965) Melinaea Subspecies Presumed species or form Plate (Fig.) refuge or where figured

erhra messatis lilis (132) I1(36), here Nechi dodona lilis (133) here Chocb ezra lilis (1 30) II(33), here (Santa Marta) la teapicalis lilis sola (1 3 1) here ‘Catatumbo lilis lilis (131) 11(34) Rancho Grande sola lilis (131) II(35), here SucrejTrinidad zamora maeonis borealis (146) Br/Ben 1974;lhere Apure maeonis maeonis (146) III(Sl), d’Alm. 1951 Nap0 tarapo tensis maenius (1 54) IV(60). here Huallaga cydon maelus (144) II1(49), Br/Ben 1974 Ucayali lamasi nov. - here Inambari mnemopsis mnemopsis (142) III(45), here Yungas kayei nov. - here ManausIGuyana flavomacula maelus madeira (143) I11(46), here Madeira maelus maelus ( 14 5) III(50), Br/Ben 1974 Tef6 erhra erhra (133) I1(37), here Bahia mneme mneme mneme (124) I1(29), here Manaus jGuiana mauensis mneme mneme (125) I1(30), Br/Ben 1974 Tapaj6s mauensis var. mneme mneme (125) here RondBnia ludovica idae idae (1 38) III(41), here Nechi vespertina idae (139) III(42) Chimborazo aurantia satevis ( 13 8) Forbes 1948, here SucreITrinidad manuelito egina (141) here Ucayali sa tevis satevis (136) II(39). here Yungas cramen‘ satevis (1 37) II(40), here Imataca ludovica egina ( 140) III(43), here Bel6m paraiya egina (141) III(44), Br/Ben 1974 Rio de Janeiro marsaeus messenina mothone (1 59) V(66), &/Ben 1974 Villavicencio macaria marsaeus (150) IV(53) Putumayo egesta (80) morhone morhone (159) V(67), here (General Andean) pkasiana phasiana (I 5 2) IV(56), here Ucayali marsaeus (80) Clara marsaeus (1 5 1) IV(SS), here Inambari egesra (80) porkere maenius (1 54) d’Alm. 1945, here RondBnia marsaeus marsaeus (149; 80) Hew. 11, Mech. 11: 10 Tef6 rileyi marsaeus (1 5 0) III(52), Br/Ben 1974 Loreto egesta (80) maenius isocomma comma (1 62) V(68) Villavicencio isocomma (81) cocana maenius (154) IV(59), here Putumayo flavosigna ta mars.egesta (149) IV(62), here Nap0 egesta egesta (80) simulator comma (162) V(69) (General Andean) isocomma (81) hicetas maenius ( 1 5 3) IV(58) Ucayali media trix mneme mneme (1 2 5) deLesse 1970, here Manaus/Guiana juruaensis maenius (1 54) d’Alm. 1943; here RondBnia maenius maenius (15 3) IV(57), Br/Ben 1974 Tef6 menophilus ernestoi nov. I here Putumayo zaneka menophilus (1 56) IV(63), here Abitagua menophilus menophilus (1 5 5) IV(61), Br/Ben 1974 Nap0 orates menophilus (1 58) V(64), Br/Ben 1974 Ucayali Differentiation of Melinaea and Mechanitis 197

~~ ~~~~~~~ Fox (1967) Mechanitis Subspecies Presumed species or form In species Map Figure (page) refuge (page) (page) or where figured polymnia lycidice i.isthmia (71) 72A(7 5) Guatemala isthmia isthmia (71) 72B(75) Chiriqui chimborazona isthmia (80) 77(75) Chimborazo caucaensis isthmia (78) 76(75) Cauca veritabilis bthmia (74) 73(75) Rancho Grande kayei isthmia (78) 75(75) Sucre/Trinidad ca. veritabilis isthmia (74) - Apure, Villavicencio donbides bthmia (80) 7 8B(7 5) Napo eurydice var. isrhmia (82) - Maranon eurydice isthmia (82) 79(75), here Chanchamayo('?) angustifacia polymnia (95) 87(93), here Yungas bolivarensis isthmia (77) 74(75), here Roraima polymnia var. p.polymnia (92) here Manaus/Guiana polymnia polymnia (9 1) 86(93), here Bel6m mauensis p.polymnia (92) here, Forbes 1948 Tapajos ca. dorimides isthmia (80) 78A(75), here Loreto casabranca polymnia (96) 88(93) Rio de Janeiro limnaea limnaea (86) 81(84), Ebert 1965 Manaus/Guiana lysimnia doryssus polymnia (87) 84(93), here Guatemala labotas pol.doryssus (90) here Chiriqui macn'nus lysimnia (1 20) 111(121), here Daribn, Nechi, Chimborazo doryssus var. pol.doryssus (87) here Rancho Grande solaria polymnia (9 1) 85(93), here Sucre/Trinidad ocona lys. elisa ( 12 3) 112B(12 1) Chanchamyo acreana 1ys.elisa (123) d'Alm. 1950, here Inambari elisa lysimnia (123) 112A( 1 2 1) Yungas nesaea lysimnia ( 12 5) 114(121), Br/Ben 1974 Bahia lysimnia lysimnia (127) 115(121), Br/Ben 1974 Rio de Janeiro menapis sa tura ta menapis (1 12) 105(114) Guatemala dan'ensis nov. - here Darih menapis menapis ( 1 16) 107(114) Nechi ca. caribensis men.samrata (1 12) - Choc6 mantineus menapis (1 18) 109(114) Chimborazo occasiva menapis (1 17) 108(114) Cauca caribensis menapis (1 14) 106(114) Rancho Grande mazaeus messenoides messenoides (107) 100(109), here Villavicencio fallax * m.mazaeus (99) here Putumayo visenda var. m.mazaeus (99) 92(100), here Nap0 deceptus messenoides (109) 101(109), here (General Andean) mazaeus* mazaeus (99), 96(105), here Ucayali eg.phasianita nigroapicalis* mazaeus (99) 93(100), here Inambari ballucams messenoides (1 10) 102(109) Y ungas beebei mazaeus (97) 90(100), here Imataca bipuncta po1.polymnia (92) here, Forbes 1948 Roraima pannifera mazaeus (98) 91(100), here Manaus/Guiana, Ventuari visenda var. maz.pannifera (9 8) Forbes 1948: Oyapock lanei lanei (84) 80(84) Belh visenda* maz.pannifera (98) here Tapajos pothetoides mazaeus (103) d'Alm. 1951, here Rondania elevata mazaeus ( 102) 94(100), here Madeira egaensis egaensis (105) 95(105), here Tef6 lucifera m.mazaeus (89) here Loreto Book notices

THE SPHECIDAE () OF a key at generic level to the known larvae. A FENNOSCANDIA AND DENMARK. By substantial bibliography, distributional tables 0. Lomholdt. Pp. 452. Scandinavian Science and an index complete the work. Press Ltd, Klampenburg, Denmark: 1975-76 D Kr. 158. (Fauna Entomologica Scandinavica, vol. 4, parts 1 and 2)

The Fauna Entomologica Scandinavica series THE OF EUROPE, Vol. 111. is conceived to give comprehensive coverage in By Kurt Harz and Alfred Kaltenback. Pp. 434. the English language of the Fauna of Dr W. Junk, The Hague, 1976. Dutch Guilders Scandinavia, with keys to genera and to 140.00, (Series Entomologica Vol. 12.) species together with descriptions, many figures and summaries of distributional and This is the third and final volume of a compre- biological information. The distributions both hensive work on the Orthoptera of Europe. It in Great Britain and in northern Germany of is the product of many years spent by the the species treated are also included, enhanc- authors studying and checking both the litera- ing both the value and the market of the series. ture and numerous large collections of Volume 4, issued in two parts, is the first Orthoptera to register the variability of treating a group of Hymenoptera. The Sphecid specific characters and to elaborate keys. The fauna of Scandinavia lacks scarcely any of the first volume dealt with the Ensifera and the 114 species present in Great Britain, and in- second with the suborder Caelifera. This cludes others which may subsequently be dis- volume, which is written in both English and covered here; the work is thus of direct value German, is concerned with the orders Cheleu- to British entomologists who may wish to toptera, Dermaptera, Mantodea, Blattoptera identify specimens. Introductory material in- and Isoptera. Following a key to the orders, cludes sections on morphology and general- each order is treated systematically with keys izations on the biology, distribution and to the superfamilks, families and subfamilies, evolution of the group. The taxonomic treat- genera and species. There are detailed descrip- ment is consistent with 3 recent reclassification tions of the included species and for each of the genera of Sphecidae and in relation to species the synonymy, detailed diagnoses and our own species list, there are few nomencla- brief distribution details are given. Line draw- tural disparities at the species level. Good keys ings are provided of all the important specific are augmented by many figures and detailed characters. An impressive list of references descriptions. Of particular interest and value is and an index completes the work.

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