Bulletin of the
British Museum (Natural History)
^36
PRESENTED
'
A review of the Miletini (Lepidoptera: Lycaenidae)
J. N. Eliot
Entomology series Vol53 Nol
25 September 1986
The Bulletin of the British Museum (Natural History), instituted in 1949, is issued in four scientific series, Botany, Entomology, Geology (incorporating Mineralogy) and Zoology, and an Historical series.
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World List abbreviation: Bull. Br. Mus. nat. Hist. (Ent.)
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Entomology series Vol53Nolppl-105
Issued 25 September 1986
(NATURAL HISTORY)
19
PRESENTED GENr
Bulletin of the
British Museum (Natural History)
Entomology series Vol 53 1986
British Museum (Natural History) London 1986
Dates of publication of the parts
No 1 25 September 1986
No 2 . . . . 30 October 1986
No 3 . . . . . . . . . . . 30 October 1986
No 4 27 November 1986
No 5 18 December 1986
ISSN 0524-6431
Printed in Great Britain by Henry Ling Ltd., at the Dorset Press, Dorchester, Dorset
Contents Entomology Volume 53
No 1 A review of the Miletini (Lepidoptera: Lycaenidae).
J.N.Eliot 1
No 2 Australian ichneumonids of the tribes Labenini and Poecilocryptini.
I. D. Gauld & G. A. Holloway ... 107
No 3 The tribe Pseudophloeini (Hemiptera: Coreidae) in the Old World tropics with a discussion on the distribution of the Pseudophloeinae. W. R. Dolling 151
No 4 The songs of the western European grasshoppers of the genus Omocestus in relation to their taxonomy (Orthoptera: Acrididae). D. R. Ragge 213
No 5 The structure and affinities of the Hedyloidea: a new concept of the butterflies.
M.J. Scoble. 251
A review of the Miletini (Lepidoptera: Lycaenidae)
J. N. Eliot
Upcott House, Bishop's Hull, Taunton, Somerset TA4 1AQ
Contents
Synopsis 1
Introduction 1
Acknowledgements 2
Checklist of the Miletini 3
Tribal characters 6
Key to the genera of Miletini 7
Genus AllotinusC.&R.Felder 7
Genus Logania Distant 57
Genus Lontalius gen. n 74
Genus Miletus Hiibner 75
Genus Megalopalpus Rober 84
References 86
Index 104
Synopsis
This review of the Miletini is based mainly on characters of the male genitalia. Keys to, and descriptions of, the genera, subgenera, species and subspecies are provided. One genus, one subgenus, eight species and seven subspecies are newly described. One species is relegated to subspecies, and three subspecies are raised to species; 41 new synonyms are established.
Introduction
The butterflies which are here considered to comprise the Oriental section of the tribe Miletini Reuter, 1897 (see Corbet, 1939b) were originally given family-group status (as the Gerydinae) by Doherty (1886), mainly on the basis of their distinctive eggs. Reuter (1897: 263) included the African genus Megalopalpus Rober, 1886, in the group, treating the latter as a tribe, the Miletidi, which is distinguished from all other Lycaenidae by the peculiar labial palpi bearing long hairs on the inner surface of the basal segment (Basalfleck). Corbet (19396) used the family group name Miletinae in place of Gerydinae because Gerydus Boisduval, 1836, is an objective synonym of Miletus Hiibner, 1819; his action is valid under Article 40b of the International Code of Zoological Nomenclature (1985). Eliot (1973) used Miletinae in a wider sense than previous authors, placing the Miletini (Miletinae sensu Corbet) as one of four tribes which he included in the subfamily.
Only two complete analyses of the Oriental Miletini (sensu Eliot) have so far been attempted, both by Fruhstorfer who based his studies in part on a number of genitalia preparations made by Reverdin. In the first (Fruhstorfer, 1913-14) it is evident that the new taxa he described had three separate origins. Many descriptions are based on specimens in his own collection, some of which bear type and/or determination labels attached either at that time or at some later date. Others relate to specimens in the collections of the British Museum (Natural History) and Walter Rothschild at Tring, to which no type or determination labels were attached; these collections were studied by Fruhstorfer during a visit to England. Finally, a few descriptions are based on records and figures by other authors of specimens which he had not seen; in Fruhstorfer's first analysis such taxa were, in most cases, based on a work by Semper (1889), so those specimens listed by Semper may be considered to comprise Fruhstorfer's type-series. In certain cases, however, it is clear that Fruhstorfer applied a name to a single specimen figured by
Bull. Br. Mus. not. Hist. (Ent.) 53 (1): 1-105 Issued 25 September 1986
2 J. N. ELIOT
Semper, and in such cases the original of the figure is automatically the holotype; in other cases, however, it has been necessary to designate a lectotype. In his second analysis (Fruhstorfer, 1916) it is evident that he had in the meantime examined Semper's collection, and he described further material to which type and determination labels were attached. There is some evidence to suggest that during this examination some of Semper's original labels were inadvertently transferred from one specimen to another. Such instances are mentioned later under the relevant taxon.
As Fruhstorfer seldom selected types, I have designated lectotypes when it is not clear that a taxon was described from a single specimen. I have also designated lectotypes for some taxa named by other authors when I have been able to recognise primary type-material in the British Museum (Natural History). When the original type-series are in museums which I have not visited, I have generally left the selection of lectotypes to future workers, except in a few cases where, to avoid confusion, I have designated figured specimens.
The following abbreviations have been used for the actual or supposed location of type- material.
BMNH British Museum (Natural History), London
CM Carnegie Museum, Pittsburgh
MNHU Museum fiir Naturkunde der Humboldt-Universitat, Berlin
RNH Rijksmuseum van Natuurlijke Historic, Leiden
SM Senckenberg Museum, Frankfurt am Main
SMT Staatliches Museum fiir Tierkunde, Dresden
ZSI Zoological Survey of India, Calcutta
Here it may be noted that there are in the BMNH collection a number of specimens bearing red type labels as well as, in some cases, labels in Corbet's handwriting identifying them as holotypes or allotypes which were not cited as such in the original descriptions. Where it is reasonable to assume that these were part of the original type-series I have normally designated them as lectotypes in preference to any other syntypes. In some cases, however, it is quite clear that the specimens cannot possibly be part of the original type-series; such specimens I have rejected as syntypic material and labelled them accordingly. These 'types' are listed, however, in the section dealing with species and subspecies since it is possible that they may have misled authors in the past or might do so in the future.
Acknowledgements
This revision is based mainly on material in the BMNH, and I am grateful to the Trustees for access to the collections and library, to Mr R. I. Vane-Wright and Mr P. R. Ackery for their help and encouragement, and to the Photographic Unit for providing the photographs for Figs 55-104. I thank especially Mr Vane- Wright and Mrs S. M. North for undertaking a review of the African genus Megalopalpus, of which I have no practical knowledge in the field, and for preparing the draft incorporated in this paper. I am also very grateful to Dr Heinz Schroder for the loan of important material from the Semper collection in the Senckenberg Museum, without which it would have been impossible to carry out a satisfactory review. I thank Dr R. de Jong, RNH, and Professor S. Murayama, who sent material for examination. Mr G. C. Treadaway placed the whole of his extensive Philippine material at my disposal and sent me the photographs for Figs 105-108. I also received valuable Philippine material through the generosity of Mr Yusuke Takanami and Dr A. Ballantine, including new taxa which I have named in their honour. Major A. Bedford Russell lent me material from Vietnam and Sulawesi, and presented the holotype of Allotinus samarensis russelli to the BMNH, and Dr E. Diehl sent me material from Simeulue. In addition the collections of the late Mr W. A. Fleming (kindly placed at my disposal by his widow), Lt-Col C. F. Cowan, Dr T. Norman and my own have been examined.
THE MILETINI
Checklist of the Miletini
Genus ALLOTINUS C. & R. Felder, [1865]
Subgenus ALLOTINUS C. & R. Felder, [1865] fallax C.&R. Felder, [1865] fallaxfallaxC. & R. Felder, [1865] sabazus Fruhstorfer, 1913 syn. n. zaradrus Fruhstorfer, 1916 fallax aphacus Fruhstorfer ,1913 ancius Fruhstorfer, 1913 syn. n. artinus Fruhstorfer, 1916 fallax eryximachus Fruhstorfer, 1913 fallax dotion Fruhstorfer, 1913 fallax tymphrest us Fruhstorfer, 1916 fallax audaxH. H. Druce, 1895 fallax apusde Niceville, 1895
michaelis Eliot, 1959 albifasciatus Eliot, 1980 subvioIaceusC. & R. Felder, [1865] subvioIaceussubviolaceusC. & R. Felder, [1865]
alkamah Distant, 1886 syn. n. manychus Fruhstorfer, 1913 syn. n. kallikrates Fruhstorfer, 1913 syn. n. silarus Fruhstorfer, 1916 syn. n. subviolaceusmirusVan Eecke, 1914 agnolia sp. n. nicholsi Moulton ,1911
nicholsi nicholsi Moulton, 191 1 nicholsi battakanus Fruhstorfer, 1913 major C. & R. Felder, [1865]
depictus Fruhstorfer, 1913 syn. n. kdldwarus Ribbe, 1926 max/musStaudinger, 1888 stat. n.
Subgenus FABITARAS subgen. n. fabius (Distant & Pryer, 1887) fabiusfabius (Distant & Pryer, 1887) caudatus Grose-Smith, 1893 syn. n. pamisus Fruhstorfer, 1914 syn. n. fa biusarrius Fruhstorfer, 1914 borneensis Moulton, 1911
elioti Corbet, 1939 syn. n. punctatus (Semper, 1889)
anaxandridas Fruhstorfer, 1916 syn. n. caesemius Fruhstorfer, 1916 syn. n. nigritus (Semper, 1889)
eretria Fruhstorfer, 1916 strigatus Moulton, 1911
strigatus strigatus Mouhon, 1911 strigatus malayanus Corbet , 1939 denalus Corbet, 1939 syn. n. brooks! sp. n. bidiensissp. n. faras(Doherty, 1889)
panormis (Elwes, 1893) syn. n. sarrastes Fruhstorfer, 1913 stat. n. mendava Riley, 1944 syn. n. porriginosus Toxopeus, 1932 syn. n.
portunus (de Niceville, 1894)
portunus port unus (de Niceville, 1894)
narsares Fruhstorfer, 1913 syn. n. portunus maitus Fruhstorfer, 1914
fruhstorferi Corbet, 1939 syn. n. portunus pyxus (de Niceville, 1894)
wdterstradti H. H. Druce, 1895
Subgenus PARAGERYDUS Distant, 1884 Miletographa Rober, 1892 horsffeldi (Moore, 1857) horsfieldi horsfieldi (Moore , 1857) horsfieldi permagnus Fruhstorfer, 1913 infumata Fruhstorfer, 1913 nessus Corbet, 1939 horsfieldi siporanus Riley, 1944 horsfieldi satelliticus Fruhstorfer, 1913 leogoron Fruhstorfer, 1916
leogoron leogoron Fruhstorfer, 1916
intricata Fruhstorfer, 1913 (unavailable name) vadosus Corbet, 1939 lindus Corbet, 1939 leogoron normani Eliot, 1967 leogoron batuensis Eliot, 1967 leogoron plessis Eliot, 1967 melos (H. H. Druce, 1896) sp. rev. reverdini Fruhstorfer, 1916 syn. n. talu Eliot, 1967 syn. n. samarensissp. n. samarensis samarensis subsp. n. samarensis russelli subsp. n. macassarensis (Holland, 1891) macassarensis macassarensis (Holland, 1891)
damodar Fruhstorfer, 1913 macassarensis menadensis Eliot, 1967 luzonensis Eliot, 1967 stat. n. albatusC. & R. Felder, [1865]
albatusalbatusC. & R. Felder, [1865] ;i//>;ifi/,v/iiriH/;/vsiihsp. n. apries Fruhstorfer, 1913
apriesapries Fruhstorfer, 1913
eupalion Fruhstorfer, 1914 syn. n. apries dositheus Fruhstorfer, 1914 apries ristus subsp. n. corbeti Eliot, 1956 unicolorC. & R. Felder, [1865]
unicolorunicolorC. & R. Felder, [1865] eurytanus Fruhstorfer, 1913 syn. n. dilutus Corbet, 1939 syn. n. unicolor continental Fruhstorfer, 1913
dtddnus Fruhstorfer, 1916 unicolor rekkia Riley & Godfrey, 1920 unicolor mooreiH. H. Druce, 1895
rebilus Fruhstorfer, 1913 syn. n. unicolor aphocAaKheil, 1884
myridndus Fruhstorfer, 1913 syn. n. unicolor posidion Fruhstorfer, 1913
molionides Fruhstorfer, 1913 syn. n.
J. N. ELIOT
niceratus Fruhstorfer, 1913 syn. n. bajanus Fruhstorfer, 1913 syn. n. enatheus Fruhstorfer, 1913 suka Piepers & Snellen, 1918 enganicus Fruhstorfer, 1913 syn. n. unicolor georgius Fruhstorfer, 1913
leitus Fruhstorfer, 1916 unicolor zitema Fruhstorfer, 1916 paetus(de Niceville, 1895) parapus Fruhstorfer, 1913 nivalis (H. Druce, 1873) nivalis nivalis (H. Druce, 1873) nivalis felderi Semper , 1 889 substrigosus (Moore, 1884) substrigosus substrigosus (Moore, 1884)
magaris Fruhstorfer, 1913 substrigosus lenaia Fruhstorfer, 1913 substrigosus sibyllinus Riley, 1944 substrigosus ballantinei subsp. n. substrigosus yusukei subsp. n. da vidis Eliot, 1959 (I mini In (Moore, [1866]) drumila drumila (Moore, [1866]) multistrigatus de Niceville, 1886 insignis (Staudinger, 1888) drumila aphthonius Fruhstorfer, 1913 grisea Riley & Godfrey, 1920
Genus LOGANIA Distant, 1884 Malais Doherty, 1889 malayica Distant, 1884 malayica malayica Distant, 1884 malayica subura Fruhstorfer, 1914 nehalemia Fruhstorfer, 1914 stat. rev. waltraudae sp. n. regina (H. Druce, 1873) regina regina (H. Druce, 1873)
evora Fruhstorfer, 1916 syn. n. regina sriwa Distant, 1886 paluana sp. n. marmorata Moore, 1884 marmorata marmorata Moore, 1884 marmorata damis Fruhstorfer, 1914 marmorata hilaeira Fruhstorfer, 1914 obscura Distant & Pryer, 1887
(nom. preocc.) nada Fruhstorfer, 1914 stenosa Fruhstorfer, 1916 syn. n. cineraria Fruhstorfer, 1916 syn. n. sora Fruhstorfer, 1916 syn. n. marmorata lahomius(K.hei\, 1884) marmorata diehJi subsp. n. marmorata munichya Fruhstorfer, 1914 marmorata javanica Fruhstorfer, 1914
glypha Fruhstorfer, 1914 marmorata palawana Fruhstorfer, 1914
distanti Staudinger, 1889 (nom. preocc.) marmorata samosata Fruhstorfer, 1914 marmorata faustina Fruhstorfer, 1914
obscura Rober, 1886
martinus (Fruhstorfer, 1913) syn. n. donussa Fruhstorfer, 1916 syn. n. distanti Semper, 1889
distanti distanti Semper , 1889 apsines Fruhstorfer, 1914 turdeta Fruhstorfer, 1916 syn. n. distanti massalia Doherty, 1891 stat. n.
luca de Niceville, 1894 syn. n. distanti drucei Moulton ,1911 distanti staudingeriH. H. Druce, 1895 hampsoni Fruhstorfer, 1914
meeki Rothschild, 1915 syn. n. masana Fruhstorfer, 1916 syn. n. watsoniana de Niceville, 1898 subfasciata Tytler, 1915
Genus LONTALIUS gen. n. eltussp. n. eltus eltus subsp. n. eltus treadawayi subsp. n.
Genus MILETUS Hubner, 1819 Symetha Horsfield, 1828 Gerydus Boisduval, 1836 Archaeogerydus Fruhstorfer, 1916 chinensisC. Felder, 1862
chinensischinensisC. Felder, 1862
chinensislearchusC. & R. Felder, [1865] irroratusH. Druce, 1874 kelantanus Corbet, 1938
chinensis assamensis (Doherty, 1891) milvius (Fruhstorfer, 1913)
chinensis longeana (de Niceville, 1898) croton (Doherty, 1889)
croton croton (Doherty, 1889) tavoyana (Evans, 1932)
croton corus Eliot, 1961
croton karennius (Evans, 1932) mallus (Fruhstorfer, 1913)
mallusmallus (Fruhstorfer, 1913)
mallus gethusus (Fruhstorfer, 1913)
mallus shanius (Evans, 1932) gaesa (de Niceville, 1895)
gaesagaesa (de Niceville, 1895)
gaesa carrinas (Fruhstorfer, 1916) nymphis (Fruhstorfer, 1913)
n ymphis n ymphis (Fruhstorfer, 1913)
nymphis porus Eliot , 1 96 1
nymphis fictus Corbet, 1939
nymphis eneus Eliot, 1961 zinckeniiC. & R. Felder, [1865]
zinckeniizinckeniiC. & R. Felder, [1865]
zinckenii improbus (H. H. Druce, 1895) gopara (de Niceville, 1890)
goparagopara (de Niceville, 1890) denticulata (Fruhstorfer, 1913)
gopara pardus Eliot , 196 1
gopara eustatius (Fruhstorfer, 1913)
THE MILETINI
gopara artaxatus (Fruhstorfer, 1913)
oichalia (Fruhstorfer, 1913) valeus (Fruhstorfer, 1913)
pallaxopas (Fruhstorfer, 1913) gaetulus (de Niceville, 1894)
gaet ulus gaet ulus (de Niceville, 1894)
gaetulus innocens(H. H. Druce, 1895)
gaetulus aphytis (Fruhstorfer, 1913) boisduvali Moore, 1857
boisduvali boisduvali Moore, 1857 vincula (H. H. Druce, 1895) heraeon (Fruhstorfer, 1916) courvoisieri (Fruhstorfer, 1915) oxylus (Fruhstorfer, 1916) lombokianus (Fruhstorfer, 1913) acragas (Doherty, 1891) buruensis (Holland, 1900) ceramensis Ribbe, 1889 dossemus (Fruhstorfer, 1913) stygianus Butler, 1884 adeus (Fruhstorfer, 1913)
boisduvali diotrophes (Fruhstorfer, 1913)
boisduvali avitus (Fruhstorfer, 1916) drucei (Semper, 1889)
druceidrucei (Semper, 1889) philippus (Staudinger, 1889) jacchus (Fruhstorfer, 1913) palanius (Fruhstorfer, 1913) epidurus (Fruhstorfer, 1913)
drucei metrovius (Fruhstorfer, 1913) phradimon (Fruhstorfer, 1915) biggsii (Distant, 1884)
biggsii biggsii (Distant, 1884) atomaria (Fruhstorfer, 1913) xeragis (Fruhstorfer, 1916) hyllus (Fruhstorfer, 1916) sebethus (Fruhstorfer, 1916) extraneus (Toxopeus, 1929)
biggsii natunensis (Fruhstorfer, 1916)
biggsii niasicus (Fruhstorfer, 1913) batunensis (Fruhstorfer, 1913)
biggsii albotignula (Van Eecke, 1914)
simalurensis (Toxopeus, 1928) cellarius (Fruhstorfer, 1913) symethus (Cramer, 1779)
symethus symethus (Cramer, 1779) pandu (Horsfield, 1828)
symethus petronius (Distant & Pryer, 1887) diopeithes (Fruhstorfer, 1913) bangkanus (Fruhstorfer, 1914) hieropous (Fruhstorfer, 1916)
symethus solitariusOkubo, 1983
symethus acampsis (Fruhstorfer, 1913)
symethus nuct us Eliot, 1961
symethus perlucidus (Fruhstorfer, 1913) megaris (Fruhstorfer, 1913)
symethus vespesianus (Fruhstorfer, 1913)
symethus batuensis (Fruhstorfer, 1914)
symethus edonus (Fruhstorfer, 1913) symethus philopator (Fruhstorfer, 1914) symethus hierophantes (Fruhstorfer, 1916) symethus phant us subsp. n.
gallus (de Niceville, 1894) gallus gallus(de Niceville, 1894) gallus leucocyon (Toxopeus, 1940) heracleion (Doherty, 1891)
heracleion heracleion (Doherty, 1891) heracleion arion Eliot, 1961 a/icon (Doherty, 1889) ancon ancon (Doherty, 1889) ancon gigas(H. H. Druce, 1895)
anconides (Fruhstorfer, 1913) archilochus (Fruhstorfer, 1913)
archilochus archilochus (Fruhstorfer, 1913) archilochus siamensis (Godfrey, 1916) gigantes(de Niceville, 1894) atimonicus Murayama & Okamura, 1973 celinus Eliot, 1961 takanamiisp. n. /eos(Guerin-Meneville, 1830) leosteos (Doherty, 1891) leos fforensis (Fruhstorfer, 1913)
eulus (Fruhstorfer, 1913) leostellus (Fruhstorfer, 1913) leos catoleucos (Fruhstorfer, 1913) leos maximus (Holland , 1 89 1 ) divisa (Fruhstorfer, 1913) sarus (Fruhstorfer, 1913) leosvaneecki (Toxopeus, 1930) leos mangolicus (Fruhstorfer, 1913) /eos/eos(Guerin-Meneville, 1830) boisduvalii (Butler, 1884) meronus (Fruhstorfer, 1913) amphiarus (Fruhstorfer, 1913) gardineri (Fruhstorfer, 1914) leos virtus (Fruhstorfer, 1913)
pentheus (Fruhstorfer, 1913) leosaronicus (Fruhstorfer, 1914) nineyanus (Fruhstorfer, 1914) acrisius (Fruhstorfer, 1914) melanion C. & R. Felder, [1865]
melanionmelanionC. & R. Felder, [1865]
albiguttatus f. n. (infrasubspecific name) melanion euphranor (Fruhstorfer, 1914) bazilan us (Fruhstorfer, 1913) stat. n.
vitelianus (Fruhstorfer, 1913) syn. n.
Genus MEGALOPALPUSRober, 1886
angulosus Griinberg, 1910 mete/eucusKarsch, 1893 simplex Rober, 1886
bicoloria (Capronnier, 1889)
similis (Kirby, 1890)
gigas Bethune-Baker, 1914 zymna (Westwood, 1851)
pallida Aurivillius, 1922
6 J. N. ELIOT
Tribal characters
The characters of Miletini (sensu Eliot, 1973) are as follows. Eyes smooth. Antennae with narrow, gradually incrassate club and with the nudum extending down the shaft to the base or very nearly so. Labial palpi asymmetrical, but there is no constancy in which palpus is the longer. In most species the palpi are unusually long and thin, more so in females than in males, and protrude well beyond the head; they are also unusual in that the 'Basalfleck' of Reuter is clothed with hairs of unknown function. Proboscis long, bearing many sensilla throughout its length. Legs more or less abnormal, with the tarsi very long, flattened and blade-like in Miletus and Megalopalpus, cylindrical but very long and thin in Allotinus and with the tibiae outwardly swollen or incrassate in Logania and Lontalius. The mid- and hind-tibiae lack the usual pair of terminal spurs. The male fore-tarsus is reduced to a single segment ending abruptly but with a small pointed process directed downwards, except in the typical species of Logania in which the tarsus tapers to a down-curved point. The claws on the mid- and hind-tarsi of both sexes are small, and minute on the fore-tarsi of females. The abdomens of males are long and protrude well beyond the hindwings, except in Allotinus major. There is a double hair tuft, sometimes inconspicuous but sometimes large and erectile, on the sternum of the eighth segment; it is not known whether it plays any part in courtship. The tergum of the eighth segment is unusually long to accommodate the peculiar genitalia, and bears a long apophysis at its proximo-ventral edge. The wing venation shows a high degree of individual variation, and does not often provide good characters for separating genera and species. The forewing always has 11 veins (vein R4 missing), and veins Sc and RI are separate throughout their lengths. In the males of most species the basal portion of vein M3 is slightly swollen and devoid of normal cover scales, but bears small to very small specialised (?scent) scales. The hindwing has a well-developed humeral vein in Megalopalpus and Lontalius, but in the other genera it is only weakly developed or absent. The male genitalia are highly characteristic. The uncus and tegumen are in the form of enormous paired plates, which are attached to the vinculum only narrowly in the dorsal region, so that they are capable of considerable freedom of movement . Articulating brachia are always present . The vinculum bears on either side two more or less triangular processes, one directed proximad and the other distad. The latter process is comparatively weakly sclerotised and overlies the sides of the tegumen. The valvae are small, with the outer dorsal portion (sometimes referred to as the ampulla) bearing a dense hair fringe. The juxta is present as a furca whose arms are united by a band above the phallus just distad of the ductus. Eliot (1973: 386) incorrectly called this band a form of transtilla. The female genitalia have not been investigated.
The early stages are very imperfectly known. According to Doherty (1889) the eggs of Miletus and Allotinus are much flattened and disc-like, but in Logania are stouter and scarcely more than twice as wide as high. They bear between two and five lateral carinae which are either simple or broken into short teeth placed one above the other, giving the appearance of a cogged wheel. The larvae, so far as known, are wholly aphytophagous, feeding on Homoptera. They are more or less cylindrical and have a particularly thick cuticle, and apparently lack the 'honey gland' on the seventh and paired eversible tubercles on the eighth abdominal segments which are present in the majority of Lycaenidae. Their relationship with the ants attending the Homoptera appears to be one of neutrality, and from this it appears at least possible that the larvae may be furnished with small glands on a number of segments, as in many other Lycaenidae (Cottrell, 1984), and that these secrete some substance which inhibits ant aggression. In one species, Miletus boisduvali, the larva pupates inside ants' nests and the pupa has attractant glands and is attended by ants; the emerging adult is clothed with fugitive scales, as in Liphyra brassolis, which confuse attacking ants (Roepke, 1918). However, in M . chinensis, the larva pupates in the open and is attached by the cremaster with or without a weak girdle (Kershaw, 1905) , and in this species, as well as in Allotinus subviolaceus, there is no evidence to suggest that fugitive scales are present in the adult (Piepers & Snellen, 1918). The adults feed on the excretions of Homoptera and do not visit flowers.
The tribe comprises four Oriental genera and one African genus (the latter erroneously described from Borneo). The Oriental genera, totalling 69 mainly Sundanian species, have been
THE MILETINI 7
subjected to a full taxonomic revision. No comparable attempt has been made to revise the four species of Megalopalpus , but a review of the genus, based on a draft kindly prepared by Mr R. I. Vane-Wright and Mrs S. M. North, is included.
In the keys and descriptions which follow, the system of veins and spaces is as in Fig. 49 (p. 74).
Key to the genera of Miletini
1 Tarsi with first segment more or less cylindrical 2
- Tarsi with first segment flattened and blade-like 4
2 Tibiae swollen or incrassate 3
- Tibiae not swollen; legs long, thin, cylindrical ALLOTINUS (p. 7)
3 Hindwing without a humeral vein. Legs comparatively short LOGANIA (p. 57)
- Hindwing with a humeral vein . Legs long and thin LONTALIUS (p. 74)
4 Hindwing without a humeral vein. Oriental MILETUS (p. 75)
Hindwing with a humeral vein. African MEGALOPALPUS (p. 84)
Genus ALLOTINUS C. & R. Felder
Allotinus C. & R. Felder, [1865]: 285. Type-species: Allotinus fallax C. & R. Felder, [1865]: 285, pi. 35, fig. 24, by designation of Scudder, 1875: 107. Gender masculine.
Legs long, thin, cylindrical. Labial palpus with third segment longer than half second segment. The wing venation shows a high degree of infra-specific individual variation. Hindwing cilia elongated into short tufts at vein endings in the male, the margin more or less crenulate and tufted in the female. Underside characteristic, white to pale buff, densely striated with small striae or spots, with heavier spots arranged in the usual lycaenid pattern. Forewing usually with very small white costal flecks at ends of veins Sc, R\, R2, /?3 and, in one species-group, R5 also.
The genus ranges from north India to Sundaland, the Philippines and Sulawesi.
Fruhstorfer (1913-15; 1916) divided the genus into two subgenera: Allotinus for species allegedly without a sex stripe in the male and Paragerydus for species with a sex stripe. I maintain these subgenera, though not in the arrangement adopted by Fruhstorfer, and add a third subgenus.
Key to the subgenera of Allotinus
1 cf valva ending in an apical point; abdominal hair tufts poorly developed, not protruding except
when genitalia are extruded. Hindwing usually with a weak humeral vein 2
- Cf valva with costa abruptly truncate (except in A . davidis) , but ending in a ventral , more or less
pointed process; abdominal hair tufts prominent. Hindwing without a humeral vein
PARAGERYDUS (p. 30)
2 Antenna barely longer than half the forewing costa, with less than 50 segments, cf forewing with
vein A/3 not, or only very briefly, swollen ALLOTINUS (p. 7)
- Antenna nearly two-thirds length of forewing costa, with 60 or more segments, cf forewing vein
M3 prominently swollen, clothed with specialised scales for one-third of its length or longer
FABITARAS (p. 20)
Subgenus ALLOTINUS C. & R. Felder
The principal characters of the subgenus are as given in the key.
Nineteenth century authors separated Allotinus from Paragerydus by the former's possession of a short upper discocellular vein, but this character does not occur in A. (A.) agnolia, wherein veins R5 and Ml have a short common stalk. Typical species, in which the males have vein M3 unswollen, have vein /?3 long, arising closer to the cell apex than to the wing apex; but in the two species in which the males have vein A/3 weakly swollen, vein /?3 is shorter, arising close to or opposite the end of vein R2.
On the basis of the male structure the subgenus can be divided into two species-groups: the/fl/fax-group in which the abdomen is of normal length for the tribe and the uncus/tegumen plates are long and narrow; and the major-group in which the abdomen is much shorter and the uncus/tegumen plates are more rounded.
The subgenus has a restricted distribution in Sundaland, the Philippines and Sulawesi, and comprises seven species.
8 J. N. ELIOT
Key to the species of subgenus Allotinus
1 cf abdomen longer than hindwing dorsum. Underside of hindwing with postdiscal spot in space
6 more or less below the spot in space 7 and remote from the spot in space 5 (fallax- group) 2
Cf abdomen same length as hindwing dorsum. Underside of hindwing with postdiscal spot in space 6 more or less equidistant from the spots in spaces 7 and 5 (major-group) 6
2 Upperside with white or grey-blue areas 3
Upperside plain brown nicholsi (p. 16)
3 Upperside with white areas 4
Upperside with greyish-blue areas subviolaceus (p. 14)
4 cf upperside of hindwing brown or diffusely sullied with white scales 5
Cf upperside of hindwing with a clearly defined white band albifasciatus (p. 13)
5 Forewing with veins R5 and MI just separate at their origins, cf forewing vein M3 not swollen nor
clothed with specialised scales. Underside with postdiscal series of spots clearly marked
fallax (p. 8)
- Forewing with veins RI and M\ stalked, cf forewing vein M3 briefly swollen and clothed with
specialised scales. Underside with postdiscal series not apparent on forewing and barely discernible on hindwing agnolia (p. 15)
6 Smaller, forewing 13-5-19-0 mm. cf upperside of forewing all brown or with a small white or
whitish spot comprising at most a small sullied area at base of space 2 and a larger white area up to 3-0 mm wide in space Ib. $ with a larger white area filling basal third of space 2 and about half of space Ib, but not reaching vein Cu\ nor entering the cell; white patch may be reduced, sullied or absent major (p. 17)
- Larger, forewing 20-0-21-0 mm. cf upperside of forewing with a white patch filling basal third of
space 2 and about half of space Ib which is basally grey. $ with white patch larger, extending above vein Cu\ and into lower edge of cell maximus (p. 19)
Allotinus (Allotinus) fallax C. & R. Felder
(Figs 1-3, cf genitalia) Allotinus fallax C. & R. Felder, [1865]: 285, partim.
The Felders confused two distinct species under this name. In pi. 35, fig. 24 they figured as a male what is currently treated as the female of A. fallax, and at figs 25, 26 as the female a different species which I describe later as the female of A. (Paragerydus) albatus mendax. Semper (1889: 163) correctly pointed out that the Felders' 'male' was in fact a female. However, he incorrectly identified their female as a male
Fig. 1 Allotinus (Allotinus) fallax fallax C. & R. Felder; Luzon. Male genitalia. Above, lateral view of right half of armature ; below, dorsal view of phallus.
THE MILETINI 9
variety of fallax, having a white area on the upperside of the hindwing such as he had seen only in a few examples from Luzon; these may have been either males or females ofmendax. Apart from these supposed variants from Luzon he correctly stated that the male of fallax has a plain brown hindwing in the Philippines.
The species is distinguished by lacking a swelling of vein A/3 in the male and by the presence of a white patch on the upperside of the dark brown forewing. The hindwing of the male is brown, except in Borneo where it is usually dusted with white scales. Except in Sumatra and the Malay Peninsula, where the hindwing is brown, the female has a white discal area (rarely vestigial or absent) which shows a high degree of individual variation, so that it is an unreliable character by which to distinguish subspecies.
On the basis of the male genitalia A. fallax can be divided into three subspecies-groups.
Type (a) (Fig. 1): valva with short apical hook and ventro-distal edge not strongly exarcate; phallus comparatively short, broad and abruptly tapered at distal end; Philippine.
Type (b) (Fig. 2): valva with short apical hook and ventro-distal edge strongly exarcate; phallus comparatively long and thin and gradually tapered at distal end; Philippine.
Fig. 2 Allotinus (Allotinus) fallax aphacus Fruhstorfer ; Mindanao. Male genitalia.
Type (c) (Fig. 3): valva with long apical hook; phallus like type (c), but still slimmer; Sundanian.
So far as my experience goes type (a) occurs exclusively in Luzon, Masbate, Bohol, Samar and Leyte, and type (b) exclusively in Mindanao and Bazilan. Both occur in Mindoro, each type having a distinctive phenotype. I have not been able to dissect males from Cebu, Panaon, Camiguin de Mindanao and the Sulu Is. , whence Fruhstorfer named subspecies, but I think it certain that type (a) will be found in Cebu and type (b) in Camiguin de Mindanao and Sulu Is. , whilst in Panaon either might occur. The occurrence of type (a) in Bohol, Samar and Leyte is rather unexpected; usually butterflies from these islands are associated with Mindanao rather than with Luzon. The dichotomy in Mindoro can be paralleled in the lycaenopsid Acytolepis puspa which occurs in Mindoro in subsp. cagaya and the very different-looking subsp. bazilana in equality and without evidence of intergradation (Eliot & Kawazoe, 1983: 183). Conceivably types (a) and (b) may represent distinct species which evolved in the northern and southern groups of islands respectively and which may now have a wider overlap in distribution than is at present known; otherwise it is difficult to see how type (b) could have reached Mindoro without also becoming established in the intervening islands.
A. fallax flies at low to moderate elevations, and is common and widespread in the Philippines, but rare in Sundaland. It has not been found in Palawan, but is likely to occur there.
Key to the subspecies of A. (A.) fallax 1 cf genitalia of type (c) . Sundanian
- cf genitalia of types (a) or (b). Philippine
2 cf genitalia of type (b). $ upperside of hindwing with white area normally confined to basal two-thirds of spaces 4 and 5 , but may be vestigial or obsolete
10 J. N. ELIOT
Cf genitalia of type (a). $ upperside of hindwing with white area normally extending below vein A/3, often as far as vein A! fallax fallax (p. 10)
3 cf upperside of forewing with white patch reaching vein Cu\ 5
Cf upperside of forewing with white patch usually not above mid-space 2; if reaching vein Cui,
upper part narrow and sullied 4
4 cf white patch comparatively large, reaching into space 2. Underside of forewing with central
area above dorsum sparsely striated fallax aphacus (p. 11)
Cf sullied white patch smaller, usually confined to space Ib. Underside of forewing with central area above dorsum as densely striated as rest of wing fallax eryximachus (p. 11)
5 Underside ground colour pale buff. $ upperside of hindwing with a small white patch
fallax dotion (p. 12) Underside ground colour whitish buff, with pale markings. $ hindwing white patch obsolete
fallax tymphrestus (p. 12)
6 Upperside of hindwing with a dusting of white scales usually present in cf and always present in
$ fallaxaudax (p. 12)
Upperside of hindwing plain brown fallax apus (p. 12)
Allotinus (Allotinus) fallax fallax C. & R. Felder (Fig. 1, cf genitalia)
Allotinus fallax C. & R. Felder, [1865]: 285, partim 'cf ', recte $ [nee $ , pi. 35, fig. 24 'cf']; Semper, 1889: 163, partim, pi. 31, figs 23cf, 26$. LECTOTYPE $, PHILIPPINES: Luzon (BMNH), here designated [examined].
Allotinus fallax fallax Felder; Fruhstorfer, 1913: 343; 1916: 809.
Allotinus fallax sabazus Fruhstorfer, 1913: 343; 1916: 809. LECTOTYPE cf, PHILIPPINES: Bohol [not located] . Syn. n.
Allotinus fallax zaradrus Fruhstorfer, 1916: 809. Syntypes, PHILIPPINES: Cebu [not located].
In the male the white patch on the forewing almost always extends across vein Cu\ into space 3. In the female the white patch on the hindwing is variable in extent, but almost always extends dorsad into space 3 and often as far as space Ib. The ground colour of the underside is pale buff, richer in the male than in the female.
The Felders did not specify a type. Their 'male', as pointed out by Semper (1889: 163), is a female, while their female belongs to another species described later as A. (Paragerydus) albatus mendax. In BMNH there is a male ex Felder coll. bearing a BMNH red type label, but as the original figure and description does not apply to this sex it seems better to reject this unpublished type selection and to designate as lectotype a female in BMNH labelled /79/Luzon Lorquin [round blue]/Felder Coll. /Rothschild Bequest 1939-17.
When naming sabazus, Fruhstorfer, evidently referring to the male, stated that the white patch on the forewing was reduced compared with subsp. fallax and that the underside was darker and more closely striated. The male from Bohol figured by Semper has the white patch entirely below vein C«i, as in occasional Luzon examples, but in a series of both sexes from Bohol in coll. Treadaway the males have the white patch extending above vein Cu\ and in neither sex can I detect any consistent differences from Luzon fallax. I therefore consider that sabazus cannot be maintained as a valid subspecies. There is in SM a female labelled /Bohol/Coll. C. Semper/2 11/Typus [red]/, but in my view it cannot be accepted as the type of sabazus since Fruhstorfer's description did not apply to this sex. I therefore designate as lectotype the male figured by Semper (pi. 31, fig. 23), which unfortunately has not been recognised in SM.
Semper recorded fallax from Cebu, but did not figure it. It is not clear whether Fruhstorfer (1916) was merely attaching a name to Semper's record or actually saw specimens from Cebu. No examples from Cebu can now be found in SM, but according to Treadaway (pers. comm.) it is thought that some may possibly exist but be temporarily mislaid. The only specimen that I have seen from Cebu is a female in coll. Treadaway. This has the underside ground colour somewhat whiter than usual, but the difference from Luzon females seems too slight to justify retaining zaradrus as a valid subspecies, and I therefore provisionally synonymise it with /a/tax.
Examples from Mindoro and Sibuyan Is. do not differ from normal fallax, but two males from Masbate in coll. Treadaway have a more whitish underside, and a female from Panay is almost pure white and relatively weakly striated below and has an exceptionally large white patch on the upperside of the hindwing. I provisionally place these under fallax .
Males from Samar and Leyte seem never to have the forewing white patch extending above vein Cui and
THE MILETINI 11
in both sexes the underside ground colour is darker buff than in examples from Luzon and Bohol. These possibly constitute a distinct subspecies to which Fruhstorfer's name artinus, given by him to the population in Panaon, may apply. Unfortunately I have seen no males from Panaon and Semper's figure of a Panaon male (pi. 31, fig. 24) shows only the underside, from which it is not possible to establish whether or not it resembles Samar and Leyte examples. The single female from Panaon which I have seen does not differ from Mindanao females, and I therefore provisionally synonymise artinus with aphacus.
DISTRIBUTION. Luzon, Mindoro, Sibuyan, Cebu, Bohol. In slightly different forms also in Masbate, Panay, Samar and Leyte. I have seen no examples from Negros, but it must occur there as well as in other islands.
Allotinus (Allotinus) fallax aphacus Fruhstorfer (Fig. 2, cf genitalia)
Allotinus fallax C. & R. Felder; Semper, 1889: 163, partim, pi. 31, figs 24 cf , 25 $ .
Allotinus fallax aphacus Fruhstorfer, 1913: 343; 1916: 809. Holotype $, PHILIPPINES: Camiguin de
Mindanao [not located]. Allotinus fallax ancius Fruhstorfer, 1913: 343; 1916: 809. LECTOTYPE cf , PHILIPPINES: Mindanao
(BMNH), here designated [examined]. Syn. n. Allotinus fallax artinus Fruhstorfer, 1916: 809. LECTOTYPE $, PHILIPPINES: Panaon (SM), here
designated [examined].
On the upperside the forewing white patch of the male is barely half as large as in subsp. fallax, seldom fully crossing space 2, and when it does so the upper part is sullied with brown scales. The female has the white patch on the hindwing almost always restricted to the basal two-thirds of spaces 4 and 5, and often it is almost obsolete. On the underside of the forewing there is an almost unstriated central area above the dorsum. The male genitalia are type (b).
Fruhstorfer named aphacus from Semper's record and figure 25 of a female from Camiguin de Mindanao. The original of this figure, which I have not been able to locate, is therefore automatically the holotype. I have seen a single male labelled /Cam. de Mind./211/Coll. C. Semper/ which does not differ from Mindanao males.
I designate as lectotype of ancius a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Mindanao/Mindanao Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/Allotinus fallax ancius [in Corbet's hand]/. A female labelled /Type [red]/Type [Fruhstorfer orange]/Mindanao/Mindanao Fruhstorfer/Fruh- storfer Coll. B.M. 1933-131/fallax ancius Fruhst. [in Fruhstorfer's hand]/ becomes a paralectotype.
I designate as lectotype of artinus a female in SM labelled /Coll. C. Semper/Panaon/211/Typus [red]/, which does not differ from Mindanao examples.
DISTRIBUTION. Mindanao, Camiguin de Mindanao, Panaon. I also provisionally place under this subspecies a female in BMNH from Talaud I.
Allotinus (Allotinus) fallax eryximachus Fruhstorfer
Allotinus fallax eryximachus Fruhstorfer, 1913: 343; 1916: 809. LECTOTYPE cf , PHILIPPINES: Mindoro (BMNH), here designated [examined].
This is the darkest subspecies. As pointed out by Fruhstorfer in his original description, the forewing white patch in the male is even more insignificant (unbedeutenderen) than in subsp. aphacus, being always more or less sullied with brown scales and sometimes restricted to a mere streak below vein Cu2. On the underside the central area above the forewing dorsum, which is not, or is comparatively weakly, striated in the other subspecies, is striated all over. The male genitalia are type (b).
Fruhstorfer stated that the type was in BMNH. However, there are no males agreeing with his description labelled as types, but a pair of subsp. fallax have been placed in the type drawer above the name eryximachus. The male, with a large white forewing patch, is labelled /Mindoro Philippine Is. Dr. Platen/Godman-Salvin Coll. 1908-168/A. fallax 107 Var. ?/, and the female is labelled like the male but with an additional label /Allotinus 'cf ' fallax Felder/. Agrees with figure of type F. A. H. 8. x. 09/. Neither specimen bears a BMNH red type label, presumably because they cannot be reconciled with Fruhstorfer's description of eryximachus. Although Fruhstorfer may have seen this pair in BMNH they are not to be considered as types, and I designate as lectotype a male from a series which Fruhstorfer may have seen at Tring and which he may have confused with what he saw in BMNH; it is labelled /Mindoro Platen/ Rothschild Bequest 1939-1/. It is interesting that Platen should have caught both subspecies in Mindoro.
DISTRIBUTION. Mindoro, where it appears to outnumber sympatric subsp. fallax.
12 J. N. ELIOT
Allotinus (Allotinus) fallax dotion Fruhstorfer
Allotinus fallox dotion Fruhstorfer, 1913: 343; 1916: 809, pi. 141h cf $ as 'dolion\ LECTOTYPE cf, PHILIPPINES: Bazilan (BMNH), here designated [examined].
On the upperside both sexes resemble subsp. fallax, with the white patch on the forewing of the male unsullied and reaching vein €1*2 broadly. On the underside the ground colour is a paler, more whitish buff than in subspp. sabazus and aphacus. The male genitalia are type b.
I designate as lectotype a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/Philippinen Bazilan II-III. 98 Doherty ex coll. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/fallax dotion Fruhst. [in Fruhstorfer's hand]/. A female labelled similarly, except that the final label reads /dotion [in Fruhstorfer's hand]/, is labelled paralectotype.
DISTRIBUTION. Bazilan.
Allotinus (Allotinus) fallax tymphrestus Fruhstorfer
Allotinus fallax tymphrestus Fruhstorfer, 1916: 809. LECTOTYPE cf, PHILIPPINES: Sulu Is. (SM), here designated [examined] .
I have seen only the original pair from which Fruhstorfer described the subspecies. The male exactly resembles subsp. dotion on the upperside, but on the underside the ground colour is slightly more whitish and the markings less well-defined. The female lacks the usual white patch on the upperside of the hindwing, but it may be that a white patch is sometimes present. The subspecies is doubtfully separate from subsp. dotion.
I designate as lectotype the male in SM labelled /Coll. C. Semper/Sulu Inseln Meyerink/211/Typus [red]/. The female is labelled paralectotype.
DISTRIBUTION. Sulu Is.
Allotinus (Allotinus) fallax audaxH. H. Druce
Allotinus audox H. H. Druce, 1895: 564, pi. 31, figs 11 cf , 12 $>. LECTOTYPE cf , BORNEO: Mt Kina Balu
(BMNH), here designated [examined]. Allotinus fallax audax Druce; Fruhstorfer, 1913: 343; 1916: 809; Corbet, 1939ft: 66.
In both sexes the forewing white patch is more extensive than in the Philippine subspecies, almost reaching the wing base in space Ib where it is dusted with grey scales. The hindwing is more or less lightly dusted with white scales in the male, but heavily dusted over most of the wing in females.
The subspecies was described from an unspecified number of specimens in coll. H. H. Druce and Staudinger. I designate as lectotype a male in BMNH labelled /Kina Balu Waterstr. A. audax co-type H. H. Druce/ex coll. Hamilton Druce 1919/Type PT [green]/Joicey Bequest Brit. Mus. 1934-120/. A female labelled $ co-type, but otherwise as the male, is a paralectotype.
DISTRIBUTION. Borneo; apparently only known from Mt Kina Balu.
Allotinus (Allotinus) fallax apus de Niceville (Fig. 3, cf genitalia)
Allotinus apus de Niceville, 1895: 27, pi. S, fig. 17 $; Fruhstorfer; 1913: 343; 1916: 809. Syntypes $,
SUMATRA (? ZSI). Allotinus fallax apus de Niceville; Corbet, 1939ft: 66; Eliot, 1962: 218; 1978: 240; 1980: 145, figs 7 cf , 14 cf
genitalia; Fleming, 1975; 22, pi. 50, fig. L50 cf . Allotinus fallax michaelis Eliot, 1959: 376, partim cf nee $ , pi. 10, fig. 8 cf . Holotype cf , WEST MALAYSIA
(BMNH) [examined]. [Synonymised by Eliot, 1962: 218.]
Differs from subsp. audax in the plain brown hindwing in both sexes.
A. apus was described from two females from north-east Sumatra, which should be in ZSI. There is in BMNH a female labelled /Type [red]/Battak Mts. N.E. Sum. 11. 94 Dr Martin/ Allotinus apus de Niceville and, [on reverse] , Collect Dr. Martin/Rothschild Bequest B.M. 1939-1/. As there is no evidence to suggest that this female was one of de Niceville's original two I reject it as a type.
DISTRIBUTION. Sumatra and West Malaysia, at elevations of about 500 m upwards.
THE MILETINI
13
Fig. 3 Allotinus (Allotinus) fallax apus de Niceville; Malay Peninsula. Male genitalia.
Allotinus (Allotinus) albifasciatus Eliot
(Fig. 4, cf genitalia)
[Allotinus fallax michaelis Eliot, 1959: 376 (partim, $ nee cf), pi. 10, fig. 9 <j>. Misidentification.] [Allotinus fallax audax H. H. Druce sensu Eliot, 1978: 241. Misidentification.]
Allotinus albifasciatus Eliot, 1980: 143, figs 5 d", 6 <j>, 14 cf genitalia. Holotype cf, SUMATRA (BMNH) [examined].
The species is distinctive in possessing a clear white band crossing the hindwing in both sexes. On the underside the ground colour is off-white, with the postdiscal markings larger and more blotchy than in A. fallax.
DISTRIBUTION. The species is extremely rare, and has so far only been found in the Malay Peninsula and Sumatra at elevations of about 1000 m.
Fig. 4 Allotinus (Allotinus) albifasciatus Eliot; Sumatra. Male genitalia.
14 J. N. ELIOT
Allotinus (Allotinus)subvioIaceusC. & R. Felder
(Fig. 5, cf genitalia) Allotinus subviolaceus C. & R. Felder, [1865]: 286.
The greyish blue areas on the upperside render this species unmistakable. As in the white areas of A. fallax these blue areas are individually variable in extent, especially on the hindwing where they may be absent or cover almost the whole wing. On average they are more extensive on both wings in the female.
The species has the widest distribution in the subgenus, occurring from Assam to Java and the Philippines.
Key to the subspecies of A. subviolaceus
1 Upperside greyish blue. Upperside of forewing in cf with space 3 all or almost all black
subviolaceus subviolaceus (p. 14)
Upperside pale grey, with only a slight blue tint , becoming whitish in lower half of forewing disc. Upperside of forewing in cf with basal half of space 3 and base of space 4 bluish grey
subviolaceus mirus (p. 15)
Allotinus (Allotinus) subviolaceus subviolaceus C. & R. Felder
(Fig. 5 cf genitalia)
Allotinus subviolaceus C. & R. Felder, [1865]: 286, pi. 35, figs 27, 28 cf ; Piepers & Snellen, 1918: 17, pi. 20, figs 20a cf , 20b $ , 20c larva. LECTOTYPE cf , JAVA (BMNH), here designated [examined].
Allotinus alkamah Distant, 18860: 452, pi. 44, 'cf ' recte <j>. Holotype <j>, WEST MALAYSIA (not located). Syn. n.
Allotinus subviolescens [sic] Felder; Swinhoe, 1910: 196, pi. 616, figs 1, la $, Ib cf.
Allotinus subviolaceus alkamah Distant; Fruhstorfer, 1913: 342; 1916: 808; Corbet, 1939a: 66; Eliot, 1978: 240; Fleming, 1975: pi. 57, fig. L37 $.
Allotinus subviolaceus subviolaceus C. & R. Felder; Fruhstorfer, 1913: 342; 1916: 808, pi. 141g £.
Allotinus subviolaceus manychus Fruhstorfer, 1913: 342; 1916: 808; Evans, 1932: 212; Cantlie, 1967: 28. LECTOTYPE <j>, BURMA (BMNH), here designated [examined]. Syn. n.
Allotinus subviolaceus kallikrates Fruhstorfer, 1913: 342; 1916: 808. LECTOTYPE cf, PHILIPPINES: Mindanao (BMNH), here designated [examined]. Syn. n.
Allotinus subviolaceus silarus Fruhstorfer, 1916: 808. LECTOTYPE cf , BORNEO (BMNH), here desig- nated [examined]. Syn. n.
Males have a broad black border on the forewing which often fills the whole of space 3 and the cell, but in some examples the blue may enter the lower half of space 3 and fill the lower two-thirds of the cell. Usually the hindwing has only a few blue scales.
In the female the forewing border is much narrower, and the hindwing varies individually from wholly fuscous to nearly all blue.
In Palawan the greyish blue colour tends to be paler in both sexes than in other areas, and this does not appear to be due to season, as in the still paler examples flying in Burma and Thailand during the dry season. In addition the forewing border tends to be wider in females, measuring an average of 2-5 mm at vein Cu2 compared with an average of 2-0 mm in normal examples. These differences seem hardly sufficient to justify erecting another subspecies. Females from Borneo and Bangka also have, on average, the forewing border similar to Palawan examples.
The Felders described subviolaceus only from the male, but did not designate a type. I therefore designate as lectotype a male in BMNH labelled /Type [red]/Java Coll. . . . [illegible]/[circular blue]/ Allotinus subviolaceous Feld. /Felder COLLN. /subviolaceus n. /Rothschild Bequest B.M. 1939-1/.
The holotype of alkamah is the 'male' (recte female) figured by Distant, which has a fuscous hindwing. In BMNH there is a Sumatran female labelled /Type [red]/alkamah Dist. /Sumatra Forbes/Rothschild Bequest B.M. 1939-1/alkamah Dist. $ Allotype [in script believed to be by Corbet]/. As the holotype is a female the allotype cannot be another female , and I reject it as such . Fortunately Corbet does not appear to have published this type selection.
Fruhstorfer did not designate a type of manychus. He had no examples from Burma in his collection, but said that he had examined examples in BMNH from Pegu and Rangoon. There is a female in BMNH labelled /Type [red]/Moore Coll. 1908-208 Pegu Magaree/May be taken as type of Allotinus subviolaceus
THE MILETINI
15
Fig. 5 Allotinus (Allotinus) subviolaceus subviolaceus C. & R. Felder; Malay Peninsula. Male genitalia.
manychus Fruh. [in script believed to be by Corbet]/. As it is likely that Fruhstorfer saw this specimen during his visit to BMNH I designate it as lectotype.
I designate as lectotype of kallikrates a male in BMNH labelled /Type [red]/Type [Fruhstorfer orangej/Mindanao Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/subviolaceus kallikrates Fr. [in Fruhstor- fer's hand]/. A female labelled /Type [red]/Type [Fruhstorfer orangej/Mindanao/Mindanao Fruhstorfer/ Fruhstorfer Coll. B.M. 1933-131/Allotinus subviolaceus kallikrates Fruh. [in Corbet's hand]/ is a paralec- totype.
I designate as lectotype of silarus a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Nordborneo Brunei Waterstradt 1890/Fruhstorfer Coll. B.M. 1933-131/Allotinus subviolaceus silarus Fruh. [in Corbet's hand]/. A female in BMNH labelled /Type [red]/W. Borneo Sintang 16. III. 10/ Fruhstorfer Coll. B.M. 1933-131/subviolaceus silarus Fr. [in Fruhstorfer's hand]/ is labelled paralectotype.
DISTRIBUTION. Manipur; Burma; Thailand; West Malaysia; Singapore; Sumatra; Bangka I.; Java; Borneo; Palawan; Mindanao.
Allotinus (Allotinus) subviolaceus mirus van Eecke
Allotinus subviolaceus mirus van Eecke, 1914: 247; Fruhstorfer, 1916, 808. LECTOTYPE cf , SIMEULUE I. (Simalur) (RNH), here designated [examined].
The subspecies is much paler than the palest dry season examples from Burma.
Above, both sexes are pale bluish grey, becoming whitish on the forewing in the middle of space Ib and basal half of space 2. In the male the forewing border is inwardly rather irregular and narrow, leaving the basal half of space 3 , the base of space 4 and the lower three-quarters of the cell grey ; it is narrowest at vein Cu2, where it is just over 2-0 mm wide. The hindwing is mostly grey, with the discocellular veins darkened and with an inwardly diffuse fuscous marginal and costal border 2-3 mm wide. The female differs as usual in having more extensive bluish grey areas on both wings.
I designate as lectotype a male in RNH labelled /60. 49/E. Jacobson Sinabang. Sim. Sum. 7. 1913/ Museum Leiden Allotinus subviolaceus mirus Det. R. v. Eecke/Type/. A further male and a female labelled /60. 50/ and /60, 51/ respectively, but otherwise as the lectotype, are paralectotypes.
DISTRIBUTION. Simeulue I. (Simalur).
Allotinus (Allotinus) agnolia sp. n.
(Figs 6 cf genitalia and scales; 55 9)
Cf forewing length 12-5 mm. Forewing with apex rounded, as in females of the genus; veins M\ and R5 with a short common stalk and vein R3 shorter than in the three preceding species, arising only a little before end
16
J. N. ELIOT
Fig. 6 Allotinus (Allotinus) agnolia sp. n.: Sumatra. Male genitalia. Lower left, androconial scale from fore wing vein M3; below, normal cover scale from the same area.
of vein R2. Upperside dark brown, with basal quarter of forewing a paler grey brown. Forewing with a rhomboidal white patch about 3-5 mm broad resting on vein A\ and reaching into lower angle of cell and base of space 3. Basal quarter of vein A/3 rather weakly swollen and clothed with very pale buff specialised scales about one-sixth size of white cover scales. Underside with a whitish area above mid-dorsum of forewing , but rest of both wings pale buff very densely striated with buff-brown striae ; usual postdiscal and submarginal markings of the genus small and only made out with difficulty.
9 . Apart from lacking the swelling of vein M3 of forewing, similar in all respects to male.
MATERIAL EXAMINED
Holotype cf , Sumatra: 'Battak Mts., N.E. Sum., II. 94 (Dr. Martin)' (BMNH).
Paratypes. Sumatra: 1 $ (allotype), 'VIII. 94', otherwise as holotype (BMNH); 1 9 'N.O. Sumatra Martin Coll. H. Fruhstorfer', 'XL 94', 'Fruhstorfer Coll. B.M. 1933-131' (BMNH).
The specific name is an anagram of Logania, given because the holotype was found in BMNH among a series of Logania distanti massalia bearing a manuscript label (in an unknown hand) reading Logania massalia Doherty.
Allotinus (Allotinus) nicholsi Moulton
(Figs 7, cf genitalia; 56 cf ; 57 $) Allotinus nicholsi Moulton, 1911: 83.
The species differs from all the preceding in being reddish brown without white or blue areas. On the forewing vein R3 is as short as in Paragerydus, arising opposite the end of vein R2. On the hindwing a weak humeral vein is present. In the male, vein M3 of the forewing is weakly swollen in its basal quarter (a fact not noticed by Moulton and Corbet) , and the specialised scales are slightly smaller than those of A . agnolia . The male genitalia possess a feature unique in the tribe, viz. a prominent cornutus in the phallus, consisting of a longer central and shorter outer bundles of minute spicules.
The species is known only from Borneo and Sumatra, and appears to be very rare. Provisionally I recognise two subspecies, but more material may show that no subdivision is necessary.
Key to the subspecies of A. (A.) nicholsi
1 Underside markings faint and comparatively broad nicholsi nicholsi (p. 17)
- Underside markings narrow, darker and sharply defined nicholsi battakanus (p. 17)
THE MILETINI
17
Fig. 7 Allotinus (Allotinus) nicholsi battakanus Fruhstorfer; Sumatra. Male genitalia.
Allotinus (Allotinus) nicholsi nicholsi Moulton
Allotinus nicholsi Moulton, 1911: 83; Fruhstorfer, 1913: 343; 1916: 809; Corbet, 19396: 66, fig. 11 cT
genitalia. Holotype cf , BORNEO: Sarawak (BMNH) [examined]. Allotinus nicholsi nicholsi Moulton; Eliot, 1967: 71.
The subspecies is known only from the male holotype, which is in worn condition. On the forewing veins MI and R5 are just separate. On the underside the markings are rather faint and comparatively wide, and resemble those of A. subviolaceus, as pointed out by Moulton.
Allotinus (Allotinus) nicholsi battakanus Fruhstorfer
(Figs 7 cf genitalia; 56 cf ; 57 9)
Allotinus taros battakanus Fruhstorfer, 1913: 370 (partim); 1916: 813, ? pi. 141g $ (partim). LECTOTYPE
Cf , SUMATRA (BMNH), here designated [examined]. Allotinus nicholsi battakanus Fruhstorfer; Eliot, 1967: 71.
The short series of one male and four females in BMNH have veins M^ and R5 of the forewing connate , and on the underside the markings are narrower, darker and more sharply defined than in subsp. nicholsi.
Although Fruhstorfer drew attention to the absence of the usual forewing brand of the male, part of his type-series, including two female syntypes in BMNH, comprise A. (Fabitaras) sarrastes, which is a common species in Sumatra. No doubt it was on this account that Fruhstorfer stated that the species was taken in great numbers by Dr Martin. It is not possible to say whether the figure of a female in Seitz represents battakanus or sarrastes.
I designate as lectotype the male in BMNH, unfortunately lacking its head, which is labelled /Type [red]/Type [Fruhstorfer orange]/CMB II. 95/Fruhstorfer Coll. B.M. 1933-131/Allotinus battakanus Frhst. [in Fruhstorfer's hand]/.
DISTRIBUTION. Known only from the Battak Mts.
Allotinus (Allotinus) major C. & R. Felder sp. rev. (Fig. 8, Cf genitalia)
Allotinus major C. & R. Felder, [1865]: 286, partim cf nee $, pi. 35, fig. 29 cf. LECTOTYPE cf, SULAWESI (BMNH), here designated [examined].
18 J- N. ELIOT
Allotinus fallax major Felder; Fruhstorfer, 1913: 343; 1916: 809.
Allotinus fallax depictus Fruhstorfer, 1913: 343; 1916: 809 [misspelled depista], pi. 141h cf [as major].
LECTOTYPE cf , SULAWESI (BMNH), here designated [examined]. Syn. n. Allotinus kalawarus Ribbe, 1926: 91. Syntypes, SULAWESI (probably in SMT).
The most remarkable feature of this species and A. maximus is the short abdomen, which in the male does not protrude beyond the hindwings; this is due to all the abdominal segments being relatively short compared with those of the other species of the genus. Both species can also be recognised by the arrangement of the postdiscal series on the underside of the hindwing, wherein the spot in space 6 is midway between those in spaces 7 and 5.
A. major shows much individual variation in size and pattern. The male is typically of medium size (forewing 17-0 mm) with a small, sullied, white patch on the forewing astride the base of vein Cu2. Occasionally the white area below vein Cu2 is unsullied and up to 3-0 mm wide; but in more than half of the examples examined the white patch is entirely absent, and such examples from east and south Sulawesi were treated as a distinct subspecies, depictus, by Fruhstorfer. However, similar males occur throughout Sulawesi, so that depictus can only be used as a varietal name. Females have, on average, a larger white patch than males. In the commonest form the ovate white patch is about 4-0 mm wide and fills the centre of space Ib, the basal half of space 2 and may enter space 3. Unmarked brown females appear to be very rare, as there is only one example in BMNH, from west central Sulawesi, without a trace of white. Other females from the same area and also from east Sulawesi form a complete connecting series leading up to the normal white-patched form.
I have not seen Ribbe's type-series of A. kalawarus, so cannot be positive that the name is applicable to A. major. Ribbe described his species from at least four examples which he did not sex. He wrote that the upperside agreed with the figure of major (Fruhstorfer, 1916: pi. 141h) which is cf-var depictus and that the underside was nearest to the figure of pyxus [A. portunus pyxus] (Fruhstorfer, 1916: pi. 141i), but differed in having a light whitish ground colour and heavier marginal flecks and other markings. He queried whether kalawarus might not be the same as damodar (= A. macassarensis macassarensis), but said that the only specimen which he had as damodar was a female with a crenulate hindwing termen (which is indeed a character of macassarensis female), whilst in kalawarus the termen was not crenulate. A non-crenulate hindwing is a feature of both sexes of A . major, so that if any of Ribbe's specimens were females this would confirm that they were major. If they were males the absence of any mention of a brand (conspicuous in A . macassarensis) also suggests that they were major. I therefore feel reasonably confident that kalawarus is a synonym of major.
Fig. 8 Allotinus (Allotinus) major C. & R. Felder; south-west Sulawesi. Male genitalia. Lower left, uncus, tegumen and brachium of an example from north-east Sulawesi: Minahassa.
THE MILETINI 19
In the male genitalia the uncus/tegumen blades are shorter and more rounded than in any other Oriental species of Miletini. Judging by the few preparations I have made they are roundest in south Sulawesi and most elongated in Minahassa, those from east-central Sulawesi being intermediate.
I designate as lectotype of major a male in BMNH labelled /Celeb Lorquin type [blue circular]/Allotinus major cf Fd./Type [red]/FELDER COLLN. /Rothschild Bequest 1939-1/. The female figured by the Felders is an example of the next species.
I designate as lectotype of depictus a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Sud-Celebes Dr. Martin Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/major fa depicta Fr. [in Fruhstor- fer's hand]/. A female labelled/Type [red]/Type [Fruhstorfer orange]/O. Celeb. Fruhstorfer/depictus Fruh. [in Corbet's hand]/ is a paralectotype; it is a typical example with a white patch on the forewing.
DISTRIBUTION. The species occurs at low levels throughout Sulawesi and in Banggai I. and Sangihe I.
Allotinus (Allotinus) maximus Staudinger stat. n. (Fig. 9, cf genitalia)
[Allotinus major C. & R. Felder, [1865]: 286, partim $ nee cf , pi. 35, figs 30, 31 $. Misidentification.] Allotinus albatus Felder var. maximus Staudinger, 1888: 269, pi. 94. Holotype cf, SULAWESI (coll.
Staudinger, probably in MNHU). Allotinus fallax major Felder $-f. albatus Felder [= maximus}; Fruhstorfer, 1913: 343; 1916: 809
[misspelled albadus and magnimus}.
Both Fruhstorfer and Staudinger confused this species with Allotinus albatus, a distinct species in subgenus Paragerydus (p. 30). The former recognised its kinship with A. major, regarding it as a montane variety from Minahassa. I think it is more likely that major and maximus are distinct sibling species. The latter may well be wholly montane, since of four males in BMNH the only two with complete data came from the Peak of Bonthain, 1000-2000 m. These examples from the extreme south-west of Sulawesi do not appear to differ from Staudinger's holotype male from Minahassa, so that it is probable that the species occurs unchanged throughout the mountains of Sulawesi.
A. maximus is larger than A. major, and in the male there is a clear white patch on the forewing filling the basal half of space 2 and the central half of space 1 b , which is grey at the wing base . The only female I have seen is the misidentified female syntype of major figured by the Felders, which has a large white patch on the forewing extending into the base of space 4 and the lower edge of the cell. It is labelled /Type [red]/Celebes Lorquin type [blue circular]/Allotinus major n $ Fd. /major n/FELDER COLLN. /Roth- schild Bequest 1939-1/.
The male genitalia (Fig. 9) of the single example dissected differ slightly from those of A. major in having a narrower, more pointed phallus and longer, more rectangular uncus blades.
DISTRIBUTION. Confined to Sulawesi.
Fig. 9 Allotinus (Allotinus) maximus Staudinger; Sulawesi (no further data). Male genitalia.
20 J. N. ELIOT
FABITARAS subgen. n.
Type-species: Paragerydus fabius Distant & Pryer, 1887: 266. Gender masculine.
Antenna nearly two-thirds length of forewing costa, with about 62-66 segments. In males, abdominal hair tufts short, not protruding except when genitalia are everted. Forewing venation particularly subject to individual variation, with apical angle of cell sometimes sharply acute, sometimes almost a right angle. Veins Ml and R5 usually stalked, but sometimes connate from cell apex, most often in A. taras and A. strigatus, while in A. bldiensis they may be just separate. Vein RT, also very variable, usually arising from about middle of vein R5, but sometimes as short as in subgen. Paragerydus, sometimes as long as in typical species of subgen. Allotinus. In males vein M3 always basally swollen and clothed with small specialised scales. Hindwing usually with a weakly developed humeral vein extending less than half-way across space 8, but sometimes absent or vestigial, most often in females. Underside characterised by a series of submarginal blackish dots which, on the forewing, are outwardly white-edged in females but not, or more indistinctly so, in males; indeed, even the blackish dots may be absent in males of A. strigatus and A. portunus. Male genitalia not unlike those of subgen. Allotinus, with valva ending in an apical hook but edges folded inwards more strongly in the form of a trough.
The species included in Fabitaras were placed in Paragerydus by earlier authors up to and including Fruhstorfer (1916) because of the swollen vein M3 of males and the common origin of veins M} and R5 in both sexes; but on the basis of the male genitalia and the frequent presence of a humeral vein it is certain that Fabitaras is more closely allied to Allotinus than to Paragerydus.
The subgenus is found from central Burma to Sundaland and the Philippines, and comprises ten species.
Key to the species of subgenus Fabitaras
1 Underside of hindwing with postdiscal spot in space 6 placed below, or just inside, that in space
7. Female with hindwing produced at vein M3 (faWus-group) 2
Underside of hindwing with postdiscal spot in space 6 placed well outside spot in space 7. Female
hindwing rounded (faras-group) 4
2 Underside of forewing with postdiscal series more or less parallel to termen except that spot in
space 5 is moved a little outwards. Postdiscal spots above vein M3 on forewing and vein M\ on
hindwing not enlarged nor blotchy 3
Underside of forewing with postdiscal series inclined towards apex; spots in spaces 4 and 5 moved out of line, the latter very close to termen. Postdiscal spots above vein M3 on forewing and vein MI on hindwing enlarged and blotchy fabius (p. 21)
3 cf upperside of forewing with vein M3 swollen for three-quarters of its length; visual brand
prominent. Underside of forewing with apical region shaded reddish brown; hindwing with submarginal black spot in space 6 at most barely larger than other submarginal spots
borneensis (p. 22)
Cf upperside of forewing with vein M3 swollen for half its length; visual brand narrow and obscure. Underside of forewing with apical region not shaded reddish brown; hindwing with submarginal black spot in space 6 triangular and larger than other submarginal spots
punctatus (p. 23)
4 Cf 9 upperside of hindwing with tornal area brown 5
Cf 9 upperside of hindwing with tornal quarter white brooksi (p. 26)
5 Underside of hindwing with submarginal black spot in space 6 at most barely larger than other
submarginal spots, cf upperside of forewing with at least indications of a visual brand astride
swollen portion of vein M3 6
Underside of hindwing with submarginal black spot in space 6 triangular and larger than other submarginal spots, cf upperside of forewing without a visual brand. 9 upperside of hindwing may have a brownish cream border nigritus (p. 24)
6 Underside of hindwing with postdiscal spot in space 6 placed roughly midway between the spots
in-spaces 7 and 5 7
Underside of hindwing with postdiscal spot in space 6 placed much further from spot in space 7 so that it is almost on an even arc with the spots in spaces 2 to 5 8
7 Underside ground colour white, cf forewing with vein M3 swollen for only one-third its length;
visual brand ill-defined and less than 1 -0 mm wide bidiensis (p. 26)
Underside ground colour pale buff, cf forewing with swollen portion of vein M3 about half its
length; visual brand about 2-0 mm wide strigatus (p. 24)
8 cf forewing with vein M3 swollen for more than half its length ; visual brand at least 1-0 mm wide 9
THE MILETINI 21
O" forewing with vein M3 swollen for half its length; visual brand very narrow, inconspicuous.
Underside ground colour greyish white ;forewing apex shaded reddish brown turns (p. 27)
9 Underside ground colour greyish white , not darkened towards forewing apex; black submargin- al spots outwardly white-edged; postdiscal series slightly closer to termen at dorsum than at costa. cf upperside of forewing with vein M3 swollen for two-thirds its length; visual brand up
to 2-0 mm wide, comparatively well defined sarrastes (p. 28)
- Underside ground colour in cf buff, becoming darker towards forewing apex; striation dense, black marginal spots absent or obscure, not white-edged. $ ground colour greyish white with forewing apex more or less shaded with brown; submarginal white-edged black spots present on forewing. Postdiscal series parallel to termen. cf upperside of forewing with vein M3 swollen for nearly four-fifths its length; visual brand ill-defined, just over 1-0 mm wide
portunus (p. 29)
Allotinus (Fabitaras) fabius (Distant & Pryer)
(Fig. 10, cf genitalia) Paragerydus fabius Distant & Pryer, 1887: 266.
The species is readily recognised by the underside markings. On the forewing the postdiscal spots in spaces 4 and especially 5 are shifted towards the termen, whilst the spots in spaces 6 and 7 are placed much further basad; in addition the spots above vein M3 are more or less enlarged and blotchy. On the hindwing the postdiscal spots in spaces 6 and 7 are placed one above the other and are large and blotchy. The female has the hindwing weakly caudate at vein M3. The male has vein M3 of the forewing swollen for about three-quarters of its length and the visual brand is about 1-5 mm wide.
The species has a restricted distribution in Sundaland, and has not been found in Java, Palawan nor the islands off the west coast of Sumatra. There are two subspecies.
Key to the subspecies of A. (F.) fabius
1 $ upperside of hindwing outwardly white fabius fabius (p. 21)
$ upperside of hindwing brown fabius arrius (p. 21)
Allotinus (Fabitaras) fabius fabius (Distant & Pryer)
Paragerydus fabius Distant & Pryer, 1887: 266; Cowan, 1966: 5. Holotype $ , BORNEO: Sandakan (BMNH)
[examined]. Allotinus caudatus Grose-Smith, 1893: 34. Holotype 'cf' recte $, BORNEO: Mt Kina Balu (BMNH)
[examined]. Syn. n.
Paragerydus caudatus (Grose-Smith); H. H. Druce, 1895: 563, pi. 31, figs 7, 8 cf - Allotinus fabius fabius (Distant & Pryer); Fruhstorfer, 1914: 22; 1916: 814; Corbet, 19396: 74. Allotinus fabius caudatus Smith; Fruhstorfer, 1914: 22; 1916: 814; Corbet, 19396: 74. Allotinus fabius pamisus Fruhstorfer, 1914: 22; 1916: 814; Corbet, 19396: 74 [misspelt pamiscus]. Holotype
$, BORNEO: south-east (BMNH) [examined]. Syn. n.
Fruhstorfer (1915) perpetuated Grose-Smith's mistake over the sex of caudatus by incorrectly stating that both sexes of the race from Kina Balu have the outer part of the hindwing white. In fact it is only females which have the hindwing partly white, and examples from Kina Balu do not differ significantly from those from the rest of Borneo nor from examples from south-west Sumatra.
DISTRIBUTION. Borneo, including Pulo Laut; south Sumatra (1 cf , 3 $ , Lebong Tandai (C. J. Brooks)).
Allotinus (Fabitaras) fabius arrius Fruhstorfer (Fig. 10, cf genitalia)
[Paragerydus panormis Elwes, 1893: 619 (partim), pi. 43, fig. 9 $. Misidentification.]
Allotinus fabius arrius Fruhstorfer, 1914: 22; 1916: 814, pi. 1411, cf $; Corbet, 19396: 74, fig. 4 cf genitalia;
Fleming, 1975: 21, pi. 58, fig. L45 cf; Eliot, 1978: 240. LECTOTYPE cf. SUMATRA (BMNH), here
designated [examined]. [Allotinus fabius panormis (Elwes); Fruhstorfer, 1916: 814. Misidentification.]
22
J. N. ELIOT
Fig. 10 Allotinus (Fabitaras) fabius arrius (Fruhstorfer); Malay Peninsula. Male genitalia.
Differs from the nominate subspecies only in the all brown female.
I designate as lectotype of arrius a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/CMB II. 94/Sumatra Monies Batlak ex coll. Fruhslorfer/Fruhstorfer Coll. B.M. 1933-131/fabius arrius Frhsl. [in Fruhslorfer's hand]/. A similarly labelled female is a paraleclolype.
DISTRIBUTION. Malay Peninsula; north Sumatra.
Allotinus (Fabitaras) borneensis Moulton (Figs 11 Cf genitalia; 58 cf)
Allotinus borneensis Moullon, 1911: 81; Fruhslorfer, 1913: 371; 1916: 814. Hololype cf , BORNEO: Sarawak
(BMNH) [examined].
Allotinus borneensis borneensis Moullon; Corbel, 19396: 76, fig. 3, cf genitalia. Allotinus borneensis elioti Corbet, 19396: 76; Fleming, 1975: 22, pi. 58, fig. L49 cf ; Eliol, 1978: 240.
Hololype cf , WEST MALAYSIA (BMNH) [examined]. Syn. n.
On Ihe upperside bolh sexes resemble A. fabius arrius, but are more reddish brown and, in Ihe male, Ihe
Fig. 11 Allotinus (Fabitaras) borneensis Moulton; Malay Peninsula. Male genitalia.
THE MILETINI
23
swollen portion of vein M3 is very slightly longer and the visual brand is very slightly wider. The underside is greyish white to pale buff-white, with the apical area of the forewing shaded with reddish brown. On the forewing the postdiscal series is parallel to the termen except that the spot in space 5 is shifted outwards, but not nearly as much as in A. fabius. On the hindwing the spot in space 6 is usually placed just inside that in space 7.
DISTRIBUTION. Borneo; Malay Peninsula; Sumatra; Bangka.
Allotinus (Fabitaras) punctatus (Semper) (Figs 12, cf genitalia; 59 9)
Paragerydus punctatus Semper, 1889: 165, pi. 31, figs 16 cf, 17 $. LECTOTYPE cf, PHILIPPINES: Mindanao (SM), here designated [examined].
Allotinus punctatus (Semper); Fruhstorfer, 1913: 371; 1916: 814; Corbet, 1939b: 74, fig. 1, cf genitalia.
Allotinus anaxandridas Fruhstorfer, 1916: 814. Holotype 9, PHILIPPINES: Mindanao (SM) [examined]. [$-morph.] Syn. n.
Allotinus caesemius Fruhstorfer, 1916: 814. LECTOTYPE $, PHILIPPINES: Mindanao (SM), here desig- nated [examined]. [$-morph.] Syn. n.
On the upperside the male has vein M3 swollen for half its length and the visual brand is rather narrow and ill-defined. The underside is pale buff with the postdiscal markings showing great individual variation in width, in some examples being as much as 2-0 mm wide. In females the ground colour is greyish white, and the postdiscal markings are much narrower, seldom as much as 1-0 mm in width. The white-edged black submarginal spots are well marked, that in space 6 of the hindwing being triangular and larger than the others. As in A. fabius and A. borneensis the postdiscal spots in spaces 7 and 6 of the hindwing are placed more or less one above the other, and the female has the hindwing weakly caudate at vein M3.
The species is chiefly remarkable for the polymorphism of the female, which was recognised by Semper who described three morphs. Fruhstorfer, however, considered that the three morphs were distinct species - a view for which there is no supporting evidence. The typical female is plain brown on both wings. In $-f. caesemius Fruhstorfer the forewing has a large white patch while the hindwing is brown; it seems to be the commonest morph. In $-f. anaxandridas Fruhstorfer the forewing resembles that of caesemius but the hindwing also has a large white patch; it appears to be much the rarest morph. Another 9 -morph was described by Fruhstorfer as 9-f- eretria, but I think that the butterfly in question was a female of A. nigritus (seep. 24).
Semper did not designate a type of punctatus, whilst Fruhstorfer named caesemius from two females and anaxandridas from a single female in Semper coll., which Semper had designated merely as varieties of punctatus. I now designate as lectotype of punctatus one of two males ex Semper coll. in SM labelled /Coll. C. Semper/Sibulan/215/Parag. punctatus typ. Semper/297c/Typus [red]/. A second male labelled /215/Coll. C. Semper/ is a paralectotype. I designate as lectotype of caesemius a female ex Semper coll. in SM labelled /Sibulan/297a/215/. The single female of anaxandridas is automatically the holotype and has been so labelled.
Fig. 12 Allotinus (Fabitaras) punctatus (Semper); Mindanao. Male genitalia.
24 J. N. ELIOT
DISTRIBUTION. The species has hitherto been recorded only from Mindanao. There is a single female in coll. Treadaway from Leyte (Catmon 450 m, 10. v. 1977) which differs from f. caesemius in having the white forewing patch reduced, with the dusky scaling at the wing base and above the dorsum almost as dark brown as the marginal and costal borders, whilst the underside is yellowish white with the markings heavier and more reddish than in females from Mindanao. It is probable that a distinct subspecies flies in Leyte, but I hesitate to name one on the basis of a single female.
Allotinus (Fabitaras) nigritus (Semper) (Figs 13, Cf genitalia, 60 $)
Paragerydus nigritus Semper, 1889: 164, pi. 31, fig. 15 cf . LECTOTYPE cf , PHILIPPINES: Mindanao (SM),
here designated [examined].
Allotinus nigritus (Semper); Fruhstorfer, 1914: 22; 1916: 814. Allotinus punctatus (Semper) $-f. eretria Fruhstorfer, 1916: 814. Holotype $. PHILIPPINES: Mindanao (not
located).
In the male vein M3 is swollen for a little under half its length and there is no visual brand. The underside is pale brownish ochreous with darker brown markings. The black submarginal spots resemble those of A. punctatus, from which A. nigritus can be separated by the postdiscal spot in space 6 of the hindwing being placed midway between the spots in spaces 7 and 5, or a little closer to the latter, as well as by its richer, more ochreous appearance.
The only female I have seen (Mindanao, Mt Apo, 15. ii. 1983, (A. Ballantine)) is dark brown above with an inwardly diffuse, sullied whitish border 2-0 mm wide on the hindwing from the tornus to vein R5. The hindwing termen is barely dentate at the end of vein M3, as in all species of the subgenus dealt with subsequently.
Fruhstorfer named as A. punctatus $ -f . eretria a female with the underside bright ochreous bearing thick brown markings and black marginal spots. The underside of females of A. punctatus is greyish white, so that I suspect that eretria really applies to A nigritus. Fruhstorfer did not mention the upperside of eretria, which was presumably unmarked brown, as in the typical female morph of A. punctatus, and if my supposition of the true identity of eretria is correct it would appear that the female of A. nigritus is dimorphic - with or without a whitish border on the hindwing.
I designate as lectotype of nigritus a male in SM labelled /Coll. C.Semper/Ost Mind./214/297b/15/Parag. nigritus typ. Semper/Typus [red]/. A second male labelled /Ost Mind./214/Typus [red]/is a paralectotype.
DISTRIBUTION. Mindanao.
Fig. 13 Allotinus (Fabitaras) nigritus (Semper); Mindanao. Male genitalia.
Allotinus (Fabitaras) strigatus Moulton
(Figs 14, cf genitalia; 61 cf ) Allotinus strigatus Moulton, 1911: 80. The species can be recognised by the fact that on the underside of the hindwing the postdiscal spot in space
THE MILETINI
25
6 is placed midway between those in spaces 7 and 5, while on the forewing the postdiscal series is much closer to the termen near the tornus than near the costa. The underside ground colour is a uniform pale buff. On the upperside the male has vein A/3 swollen for half its length, and the visual brand is quadrate, about 2 mm wide and not sharply outlined.
The species is strictly Sundanian, and has not yet been found in Java, Palawan or the islands off the west coast of Sumatra. There are two subspecies.
Key to the subspecies of A. (F.) strigatus
1 On the underside of the forewing the blackish submarginal dots are inconspicuous and not
outwardly white-edged strigatus strigaius (p. 25)
- On the underside of the forewing the blackish submarginal dots are outwardly white-edged in $
and in the apical half of the wing in cf strigatus malayanus (p. 25)
Allotinus (Fabitaras) strigatus strigatus Moulton (Fig. 61 cf)
Allotinus strigatus Moulton, 1911: 80. Holotype cf , BORNEO: Pulo Laut (BMNH) [examined]. Allotinus strigatus strigatus Moulton; Fruhstorfer, 1914: 22; 1916: 813; Corbet, 1939ft: 75, fig. 5, cf genitalia.
On the underside the submarginal blackish dots are not white-edged, and the usual markings are comparatively broad and well defined.
DISTRIBUTION. Borneo, including Pulo Laut.
Allotinus (Fabitaras) strigatus malayanus Corbet (Fig. 14, cf genitalia)
Allotinus strigatus malayanus Corbet, 1939ft: 75; Fleming, 1975: 22, pi. 58, fig. L48 cf ; Eliot, 1978: 240.
Holotype cf , WEST MALAYSIA (BMNH) [examined]. Allotinus strigatus denalus Corbet, 1939ft: 75. Holotype cf , SUMATRA: Battak Mts (BMNH) [examined].
Syn. n.
On the underside of the forewing the blackish submarginal dots are outwardly edged with whitish in the female, but in the male less distinctly and only in the apical half of the wing. The markings are narrower and less well defined than in the nominate subspecies. In the male the swelling of vein M3 is slightly shorter, being just under half its length, and the visual brand is usually a little narrower.
DISTRIBUTION. Malay Peninsula, including Singapore; Sumatra.
Fig. 14 Allotinus (Fabitaras) strigatus malayanus Corbet; Malay Peninsula. Male genitalia.
26
J. N. ELIOT
Allotinus (Fabitaras) brooksisp. n.
(Figs 15, cf genitalia; 63 cf )
Cf forewing length 15-0 mm. Venation normal for the subgenus, with veins M j and R5 of forewing having a moderately long common stalk. Upperside brown, with tornal quarter of hindwing white, very sparsely dusted with brown scales and with adjoining cilia white chequered with dark brown at vein endings. Forewing with vein M3 swollen for slightly less than one-third of its length; visual brand very narrow, obscure. Underside very pale greyish white, sparsely striated; postdiscal spots small , more or less parallel to termen; hindwing with postdiscal spot in space 6 a little closer to that in space 7 than to that in space 5; cell-end bars unusually heavy, especially on hindwing; forewing with submarginal blackish dots outwardly white-edged. Genitalia similar to those of A. strigatus, A. taras and A. sarrastes, but apical process of valva slightly larger.
$ forewing length 15-0 mm. Differs from male in that inwardly diffuse white area on hindwing a little larger, occupying one-third of wing. At first sight this sex might be mistaken for A. fabius fabius , but the rounded hindwing termen separates it readily.
MATERIAL EXAMINED
Holotype cf , Borneo: 'Bau Feb Feb 10', 'Sarawak C. J. Brooks', '192', 'C. J. Brooks Bequest. B.M. 1953-173' (BMNH).
Paratype. Borneo: 1 $ (allotype), 'Bau Dec. 09', '192', 'C. J. Brooks Bequest B.M. 1953-173' (BMNH).
Fig. 15 Allotinus (Fabitaras) brooksi sp. n. ; Borneo. Male genitalia.
Allotinus (Fabitaras) bidiensissp. n.
(Figs 16, cf genitalia; 62 cf )
Cf forewing length 15-0-16-0 mm. In the two examples examined forewing venation differs from rest of subgenus in that veins MI and R5 are just separate at their origins; vein /?3 rather short, arising just before end of vein R2. Upperside brown, with basal third of vein M3 swollen; visual brand narrow, obscure. Underside with markings generally arranged as in sympatric A. strigatus, but differing in the pale greyish white ground colour, the smaller, more rounded postdiscal spots and the prominently white-edged blackish submarginal dots on the forewing. Male genitalia distinguished by valva, in which the lower process extends beyond the short apical hook, and comparatively short, stout phallus.
9 forewing length 14-0 mm in the single example seen, wherein veins MI and R5 of forewing connate from cell apex. Upperside brown. Underside similar to male.
MATERIAL EXAMINED
Holotype cf , Borneo: Sarawak, '647 20/5/08', '190f , 'C. J. Brooks Bequest. B.M. 1953-173' (BMNH).
Paratypes. Borneo: 1 $ (allotype), Sabah, 'N. Born.', 'Madai 3.2.92', 'Joicey Bequest. Brit. Mus. 1934-120' (BMNH); cf , 'Bidi Sarawak 1907 C. J. Brooks', '190f , 'C. J. Brooks Bequest. B.M. 1953-173' (BMNH).
THE MILETINI
27
Fig. 16 Allotinus (Fabitaras) bidiensis sp. n. ; Borneo. Male genitalia.
Allotinus (Fabitaras) taras (Doherty)
(Fig. 17, Cf genitalia)
Paragerydus taras Doherty, 1889: 437, pi. 23, fig. 10 cf . Syntypes, BURMA: Tenasserim (not located). Parageryduspanormis Elwes, 1893: 619, partim 'cf ' recte £ , nee $ , pi. 43, fig 8 'cf '. Holotype 'cf ' recte $ ,
BURMA: East Pegu (BMNH) [examined]. Syn. n. Allotinus taras (Doherty) Bingham, 1907: 300; Swinhoe, 1910: 199, pi. 617, figs 2, 2b cf , 2a, 2c $; Evans,
1932: 212; Cantlie, 1963: 27. Allotinus panormis (Elwes) Bingham, 1907: 301; Swinhoe, 1910: 197 partim, pi. 616, figs 3, 3b 'cf' recte $,
nee $.
Allotinus taras taras (Doherty); Fruhstorfer, 1913: 370; 1916: 813; Corbet, 19396: 74, partim. Allotinus fabius panormis (Elwes); Fruhstorfer, 1916: 814, partim; Evans, 1932: 212. Allotinus panormis panormis (Elwes); Corbet, 19396: 74; Cantlie, 1963: 27.
On the upper surface of the forewing the male has vein M3 swollen for half its length, and the visual brand is very narrow and inconspicuous; indeed Doherty, in his original description, gave absence of the brand as
Fig. 17 Allotinus (Fabitaras) taras (Doherty); Burma. Male genitalia.
28
J. N. ELIOT
one of the characters of the species. The underside of both sexes is greyish white turning to reddish brown towards the forewing apex and, occasionally, at the hindwing apex also. The postdiscal series of spots is rather lightly marked, and though these spots were mentioned by Doherty they are wanting in the example chosen for his figure. Such examples cannot be regarded as typical, only two out of 28 males in BMNH being similar to Doherty's figure.
The 'male' (recte female) of Paragerydus panormis is a quite typical female of A taras, and I am at a loss to understand why Corbet regarded it as a different species conspecific with A. portunus.
The male genitalia of A. taras and A. sarrastes are similar, so there are grounds for regarding these two taxa as conspecific. However, they overlap in south Burma between Tavoy and Mergui over a distance of some 160 km without any evidence of intergradation, so that it seems highly probable that their period of isolation, when there was a sea barrier in the region of the Isthmus of Kra, was sufficiently prolonged for interbreeding to be impossible when they again met.
I have seen none of Doherty's type-series and as the figured male is atypical I do not designate it as lectotype.
DISTRIBUTION. Burma, from the Karen Hills to Mergui.
Allotinus (Fabitaras) sarrastes Fruhstorfer stat. n. (Fig. 18, cf genitalia)
Allotinus taras sarrastes Fruhstorfer, 1913: 370; 1916: 813; Corbet, 19396: 74; Eliot, 1961: 71; 1978: 240;
Fleming, 1975: 21, pi. 58, fig. L46 cf . LECTOTYPE cf , BORNEO (BMNH), here designated [examined]. [Allotinus taras battakanus Fruhstorfer, 1913: 370, partim; 1916: 813 partim, ? pi. 141g $. Misidentifica-
tion.]
Allotinus porriginosus Toxopeus, 1932: Ixxvii. Holotype cf , JAVA: south (not located). Syn. n. [Allotinus taras taras (Doherty); Corbet, 19396: 74, fig. 2, cf genitalia. Misidentification.] Allotinus taras mendava Riley, 1944: 253, pi. 2, fig. 30 cf, 31 $. Holotype cf, MENTAWAI Is.: Sipora
(BMNH) [examined]. Syn. n.
This Sundanian species is obviously very closely related to A. taras, but it is best regarded as specifically distinct. It differs from A. taras on the underside by the absence of reddish brown shading towards the forewing apex. In addition, on the upperside of the male forewing the swelling of vein M3 extends over about two-thirds of its length and the visual brand is prominent and usually about 1-5 mm wide. On the underside the white edges to the blackish submarginal spots are particularly well marked, especially in the female.
I designate as lectotype a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/Kina Balu ex coll. H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/sarrastes Frhst. [in Fruhstorfer's hand]/. A female labelled /Type [red]/Type [Fruhstorfer orange]/Nord-Borneo Brunei Waterstradt 1890/Fruhstorfer Coll. B.M. 1933-131/sarrastes Frhst. [in Fruhstorfer's hand]/ is a paralectotype.
Fig. 18 Allotinus (Fabitaras) sarrastes Fruhstorfer; Malay Peninsula. Male genitalia.
THE MILETINI 29
DISTRIBUTION. Burma, from Tavoy southwards; Malay Peninsula; Sumatra; Mentawai Is.; Borneo; Java; Mindanao (SM, 1 cf labelled /Ost Mind./298a/213/Coll. C. Semper/re verdini/; this specimen has the visual brand obscure and ill-defined, but this may be due partly to its rubbed condition.)
Allotinus (Fabitaras) portunus (de Niceville)
(Fig. 19, cf genitalia) Paragerydus portunus de Niceville, 1894: 27.
Males are easily recognised by the long swelling of vein M3 of the forewing, extending over nearly four-fifths of its length, and the rather diffuse and narrow visual brand, as well as by the buff or pale rufous underside on which the usual blackish submarginal spots are absent or, if present, are not outwardly white-edged. Females most nearly resemble that sex of A. taras on the underside, but the subapical darkening on the forewing is less developed and brownish ochreous without the red tinge of taras. In both sexes the postdiscal series of spots on the forewing is more exactly parallel to the termen than in the other species of the subgenus.
As already pointed out (p. 28) Corbet (19396) subordinated A. portunus and its subspecies under A. panormis, which is a synonym of the Burmese species A. taras (Doherty).
The species is purely Sundanian, not extending to Palawan or the islands off the west coast of Sumatra, and, at least in the Malay Peninsula, flies at higher average elevations than the other species of the subgenus, being seldom found below 800 m.
I recognise three weak subspecies based on a mean of differences.
Key to the subspecies of A. (F.) portunus
1 cf underside of forewing with blackish submarginal spots normally absent 2
- cf underside of forewing with blackish submarginal spots normally present... portunus pyxus (p. 30)
2 cf underside of forewing with postdiscal series ill-marked. $ underside usually pale greyish buff,
with forewing apex shaded with ochreous portunus portunus (p. 29)
Cf underside of forewing with postdiscal series narrow but usually clearly defined. $ underside greyish white with forewing apex only narrowly and faintly shaded portunus maitus (p. 29)
Allotinus (Fabitaras) portunus portunus (de Niceville)
Paragerydus portunus de Niceville, 1894: 27, pi. 5, fig. 14 cf . Syntypes, JAVA (?ZSI).
Allotinus taras narsares Fruhstorfer, 1913: 370; 1916: 813; Corbet, 19396: 74, partim. LECTOTYPE $,
JAVA (BMNH), here designated [examined]. Syn. n.
Allotinus portunus portunus (de Niceville) Fruhstorfer, 1914: 22; 1916: 813. [Allotinus strigatus dositheus Fruhstorfer, 1914: 22 partim, $ nee cf .] [Allotinus taras (Doherty); Piepers & Snellen, 1918: 14, pi. 19, fig. 16 $. Misidentification.] Allotinus portunus (de Niceville); Piepers & Snellen, 1918: 15, pi. 20, figs 18a cf , 18b $. Allotinus panormis portunus (de Niceville); Corbet, 19396: 75.
On the underside the ground colour is variable in both sexes. In the male it varies from pale buff to rufous buff, and generally the postdiscal markings are ill-defined and may be absent as in the example figured by de Niceville. In the female the ground colour may be as buff as in the male, but in some examples, as in the lectotype of narsares, which apparently was the model for the figure of A. taras in Piepers & Snellen (1918), it is pale greyish. The blackish submarginal spots on the underside of the forewing are usually missing in the male but present in the female and at most only weakly edged with white.
I designate as lectotype of narsares a female in BMNH labelled /Type [red]/Java Occident. Sukabumi 2000' ex coll. H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/narsares Frhst. [in Fruhstorfer's hand]/.
DISTRIBUTION. Java.
Allotinus (Fabitaras) portunus maitus Fruhstorfer (Fig. 19, Cf genitalia)
Allotinus portunus maitus Fruhstorfer, 1914: 21; 1916: 813. LECTOTYPE cf, SUMATRA (BMNH), here designated [examined].
30 J. N. ELIOT
Fig. 19 Allotinus (Fabitaras) portunus maitus Fruhstorfer; Malay Peninsula. Male genitalia.
Allotinus panormis fruhstorferi Corbet, 19396: 74, fig. 7, cf genitalia; Fleming, 1975: 22, pi. 58, fig. L47 $ ;
Eliot, 1978: 240. Holotype cf , WEST MALAYSIA (BMNH) [examined]. Syn. n. Allotinus panormis maitus Fruhstorfer; Corbet, 19396: 74.
In the male the postdiscal markings on the underside are usually more clearly defined than in subsp. portunus and the submarginal blackish spots are missing; in the female the ground colour is more greyish white.
I designate as lectotype of maitus a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/CMB 111. 94/Fruhstorfer Coll. B.M. 1933-131/portunus maitus Frhst. [in Fruhstorfer's hand]/. A female labelled /Type [red]/Type [Fruhstorfer orange]/Sumatra Monies Battak ex coll. H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/allotype Allotinus portunus maitus Fruh. [in Corbet's hand]/ is a paralectotype.
DISTRIBUTION. Sumatra; West Malaysia.
Allotinus (Fabitaras) portunus pyxus (de Niceville)
Paragerydus pyxus de Niceville, 1894: 27, pi. 5, fig. 2 cf . Holotype cf , BORNEO: Kina Balu (? ZSI). Paragerydus waterstradti H. H. Druce, 1895: 562, pi. 31, figs 1 cf, 2 $. LECTOTYPE cf, BORNEO
(BMNH), here designated [examined]. [Synonymised by Fruhstorfer, 1914: 22.] Paragerydus waterstradti ab. absens H. H. Druce, 1895: 562.
Allotinus portunus pyxus (de Niceville); Fruhstorfer, 1914: 22; 1916: 813, pi. 141i, cf $. Allotinus panormis pyxus (de Niceville); Corbet, 19396: 75.
Females appear to be inseparable from the nominate subspecies but in males the blackish submarginal spots on the underside of the fore wing are more often present.
De Niceville, who named the subspecies from a single male, differentiated it from subsp. portunus by the more rufous tone of the upperside and the pale rufous instead of pale ochreous colour of the underside. However, the ground colour is very variable in all three subspecies and is not a reliable character. De Niceville's figure is misleading, as it does not show the visual brand which is mentioned in his description.
H. H. Druce described waterstradti from syntypes in coll. Staudinger and in his own collection. I designate as lectotype a male in BMNH labelled /Kina Balu Waterstr./P. waterstradti co-type H. H. Druce/ex coll. Hamilton Druce/Joicey Bequest Brit. Mus. 1934-120/. As there is no specimen in BMNH of ab. absens H. H. Druce, it is likely that the name applies to a specimen in coll. Staudinger.
DISTRIBUTION. Borneo, apparently only known from Mt Kina Balu.
Subgenus PARAGERYDUS Distant
Paragerydus Distant, 1884: 207. Type-species: Miletus horsfieldi Moore, 1857: 19, pi. la, fig. 1, by
designation of Kirby, [1885]: 191. Miletographa Rober, 1892: 277. Type-species: Miletus drumila Moore, [1866]: 777, pi. 41, fig. 12, by
monotypy. [Synonymised by Fruhstorfer, 1913: 371.]
THE MILETINI 31
Antenna a little over half length of costa, slightly longer than in subgen. Allotinus and shorter than in subgen. Fabitaras. Antennal segments number about 40 to 60. Shaft segments of the smaller species relatively longer, consequently fewer in number; thus in the nivalis-group and in A. corbeti there are about 40 segments, whereas in the larger species, such as A. horsfieldi and A. apries, there are about 58 segments. Species of intermediate size have an intermediate number of segments, for example in A. unlcolor there are usually about 45. In dwarf individuals, occurring most frequently in A. horsfieldi and A. unicolor, there is no diminution in the number of segments. Venation shows less individual variation than in Fabitaras. Hindwing without humeral vein. Forewing veins MI and R5 usually connate or briefly stalked, but in the nivalis-group they may be just separate at their origins. In males, vein M3 of forewing basally swollen, clothed with small specialised scales, those of the nivalis-group being relatively large. In males, abdominal hair tufts prominent, permanently extruded and, in the genitalia, the valva has a truncate apex, except in A. davidis, and its ventral edge prolonged into a more or less pointed process. With only a few exceptions, the interspecific differences in the valva are very slight and perhaps inconstant; in general the genitalia give little assistance in identification.
Early authors, up to Fruhstorfer (1913; 1916), used Paragerydus in a wider sense, either as a genus or subgenus (Artengruppe) to include the species here placed in Fabitaras.
The subgenus ranges from north India to Sundaland, the Lesser Sunda Is., Philippines and Sulawesi (including Sula Is.). It comprises 16 species.
Key to the species of subgenus Paragerydus
1 Underside mottled with brown specks and striae; postdiscal series not outlined by darker lines
nor catenulate. Smaller, forewing 9-0-23-0 mm 2
- Underside mottled with brown spots ringed with pale buff; postdiscal series catenulate and
outlined by darker lines. Larger, forewing usually over 23-0 mm drum Ha (p. 56)
2 Underside of forewing without a white fleck at end of vein R5 (horsfieldi-group) 3
- Underside of forewing with a white fleck at end of vein R5 (nivalis-group) 14
3 Underside with postdiscal series more or less the same size throughout. Upperside of hindwing
unmarked brown 4
Underside with postdiscal spots below vein M3 on forewing much larger than those above, and spots above vein MI on hindwing much larger than those below. Upperside of hindwing partly white in cf $ or in $ alone 13
4 Upperside of forewing brown; white patch or streak present in 9 of two species. Hindwing
brown, of same shade as forewing; termen crenulate (except in A . apries) 5
Upperside of forewing white to wing base, with dark brown margin and costa. Hindwing uniform pale buff-brown; termen not crenulate parapus (p. 51)
5 $ upperside brown, cf with visual brand, if present, not extending into cell basad of origin of
vein Cui 6
- $ upperside of forewing with white patch or streak, cf (where known) with visual brand
extending obscurely into cell basad of origin of vein Cu2 samarensis (p. 37)
6 Underside of forewing with postdiscal spots in spaces 4, 5 and 6 more or less in line and
equidistant , or with the spot in space 5 nearer to , and often touching, the spot in space 6 7
Underside of forewing with postdiscal spots in spaces 4, 5 and 6 on an uneven curve, that in space 5 overlapping that in space 4 and well separated from that in space 6 macassarensis (p. 39)
7 Forewing veins MI and R5 connate or with a common stalk not more than 1-0 mm long.
Underside of forewing with postdiscal spot in space 2 nearer termen than the spots in spaces 3 and Ib (if present), cf $ with hindwing cilia elongated into tufts at vein endings, termen crenulate in $
- Forewing veins MI and R5 with a stalk more than 1-0 mm long. Underside of forewing with
postdiscal spots in spaces 3, 2 and Ib (if present) in line and parallel to termen. cf $ with hindwing cilia barely longer at vein endings and termen not crenulate in $ apries (p. 42)
8 cf upperside of forewing with a visual brand , vein M3 swollen for about half its length or longer 9 Cf upperside of forewing without a visual brand, vein M3 swollen for only one-quarter of its
length. Small, forewing 10-0-12-0 mm corbeti (p. 44)
9 Forewing veins Ml and R5 usually with a very short common stalk, cf valva with terminal
process not rising above costa ; tip of uncus not , or only a little , produced 10
- Forewing veins M1 and R5 usually connate, cf valva with terminal process curved upwards,
ending just above costa; tip of uncus produced , rostriform horsfieldi (p. 32)
32 J. N. ELIOT
10 cf forewing with vein A/3 swollen for at least three-fifths of its length; visual brand comparative-
ly wide 11
- cf forewing with vein M3 swollen for only half its length or slightly over; visual brand comparatively narrow, less than 1-5 mm wide (except in subspp. continentalis , and moorei sometimes, of A. unlcolor) 12
11 cf valva with terminal process comparatively long and narrow (Fig. 21). Underside of
sympatric taxa greyish white; on hindwing postdiscal spot in space 6 usually more or less
below that in space 7 leogoron (p. 34)
Cf valva with terminal process short and broad (Fig. 22). Underside ground colour very pale buff; on hindwing postdiscal spot in space 6 nearly always inside the spot in space 7 and sometimes nearly mid-way to the end-cell bar melos (p. 36)
12 Underside ground colour greyish white to pale buff. Comparatively small, with forewing
10-5-18-0 mm. cf valva (Figs 29, 30) comparatively broad, with terminal process in centre
line unicolor (p. 45)
Underside chalky white. Larger, forewing 18-0-20-0 mm. cf valva narrower (Fig. 31), with terminal process nearly in line with ventral edge paetus (p. 50)
13 cf upperside greyish brown. $ with a whitish discal patch on forewing and an obscure whitish
streak in space 5 on hindwing. cf 9 discocellular veins on hindwing not darkend
luzonensis (p. 40) Cf $ upperside partly white on both wings. Discocellular veins blackened albatus (p. 41)
14 Underside of hindwing with central spot in space 7 at least partly blackened. Smaller, forewing
9-0-14-5 mm 15
Underside of hindwing with central spot in space 7 not blackened (may be darker brown than other spots). Larger, forewing 12-5-15-5 mm nivalis (p. 51)
15 cf valva with costa incised shortly before apex (as in all preceding species). Underside of
hindwing with postdiscal spot in space 7 not blackened on its inner edge, except sometimes in the dry season form in Burma; ground colour greyish white, except in Philippines where it is
pale buff substrigosus (p. 53)
Cf valva with costa entire. Underside of hindwing with postdiscal spot in space 7 blackened on its inner edge; ground colour pale buff. Not found in Burma or Philippines davidis (p. 55)
Allotinus (Paragerydus) hors field! (Moore)
(Fig. 20, cf genitalia) Miletus horsfieldi Moore, 1857: 19.
This and three succeeding species, leogoron, melos and macassarensis , form a confusing group difficult to separate by superficial characters. In the males of all four species the swelling of vein M3 and the visual brand on the forewing vary according to locality. Venation is individually variable, but in A. horsfieldi veins MI and R5 of the forewing are nearly always connate , whereas in the other three species they usually share a very short common stalk. In both sexes the hindwing cilia are elongated into short tufts at the vein endings and females have the hindwing termen crenulate. In the female of A. horsfieldi the hindwing is particularly strongly crenulate. In A. leogoron the crenulations are much less pronounced, while in A. melos and A. macassarensis they are intermediate. On the underside the markings show much individual variability in their density and in the position of the postdiscal series. On the forewing this series is dislocated at vein M3, with the stria in space 3 moved basad and often forming with the stria in space 2 an irregular, oblique stripe; but the degree of dislocation is variable and usually slight in A. macassarensis. In addition the stria in space Ib, if present, is moved basad in relation to that in space 2.
Males of A. horsfieldi can always be identified with certainty by the genitalia (Fig. 20), wherein the ventral tip of the uncus is much produced and the terminal process of the valva is long and curved up above the costa.
In both sexes A. horsfieldi shows great variability in size. The smaller males, with forewing length as little as 14 mm, have the forewing apex and termen more rounded and the visual brand relatively narrow, but they are connected by a complete range of intermediates to the largest males, with forewing length up to 23 mm, which have the forewing apex more pointed and the termen straighter.
A. horsfieldi has a restricted distribution confined to Sundaland excluding Palawan. Where it occurs it is usually the commonest species of the subgenus apart from A. unicolor.
THE MILETINI
33
Key to the subspecies of A. (P.) horsfieldi
1 Underside pale buff, postdiscal series not quadrate 2
- Underside greyish white, postdiscal series comprising large, dark, quadrate spots
horsfieldi siporanus (p. 34)
2 Upperside brown without a reddish tinge . Underside usually lightly marked 3
- Upperside brown with a reddish tinge , especially strong in 9 and in basal half of forewing in cf .
Underside more strongly marked horsfieldi permagnus (p. 33)
3 cf with visual brand broad, for about half its length contiguous with vein Cu\. 9 with forewing
disc not conspicuously paler horsfieldi horsfieldi (p. 33)
- cf with visual brand narrower, only its basal quarter touching vein Cu\. 9 with forewing disc
conspicuously paler horsfieldi satelliticus (p. 34)
Allotinus (Paragerydus) horsfieldi horsfieldi (Moore)
Miletus horsfieldi Moore, 1857: 19, pi. la, fig. 2 cf . LECTOTYPE cf , JAVA (BMNH), here designated
[examined].
Allotinus horsfieldi horsfieldi (Moore) Fruhstorfer, 1913: 367; 1916: 812; Corbet, 1939ft: 73. Allotinus horsfieldii [sic] (Moore); Piepers & Snellen, 1918: 12, pi. 19, figs 12a Cf , 12b 9, 14a cf , 14b $.
In the male the visual brand is broad, and touches vein Cu^ for about half its length. The female is dull brown without a reddish tinge, and the forewing disc is only a little paler. On the underside, especially in females, the usual lycaenid markings are generally light and may be faded out, as in the extremes shown in Piepers & Snellen (1918: pi. 19, figs 14a, 14b), and on the hindwing the postdiscal spot in space 6 is usually placed well inside the spot in space 7.
I designate as lectotype a male in BMNH labelled /Type [red]/60-15 E. E.G. /Miletus horsfieldi cf M/GENITALIA Slide No ASC 23 Allotinus/. A female labelled /Type [red]/60-15 E.E.C./ is a paralecto- type.
DISTRIBUTION. Java.
Allotinus (Paragerydus) horsfieldi permagnus Fruhstorfer
(Fig. 20, cf genitalia) Paragerydus horsfieldi (Moore); Distant, 1884: 207, pi. 20, fig. 7 '9' recte cf .
Fig. 20 Allotinus (Paragerydus) horsfieldi permagnus Fruhstorfer; Rhio Archipelago: Great Karimon I. Male genitalia.
34 J. N. ELIOT
Allotinus horsfieldi permagnus Fruhstorfer, 1913: 366; 1916: 812. LECTOTYPE d", SUMATRA (BMNH),
here designated [examined]. Allotinus horsfieldi nessus Corbet, 19396: 72, fig. 15, cf genitalia; Eliot, 1978: 240. Holotype cf , WEST
MALAYSIA (BMNH) [examined]. [Synonymised by Eliot, 1967: 66.] [Allotinus leogoron lindus Corbet, 19396: 73, partim $ nee d*. Misidentification.] Allotinus horsfieldi permagnus Fruhstorfer; Eliot, 1967: 66, fig. 1, cf genitalia; Fleming, 1975: 21, pi. 58,
fig. L43 O".
The subspecies is best distinguished by the female which is reddish brown with the forewing disc only very slightly paler. The female named by Fruhstorfer as f . infumata is well within the normal range of individual variation. In the male the visual brand is comparatively narrow, only touching vein Cui at its base, and the wing base of the forewing has a more reddish tinge than the darker apical region.
I designate as lectotype of permagnus a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/W. Sumatra H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/horsfieldi permagnus Fruhst. [in Fruhstorfer's hand]/. The specimen, which is rather more lightly marked on the underside than usual, has lost its abdomen. A female labelled /Type [red]/Sumatra Montes Battak ex coll. H. Fruhstorfer/^ horsfieldi Selesseh 19. xii. 94/Fruhstorfer Coll. B.M. 1933-131/horsfieldi permagnus Fruhst. [in Fruhstor- fer's hand]/ is a paralectotype.
I designate as lectotype of infumata a female in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/N. Oekor viii. 94/Fruhstorfer Coll. B.M. 1933-131/infumata Frhst. [in Fruhstorfer's hand]/.
DISTRIBUTION. Malay Peninsula, including Singapore; Sumatra; Bangka; Borneo; Peninsular Thailand (Pinratana). Three females in BMNH from Batu Is. are provisionally included in this subspecies; they are small with the underside ground colour whiter and the postdiscal markings rather dark and heavy, showing an approach to subsp. siporanus. They are also rather similar to A. leogoron batuensis, with which Eliot (1967) originally confused them and from which they can be distinguished by their more crenulate hindwingtermen.
Allotinus (Paragerydus) horsfieldi siporanus Riley
Allotinus horsfieldi siporana Riley, 1944: 253. Holotype $. MENTAWAI Is.: Sipora (BMNH) [examined].
The subspecies was described from a single large female in extremely battered condition; the whole of the outer half of the left hindwing and the tornal quarter of the right hindwing below vein Cu\ are missing. The greyish white ground colour of the underside and heavy dark brown markings suggest that it might pertain to A. leogoron, but such crenulations as remain on the termen of the right hindwing suggest A. horsfieldi, and it seems best to leave Riley's combination unchanged pending the discovery of the male.
DISTRIBUTION. Mentawai Is.
Allotinus (Paragerydus) horsfieldi satelliticus Fruhstorfer
Allotinus horsfieldi satelliticus Fruhstorfer, 1913: 366; 1916: 812; Eliot, 1967: 66. LECTOTYPE $, ENGANO I. (BMNH), here designated [examined].
In the male the visual brand is like that of subsp. permagnus, but in other respects the subspecies more nearly resembles the nominate subspecies, especially in the female which is without a reddish tinge on the upperside. However, this sex differs from Javanese females by having a very prominent paler discal patch on the forewing.
I designate as lectotype a female in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/Engano April-Juli Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/horsfieldi satelliticus Frhst. [in Fruhstorfer's hand]/.
DISTRIBUTION. Engano I.
Allotinus (Paragerydus) leogoron Fruhstorfer
(Fig. 21, cf genitalia) Allotinus leogoron Fruhstorfer, 1916: 811.
This species has been much confused in the past with A. horsfieldi, from which the male is readily separated by the genitalia, wherein the ventral tip of the uncus is not produced and the terminal process of the valva is
THE MILETINI
35
fairly long and narrow but not upturned. Females are best distinguished by the hindwing termen, which is less crenulate than in horsfieldi, but the cilia bear slightly longer and narrower tufts at the vein endings. On the forewing veins MI and R5 usually have a short common stalk and, except in Java, the ground colour of the underside is whiter with darker, more contrasted markings than in horsfieldi. A. leogoron has the same distribution as A. horsfieldi. I recognise four subspecies.
Key to the subspecies of A. (P.) leogoron
1 cf visual brand narrow, only its base touching vein C«i 2
- cf visual brand wide, touching vein Cu\ throughout half its length leogoron leogoron (p. 35)
2 Underside greyish white with dark markings 3
- Underside pale buff with markings smaller and paler brown leogoron plessis (p. 36)
3 d" visual brand with only its extreme base touching vein Cu\. Underside of hindwing with
postdiscal spot in space 6 below that in space 7, as in the nominate subspecies
leogoron nor muni (p. 36)
- cf visual brand a little wider, touching vein Cu\ for a quarter of its length . Underside of hindwing
with postdiscal spot in space 6 placed inside that in space 7, sometimes almost half-way to cell-end bar leogoron batuensis (p. 36)
Allotinus (Paragerydus) leogoron leogoron Fruhstorfer
(Fig. 21, cf genitalia)
Allotinus horsfieldi permagnus $-f. intricata Fruhstorfer, 1913: 366; 1916: 812. LECTOTYPE $, SUMATRA
(BMNH), here designated [examined]. [Unavailable name.] [Synonymised by Eliot, 1967: 68.] Allotinus leogoron Fruhstorfer, 1916: 811. LECTOTYPE cf, SUMATRA (BMNH), here designated
[examined]. Allotinus continental vadosus Corbet, 19396: 72. Holotype cf , WEST MALAYSIA (BMNH) [examined].
[Synonymised by Eliot, 1967: 69.] Allotinus leogoron leogoron Fruhstorfer; Corbet, 19396: 73; Eliot, 1967: 68, fig. 5, cf genitalia; 1978: 240;
Fleming, 1975: 21, pi. 58, fig. L44 cf . Allotinus leogoron lindus Corbet, 19396: 73, partim cf nee $. Holotype cf, WEST MALAYSIA (BMNH)
[examined]. [Synonymised by Eliot, 1967: 69.]
The male can be separated from that sex of sympatric A. horsfieldi by the wider, more sharply defined visual brand which touches vein Cu\ throughout its basal half, and the female by the less crenulate hindwing termen. In addition, the underside is whiter and the markings are darker brown.
Fig. 21 Allotinus (Paragerydus) leogoron leogoron Fruhstorfer; Malay Peninsula. Male genitalia.
36 J. N. ELIOT
I designate as lectotype of leogoron a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ CMB viii. 94/A. leogoron Fr. [in Fruhstorfer's hand]/ and of intricata a female in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/Sumatra Montes Battak ex coll. H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/$ horsfieldi CMB viii. 94/intricata [in Fruhstorfer's hand]/.
DISTRIBUTION. Malay Peninsula; Sumatra; Bangka; Peninsular Thailand (Pinratana).
Allotinus (Paragerydus) leogoron normani Eliot Allotinus leogoron normani Eliot, 1967: 69. Holotype cf , BORNEO (BMNH) [examined].
Differs from the nominate subspecies only in that the male has the visual brand much narrower, only touching vein Cui at its extreme base.
DISTRIBUTION. Borneo.
Allotinus (Paragerydus) leogoron batuensis Eliot Allotinus leogoron batuensis Eliot, 1967: 69. Holotype cf , BATU Is. (BMNH) [examined].
A small subspecies, with a slightly more reddish brown tinge than in the two preceding subspecies. In the male the visual brand is intermediate between that of leogoron and that of normani, touching vein Cui along its basal quarter. It differs on the underside of the hindwing in that the postdiscal spot in space 6 is placed inside the spot in space 7.
DISTRIBUTION. Batu Is.
Allotinus (Paragerydus) leogoron plessis Eliot
Allotinus leogoron plessis Eliot, 1967: 69. Holotype cf , JAVA (BMNH) [examined].
In the male the visual brand is narrow, as in subsp. normani, whereas the brand of sympatric A horsfieldi horsfieldi is as wide as in subsp. leogoron. The subspecies is further distinguished by the pale buff underside on which the markings are smaller and less contrasted.
DISTRIBUTION. Java.
Allotinus (Paragerydus) melos (H. H. Druce)
(Figs 22, cf genitalia; 64 cf )
[Paragerydus horsfieldi (Moore) sensu Semper, 1889: 164 partim, pi. 31, figs 19 cf, 20 $. Misidentifica-
tion.] Paragerydus melos H. H. Druce, 1896: 652. LECTOTYPE cf , PHILIPPINES: Cagayan Sulu (BMNH), here
designated [examined]
Allotinus melos (Druce) Fruhstorfer, 1913: 369. Allotinus horsfieldi leos [sic] (Druce); Fruhstorfer, 1916: 812. Allotinus horsfieldi reverdini Fruhstorfer, 1916: 812. LECTOTYPE cf , PHILIPPINES (SM), here designated
[examined]. Syn. n.
melos (H. H. Druce); Eliot, 1967: 68. re verdini Fruhstorfer; Eliot, 1967: 68. talu Eliot, 1967: 68. Holotype cf , BORNEO: Pulo Laut (BMNH) [examined].
Allotinus macassariensis [sic Allotinus macassariensis [sic Allotinus macassariensis [sic Syn. n.
This species replaces A. horsfieldi, with which it has been confused in the past, in the southern Philippines. Both species are found in Borneo, but A. melos seems to be the commoner throughout most of the island. Sympatric examples of the two species are very difficult to separate by external characters, but males may be separated by the genitalia without difficulty; in melos the terminal process of the valva is short and broad and the tip of the uncus is not produced. Females may be impossible to separate with certainty, but on the underside of the hindwing of melos the postdiscal spot in space 6 is usually placed inside the spot in space 7, sometimes almost half-way to the end-cell bar, whereas in horsfieldi these spots usually overlap. In addition the hindwing termen of melos is very slightly less crenulate.
Eliot (1967) treated melos as conspecific with the allopatric A. macassarensis , but in view of constant small differences in the male genitalia and in the arrangement of the postdiscal spots in spaces 4, 5 and 6 on the underside of the forewing (see p. 39), I no longer maintain this combination.
THE MILETINI
37
Fig. 22 Allotinus (Paragerydus) melos (H. H. Druce); Mindanao. Male genitalia.
Males have vein M3 swollen for three-fifths of its length and the visual brand is clearly defined and about 1-75 mm wide. The underside shows very great individual variation in the intensity of the markings and in the placing of the postdiscal series. The postdiscal series is quite well separated from the wing margins in typical examples but, especially in north-east Borneo, examples occur in which this series is placed close to the termen, as in A. macassarensis . On the hindwing the postdiscal spot in space 6 is usually placed inside that in space 7, sometimes as much as mid-way to the end-cell bar, and this character may be of assistance in separating Bornean females from sympatric A. horsfieldi permagnus .
H. H. Druce stated that the types of melos were in Mus. Cator and Druce. There is in coll. Cator (now in BMNH) a long series of both sexes, none of which has been labelled type, and in the main BMNH coll. a pair ex H. H. Druce coll. labelled as paratypes. Of the latter I designate the male as lectotype; it is labelled /Paratype [yellow]/Cagayan 3. 6. 94/Paragerydus melos d" co-type H. H. Druce/ex coll. Hamilton Druce 1919/Joicey Bequest Brit. Mus. 1934-120/. The female, labelled /Paratype [yellow]/Cagayan 2. 6. 94/P. melos $ co-type H. H. Druce/ex Coll. Hamilton Druce 1919/Joicey Bequest Brit. Mus. 1934-120/, as well as the series in coll. Cator, are paralectotypes.
When naming reverdini, Fruhstorfer referred to large males in coll. Semper. I designate as lectotype a male in SM labelled /19/Bohol/298b/213/Coll. C. Semper/Original of Semper PI. 31, fig. 19 from Mindanao. ? loc. label changed. Det. as A. melos reverdini Fruh. C? J. N. Eliot, ix. 1982/. The male figured by Semper on pi. 31, fig. 19 was said by him to have come from Mindanao. I think it more likely that the label 'Bohol' was inadvertently transferred from the male shown in fig. 18 (which is the holotype of A. posidion georgius and which is without a locality label - see p. 49) than that Semper made a mistake in the legend to his plate 31.
DISTRIBUTION. Cagayan Sulu; Mindanao; Palawan; Balabac I.; Borneo (many localities, including Pulo Laut). There is a male in BMNH labelled /Sula Mangoli, Oct. '97, W. Doherty/, which I suspect may be wrongly labelled although Doherty certainly collected there at that date. Examples from Mindanao tend to be rather heavily marked on the underside. Males from Palawan and Balabac tend to have the visual brand slightly longer and narrower. Examples from Pulo Laut are slightly richer buff on the underside than typical examples from Cagayan Sulu, but the difference is hardly sufficient to justify maintaining talu as a distinct subspecies. There is in coll. Cator a series of seven males and one female from Melikop and Sapagaya (small islands off the north coast of Sabah) which Cator had placed separately from his series of melos as an unrecognised species; they are small, darker brown above and with a greyish white underside bearing darker markings. Superficially they resemble A. leogoron normani, but the male genitalia prove them to be A. melos, of which they could well be regarded as a microsubspecies.
Allotinus (Paragerydus) samarensis sp. n.
(Figs 23, cf genitalia; 65, 66 $, 105 cf )
Upperside of both sexes blackish brown, almost as dark as the shade of A. macassarensis. Male, only known from the nominate subspecies, distinctive in that visual brand extends obscurely as a paler streak
38
J. N. ELIOT
Fig. 23 Allotinus (Paragerydus) samarensis samarensis sp. n. ; Samar. Male genitalia. Phallus with juxta attached.
into fore wing cell basad of origin of vein Cu2. Female distinguished by the possession of a white area on forewing. Underside marked like A melos.
Male genitalia hardly differ from those of A. leogoron, of which it may conceivably be a subspecies; because of the unusual visual brand and shorter swelling of vein M3 in the male, and the distinctive white-marked female, I think it is best regarded as a distinct species.
The species is only known from a very few examples from the southern Philippines and Sulawesi in slightly different subspecies.
Key to the subspecies of A. (P.) samarensis
1 Underside pale greyish white. $ upperside of forewing with white area extending into cell well
basad of origin of vein Cu2 samarensis samarensis (p. 38)
Underside very pale buff. $ upperside of forewing with white area not, or only just, entering cell
samarensis russelli (p. 38)
Allotinus (Paragerydus) samarensis samarensis subsp. n.
(Figs 23, cf genitalia; 65 $, 105 cT)
Cf upperside blackish brown, with vein M3 swollen for just over half its length, visual brand about 1 -25 mm wide; a pale streak above cubitus extending a little basad of origin of vein Cu2 appears to be an extension of the brand. Underside marked as in sympatric A. melos, but with ground colour more greyish white.
$ upperside blackish brown. Forewing with a creamy white streak in spaces 4 and 3 ending 2-5 mm from termen and extending into cell along cubitus to beyond origin of vein Cu2. Underside similar to male.
MATERIAL EXAMINED
Holotype cf , Philippines: Samar, Bagacay, 2000', 31. v. 1979 (C. G. Treadaway) (coll. Treadaway).
Paratypes. Philippines: 1 $ (allotype), west Samar, Hinabangan, 1000 m, 5.ii.l984 (coll. Treadaway); 2 Cf , data as holotype (coll. Treadaway).
Excluded from type-series. Mindanao: 1 $, Surigao Sur, Tandag, viii. 1981 (ex Takanami coll.) (BMNH), with the white streak wider and extending into space 2.
Allotinus (Paragerydus) samarensis russelli subsp. n.
(Fig. 66 $)
Cf unknown.
$ forewing 19-0-21-0 mm. Upperside dark brown with a white discal patch divided by dark veins on forewing, measuring 4-0 mm in space 2 and 8-0 mm in space 3, entering base of space 4 and lower angle of cell where it is sullied. In a second specimen the white patch is smaller, not extending above vein M3 nor entering cell. Underside very pale buff with brown markings arranged as in nominate subspecies.
THE MILETINI
39
MATERIAL EXAMINED Holotype $?> Sulawesi: east-central, north-west of Morowali, Kabalo, 450 m, 10.iii.1980 (A. Bedford
Paratype. 1 $ , data as holotype (coll. Bedford Russell).
Allotinus (Paragerydus) macassarensis (Holland)
(Figs 24 cf , genitalia; 67 cf , 68 £) Paragerydus macassarensis Holland, 1891: 70.
The species name has been consistently misspelled macassariensis in subsequent literature.
Previously (Eliot, 1967) I united this species and A. melos, but in view of small but apparently constant differences in facies and male genitalia it seems best to treat them as separate allopatric species.
The upperside in both sexes is slightly blacker brown than in the other species of the subgenus. On the underside the postdiscal markings, which are usually heavy and dark chocolate brown on an off-white ground, are placed closer to the termen than in the preceding species. A distinguishing character is that on the forewing the postdiscal spots in spaces 4, 5 and 6 are on an irregular curve, with the spot in space 5 overlapping that in space 4 and well separated from that in space 6. In the male genitalia the valvae differ from those of A. melos in two respects; the terminal process is narrower and slightly longer from whichever angle it is viewed and, in lateral view, there is a distinct concavity (indicated in Fig. 24 by an arrow) about two-thirds from the base.
The species is confined to Sulawesi and its satellite islands and is represented by two subspecies.
Key to the subspecies of A. (P.) macassarensis
1 cf vein M3 of forewing swollen for over two-thirds of its length, visual brand usually less than 1
mm wide .................................................................. macassarensis macassarensis (p. 39)
- Cf vein M3 of forewing swollen for only half its length, visual brand more than 1 mm wide
macassarensis menadensis (p. 40)
Allotinus (Paragerydus) macassarensis macassarensis (Holland)
(Figs 24, cf genitalia; 67 cf , 68 £)
Paragerydus macassarensis Holland, 1891: 70, pi. 4, fig. 5 $ . Holotype $ , SULAWESI (not located, probably
in CM). Allotinus horsfieldi macassariensis [sic] (Holland) Fruhstorfer, 1913: 368; 1916: 812, pi. 141h.
Fig. 24 Allotinus (Paragerydus) macassarensis macassarensis (Holland); Sulawesi. Male genitalia. Lower left, right valva enlarged.
40 J. N. ELIOT
Allotinus unicolor damodar Fruhstorfer, 1913: 369; 1916: 811. LECTOTYPE cf , SULAWESI (BMNH),
here designated [examined]. [Synonymised by Eliot. 1967: 68.] Allotinus macassariensis [sic] (Holland); Corbet, 19396: 72, fig. 14 cf genitalia. Allotinus macassariensis macassariensis [sic] (Holland); Eliot, 1967: 68.
In the male the swelling of vein A/3, extending just over two-thirds of its length, is the longest in the subgenus and the visual brand is typically very narrow. But in east-central Sulawesi there is a tendency for the brand to become wider and less well-defined; in one extreme example in coll. Bedford Russell (Fig. 64) there is a diffuse lighter patch which reaches across vein Cu\ more than half-way to vein Cu2. The female has the forewing disc at most only slightly paler.
I designate as lectotype of damodar a small but otherwise normal male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/S. Celebes Tonus 27.xi.06/Fruhstorfer Coll. B.M. 1933-131/unicolor damodar Frhst. [in Fruhstorfer's hand]/. A female labelled /Type [red]/S. Celebes Samangi 17.xi.06/ Fruhstorfer Coll. B.M. 1933-13 1/damodar Frhst. [in Fruhstorfer's hand]/ is a paralectotype.
DISTRIBUTION. South to central Sulawesi; Banggai I.
Allotinus (Paragerydus) macassarensis menadensis Eliot
Allotinus macassariensis [sic] menadensis Eliot, 1967: 68. Holotype cf, SULAWESI: north (BMNH), [examined].
Differs from the nominate subspecies in that the swollen portion of vein M3 in the male is only half its length and the visual brand is quite well-defined and from 1-25 to 2-0 mm wide. In both sexes the forewing disc may be lightened by a scattering of white scales, extreme examples showing an approach to A. samarensis russelli.
DISTRIBUTION. North Sulawesi as far south as Paloe Bay (Lat. 0° 35' S). A single male from Bangka I. (off the tip of the Minahassa Peninsula) is more lightly marked beneath and the ground colour has a buff tint.
Allotinus (Paragerydus) luzonensis Eliot stat. n.
(Figs 25, cf genitalia; 69 cf , 70 9)
Allotinus macassariensis [sic] luzonensis Eliot, 1967: 68, fig. 3 cf genitalia. Holotype cf, PHILIPPINES: Luzon (BMNH) [examined].
The male is greyish brown above; on the forewing vein M3 is swollen for half its length and the visual brand is rather diffuse. The underside is buff, closely striated, and the postdiscal markings in spaces 2 and 3 on the
Fig. 25 Allotinus (Paragerydus) luzonensis Eliot; Luzon. Male genitalia.
THE MILETINI 41
forewing and 6 and 7 on the hindwing are broad and rather blotchy. The female has a small white patch on the forewing disc and an obscure whitish streak in space 5 of the hindwing. In these white patches it shows an approach to the female of A. albatus mendax, but they are less developed than in that species, and the discocellular veins on the hindwing are not blackened.
The species must be closely related to A . albatus because of the same arrangement of the markings on the underside. The male genitalia of the only example dissected are intermediate between those of A. albatus and A. melos.
DISTRIBUTION. Luzon.
Allotinus (Paragerydus) albatus C. & R. Felder
(Figs 26, cT genitalia; 71 £, 72 c?, 73 £) Allotinus albatus C. & R. Felder, [1865]: 287.
The species is readily recognised by the presence in both sexes of white areas on both wings and by the heavily blackened discocellular veins on the hindwing. The underside is white marked more or less as in A. luzonensis, and like this species veins MI and R5 of the forewing usually have a very short common stalk. The male genitalia most nearly resemble those of A leogoron, with the terminal process of the valva rather longhand slender. The species appears to be very rare and to be confined to Wallacea.
Key to the subspecies of A. (P.) albatus
1 $ upperside with larger white areas, on forewing extending from space Ib to base of space 5, on
hindwing from outer part of space 6 to dorsum albatus albatus (p. 41)
- C? $ upperside with smaller white areas, on forewing not below mid-space Ib and not entering
spaces 4 and 5, on hindwing usually not below vein 4 albatus mendax (p. 41)
Allotinus (Paragerydus) albatus albatus C. & R. Felder
(Fig. 71$)
Allotinus albatus C. & R. Felder, [1865]: 267. Holotype $, SULAWESI (BMNH) [examined]. [Allotinus fallax major Felder f. albatus Felder (= maximus Staudinger) sensu Fruhstorfer, 1913: 343, partim; 1916: 809, partim, asalbadus [sic].]
The subspecies is known only from the female holotype, which the authors stated was from 'Celebes Lorquin'. It was therefore surprising to find the holotype bearing labels reading /Halmaheira Lorquin [round blue]/Allotinus albatus Feld./Type [red]/FELDER COLLn. /albatus n./Rothschild Bequest B.M. 1939-1/. As no Allotinus species is known from east of Weber's Line I feel confident that the locality 'Halmaheira' is the result of a wrongly acquired label and that the specimen came from north Sulawesi, where Lorquin is known to have collected.
The forewing is white dusted with brown basally and along the dorsum below vein A\, and with an irregular dark brown border narrowest at vein Cu2, where it measures 2-5 mm; above vein Cu-i it curves to the wing base through the upper third of the cell. The hindwing is white, becoming sullied towards the dorsum, except for most of the cell and spaces 8, 7 and the basal half of space 6 which are brown; the discocellular veins are heavily blackened. The underside is white with heavy brown markings.
Fruhstorfer confused A. albatus with large examples of A. major with extensive white areas on the forewing which were named f . maximus by Staudinger.
DISTRIBUTION. Sulawesi.
Allotinus (Paragerydus) albatus mendax subsp. n.
(Figs 26, c? genitalia; 72 a", 73 $)
[Allotinus fallax C. & R. Felder, 1865: 285, partim, pi. 35, figs 25, 26 $; Semper, 1889: 163, partim. Misidentifi cations . ]
Cf forewing length 20-0 mm. Upperside blackish brown. Forewing with a discal white patch 7-0 mm wide at the bases of spaces 3, 2 and upper part of Ib and entering the lower angle of the cell; the inner and lower part of the cell and lower part of space Ib paler brown than the marginal and costal border. Vein M3 swollen
42
J. N. ELIOT
: .OP
Fig. 26 Allotinus (Paragerydus) albatus mendax subsp. n.; Luzon. Male genitalia. Lower left, internal view of left valva enlarged; lower centre, lateral view of phallus.
for just under three-fifths of its length. Hindwing with a white streak filling most of space 5, the upper basal part of space 4 and just entering the cell; discocellular veins heavily blackened. Underside white, with reddish brown markings arranged as in sympatric A luzonensis.
9 similar to the male, except that the white streak on the hindwing is wider and enters space 6.
MATERIAL EXAMINED
Holotype cf, Philippines: Luzon, bearing labels /Allotinus mendax Bd. Manille/Ex Musaeo Dris BOISDUVAL/ex Oberthur Coll. Brit. Mus. 1927-3/ Allotinus albatus Feld. [in Corbet's hand]/.
Paratypes. Philippines: 1 $ (allotype) Luzon, labelled /N. Luzon, Whitehead. 94/Rothschild Bequest B.M. 1939-1/ (BMNH); 1 cf , Luzon, Bicol Nat. Park, 29.viii.1980 (Y. Takanami) (BMNH); 1 cf , Luzon, Quezon Nat. Park, 27.iv.1983 (Y. Takanami) (BMNH); 1 cf, Luzon, Banahao Ridge, v.1982 (A. Ballantine) (coll. Ballantine); 1 <j>, Luzon, Quezon Nat. Park, 1000', Altimonan Rd, 20.vi.1954 (coll. Treadaway); 1 $, Quezon Nat. Park, Altimonan area, 28. iv. 1969 (coll. Treadaway).
Excluded from type-series. Marinduque: 1 $ , Nu Boac, xi.1980 (coll. Treadaway) with the white area on the hindwing more extensive and nearly reaching the dorsum and the discocellular veins less heavily blackened. Samar: 1 $, east, Borongan, 100', 10.viii.1979 (coll. Treadaway), resembling the allotype above, but beneath with the postdiscal markings diffuse and obscure.
Allotinus (Paragerydus) apries Fruhstorfer
(Figs 27 Cf genitalia; 74, 106 cf ) Allotinus horsfieldi apries Fruhstorfer, 1913: 344, partim.
Fruhstorfer 's type-series probably comprised more than one species; in addition he confused apries with A. strigatus. I use apries here for the species identified as such by Corbet (19396).
The species has several characters which enable it to be recognised with comparative ease. On the forewing veins Ma and R5 share a long common stalk averaging 1-5 mm. On the underside of the forewing the postdiscal series is not, or only very slightly, dislocated at vein M3 and the spots in spaces 4,3,2 and Ib (if present) are small, rounded and well separated from, and parallel to, the termen. In the female the hindwing termen is almost evenly rounded and the cilia are a little longer, but not tufted, at the vein endings. In the male genitalia the comparatively slender phallus is distinctive.
The species occurs throughout Sundaland.
Key to the subspecies of A. (P.) apries
1 Underside pale buff, tending to become darker towards forewing apex in cf, with darker
buff-brown markings, cf visual brand 2-0 mm wide, with half its lower edge touching vein Cui 2
Underside greyish white, with darker greyish brown markings, cf visual brand 1-5 mm wide, with only basal fifth of its lower edge touching vein Cui apries ristus (p. 44)
2 Upperside reddish brown apries apries (p. 43)
- Upperside brown without a reddish tint apries dositheus (p. 44)
THE MILETINI
43
Allotinus (Paragerydus) apries apries Fruhstorfer
(Figs 27, Cf genitalia; 74 cf )
[Allotinus horsfieldi (Moore) sensu Swinhoe, 1910: 198, partim, pi. 617, fig. 1 cf . Misidentification.] Allotinus horsfieldi apries Fruhstorfer, 1913: 344, partim; 1916: 812, partim, pi. 141g cf nee $. LECTO-
TYPE cf , BORNEO (BMNH), here designated [examined]. Allotinus strigatus eupalion Fruhstorfer, 1914: 22; 1916: 813. LECTOTYPE cf , SUMATRA (BMNH), here
designated [examined]. Syn. n. Allotinus apries apries Fruhstorfer; Corbet, 1939ft: 70. Allotinus apries eupalion Fruhstorfer; Corbet, 1939ft: 70; Fleming, 1975: 21, pi. 58, fig. L42 cf ; Eliot, 1978:
240.
In both sexes the underside is nearly always pale buff, with darker buff-brown markings, but occasionally it is more greyish white with only a slight buff tint. In the female the postdiscal series is lightly marked and may be obsolete on the forewing. The male has vein A/3 swollen for a little over three-fifths its length, and the visual brand is clearly defined and 2-0 mm wide. The female is more reddish brown above, with the forewing disc a little paler.
Fruhstorfer described A. horsfieldi apries from several males from Sintang (south-west Borneo) and 10 females from north Borneo. He wrote that the underside was bluish white speckled with pale brown in the male and thicker grey-brown in the female. This description does not accord well with the taxon here treated as apries. However, there is a male from Sintang in BMNH labelled apries by Fruhstorfer and with the underside paler and greyer than usual, which presumably formed part of the type-series. In addition there are seven males of A. horsfieldi permagnus from Sintang ex Fruhstorfer coll. , and it seems likely that these also formed part of Fruhstorfer's type-series even though none had been labelled by him as apries. The only other males of apries ex coll. Fruhstorfer in BMNH are one labelled by Fruhstorfer as A. strigatus Moulton and one which was apparently regarded by Corbet as the holotype of apries and is labelled /Type [red]/Type [Fruhstorfer orange]/Kina Balu ex coll. H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/ apries/. As it did not come from Sintang it cannot have formed part of the type-series, and I reject it as a type. In its place I designate as lectotype of apries the male from Sintang, which is labelled /Sintang Dr Martin H. Fruhstorfer/21.IV.10/Fruhstorfer Coll. B.M. 1933-131/apries Fr. [in Fruhstorfer's hand]/A. apries apries Det. by Dr. A. S. Corbet [in Corbet's hand]/.
There are also two females of apries in BMNH which have been treated as syntypes. One is labelled /Type [red]/Type [Fruhstorfer orange]/Kina Balu Borneo/Fruhstorfer Coll. B.M. 1933-131/horsfieldi apries Frhst. [in Fruhstorfer's hand]/ and is a paralectotype. The other, labelled /Type [red]/Type [Fruhstorfer orange]/Sintang 10.IV.10/Fruhstorfer Coll. B.M. 1933-131/ is rejected as a paralectotype as it did not come from north Borneo and therefore cannot have formed part of the type-series.
Fig. 27 Allotinus (Paragerydus) apries apries Fruhstorfer; Malay Peninsula. Male genitalia. Lower left, lateral view of phallus.
44
J. N. ELIOT
I designate as lectotype of eupalion a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ CMB II.94/Fruhstorfer Coll. B.M. 1933-131/strigatus eupalion Frhst. [in Fruhstorfer's hand]/GENITA- LIA Slide No. ASC 9 Allotinus/.
Here it may be mentioned that the female figured by Fruhstorfer (1916: pi. 141g) as apries is a different species, either A. horsfieldi or A. melos, and also that the male and female figured by Swinhoe (1910: pi. 617, figs 1, la) as A. horsfieldi are in BMNH and are in fact Bornean specimens of A. apries and A. melos respectively.
DISTRIBUTION. Borneo, including Pulo Laut; Malay Peninsula; Sumatra.
Allotinus (Paragerydus) apries dositheus Fruhstorfer
Allotinus strigatus dositheus Fruhstorfer, 1914: 22; 1916: 813. LECTOTYPE cT, JAVA (BMNH), here
designated [examined].
[Allotinus strigatus Moulton sensu Piepers & Snellen, 1918: 22, pi. 20, fig. 17 cf • Misidentification.] Allotinus apries dositheus Fruhstorfer; Corbet, 19396: 72.
The male does not differ from that sex of the nominate subspecies. The female differs in being brown without a reddish tint, in this respect showing parallel variation with the female of A. horsfieldi.
I designate as lectotype of dositheus a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Java Occident. Sukabumi 2000' ex coll. H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/strigatus dositheus Frhst. [in Fruhstorfer's hand]/GENITALIA Slide No. ASC 8 Allotinus/. A female, similarly labelled, which has been treated as a syntype, is a misidentified example of A. portunusportunus.
DISTRIBUTION. Java.
Allotinus (Paragerydus) apries ristus subsp. n.
(Fig. 106 cf )
Cf on the upperside differs from subsp. apries in having a narrower forewing visual brand 1-5 mm wide. The underside differs by its greyish white ground colour and darker greyish brown markings. The striations are fine and the postdiscal spots small.
MATERIAL EXAMINED Holotype cf , Philippines: central Palawan, Languan, i.1981 (Treadaway Coll.).
Allotinus (Paragerydus) corbeti Eliot
(Fig. 28, cf genitalia)
[Allotinus dilutus Corbet sensu Corbet, 19406: pi. 1, figs 13 cf , 14 $. Misidentification.] Allotinus corbeti Eliot, 1956: 34; 1984: 100. Holotype cf , WEST MALAYSIA (BMNH) [examined].
Fig. 28 Allotinus (Paragerydus) corbeti Eliot; Borneo: Pulo Laut. Male genitalia.
THE MILETINI 45
[Allotinus felderi felderi Semper sensu Eliot, 1967: 70, partim. Misidentification.]
Allotinus felderi corbeti Eliot; Eliot, 1967: 70, fig. 2 cf genitalia; 1978: 240; Fleming, 1975: 21, pi. 21, fig. L41cf.
This very small species has an uniform reddish brown upperside in both sexes, and the underside marked as in A. leogoron leogoron. In the male the forewing apex is rounded, and the swelling of vein M3 is confined to the basal quarter of the vein; there is no visual brand. The female has the hindwing termen more strongly crenulate than A leogoron.
Although smaller, it is possible that A corbeti might be confused with A nicholsi, since the males of both species are similar on the upperside. Females are also a similar shade of uniform reddish brown, but those of corbeti are readily separated by their crenulate hindwing with the cilia elongated into tufts at the vein endings. In addition, corbeti has no humeral vein on the hindwing, this vein being present in all six examples of nicholsi which I have examined.
DISTRIBUTION. West Malaysia; Singapore; Sumatra; Pulo Laut; Mindanao; Peninsular Thailand (Pinrata- na). The species appears to be extremely rare, but is perhaps often overlooked.
Allotinus (Paragerydus) unicolorC. & R. Felder
(Figs 29, 30, O" genitalia) Allotinus unicolor C. & R. Felder, [1865]: 286.
This, the most wide-ranging species of the genus, shows great individual variation in size, wing shape and underside pattern. In general the smaller males, with forewing length as little as 11-0 mm, have a more rounded forewing apex and termen and a relatively shorter brand, whilst the bigger males have a more acute apex and straighter termen and a brand which is slightly longer and narrower in relation to wing span. Examples, such as the holotype of unicolor, in which the usual lycaenid markings on the underside are heavy, are comparatively sparsely striated. In examples in which the underside is more densely striated there is usually a reduction in the size and darkness of these markings. On the forewing the postdiscal series is typically on an almost even curve, but examples in which it is dislocated at vein M3 to a greater or less extent are equally frequent.
The best characters for identifying the species are the male genitalia (Figs 29, 30) with a broad valva with the terminal process almost in the centre line, the short stalk of veins MI and R5 of the forewing and the relatively short swelling of vein M3 in the male.
Fruhstorfer grouped the complex into three sympatric 'species': unicolor, aphocha andposidion. Under unicolor he grouped comparatively small specimens with a whitish ground colour on the underside and heavy postdiscal markings, and under aphocha specimens of similar size and wing shape but with less pronounced markings. He reserved posidion for large specimens in which the male forewing was more produced. Corbet (1939ft), though at first inclined to agree with Fruhstorfer's arrangement, finally decided that all three constituted a single, variable species, but at the same time erected a new 'species', A dilutus,
Fig. 29 Allotinus (Paragerydus) unicolor continentalis Fruhstorfer; Burma. Male genitalia.
46
J. N. ELIOT
Fig. 30 Allotinus (Paragerydus) unicolor zitema Fruhstorfer; Sulawesi. Male genitalia.
which he later (1956) relegated to subspecies, for very small examples from the Malay Peninsula. The various sizes, wing shapes and patterns are connected iby intermediates and, like Corbet, I can find no differences in the male genitalia of the largest and smallest specimens. Therefore I also consider that there is only one species, to which I now add the taxon continentalis , which has previously been treated as a subspecies of A. horsfieldi or as a distinct species.
It is extraordinarily difficult to decide what constitutes a valid subspecies because of the great range of phenetic variation. When large series are available for comparison, as in coll. BMNH, it is possible to detect differences between the average phenotype of different geographical areas, but because of the overlap of phenetic characters it may often be impossible to ascribe individual specimens, if deprived of their locality labels, to any particular country of origin. I have opted to retain as subspecies those groups of populations which can be distinguished by a mean of differences and which are contained within generally accepted faunal areas in preference to lumping together into just two or three polytypic subspecies the populations of widely separated geographical areas which in all probability differ genetically to a considerable degree. In using the key below this limitation should be borne in mind.
The species ranges from Assam to the Lesser Sunda Is. , Philippines and Sulawesi (including the Sula Is.) , and is most abundant in low level primary or secondary forest.
Key to the subspecies of A. (P.) unicolor
1 Underside of hindwing with the postdiscal spot in space 6 well inside that in space 7 and often
mid-way to the end-cell spot 2
Underside of hindwing with the postdiscal spot in space 6 much closer to the spot in space 7 than to the end-cell spot 7
2 Underside ground colour greyish white . cf vein M3 of fore wing swollen for not more than half its
length. £ upperside of forewing with disc not conspicuously paler 3
Underside ground colour pale buff, cf vein M3 swollen for a little over half its length. $ forewing disc conspicuously paler and often sullied whitish unicolor continentalis (p. 47)
3 cf forewing brand 1 -0 mm or more wide. Continental Asia and Borneo 4
Cf brand less than 1-0 mm wide. Philippines and Sulawesi 6
4 9 upperside reddish brown 5
9 upperside brown without a reddish tint unicolor rekkia (p. 48)
5 cf forewing brand about 1-0 mm wide unicolor unicolor (p. 47)
Cf brand comparatively short and broad, about 1-75 mm wide in large specimens.
Underside rather lightly marked unicolor moorei (p. 48)
6 Underside with postdiscal markings usually heavy and sharply defined, cf vein A/3 swollen for
just under half its length, brand sharply defined unicolor georgius (p. 49)
Underside with postdiscal markings usually rather light and tending to be blurred, cf vein M3 swollen for half its length and brand usually rather inconspicuous unicolor zitema (p. 50)
THE MILETINI 47
7 Upperside hindwing with postdiscal spot in space 6 often below and conjoined to the spot in
space 7. $ upperside brown without a reddish tint, forewing disc paler . . . unicolor postilion (p. 48) Underside hindwing with the postdiscal spot in space 6 nearly always inside the spot in space 7.
9 upperside with a slight reddish tint, forewing disc only slightly paler... unicolor aphocha (p. 48)
Allotinus (Paragerydus) unicolor unicolor C. & R. Felder
Allotinus unicolor C. & R. Felder, [1865]: 286. Holotype 'cf' recte $, SINGAPORE (BMNH) [examined].
[Allotinus posidion myriandus Fruhstorfer, 1913: 368 (partim); 1916: 811 (partim).]
Allotinus posidion eurytanus Fruhstorfer, 1913: 368; 1916: 811. LECTOTYPE cf , BORNEO (BMNH), here
designated [examined]. Syn. n. Allotinus unicolor unicolor Felder; Fruhstorfer, 1913: 369; 1916: 809, pi. 141i; Corbet, 19396; 68, pi. 1, figs
3 $ holotype, 4 cf; 1956: 269, pi. 44, fig. 153 cf; Eliot, 1978: 240, pi. 20, figs 5 cf, 6 $. [Allotinus aphocha aphocha (Kheil); Fruhstorfer, 1913: 370 (partim); 1916: 810 (partim).] Allotinus unicolor eurytanus Fruhstorfer; Corbet, 19396: 70, pi. 1, figs 9 cf 'holotype of eurytanus' (in
error), 10 cf 'holotype of eurytanus f. rebilus' (in error).
Allotinus dilutus Corbet, 19396: 70. Holotype cf , WEST MALAYSIA (BMNH) [examined]. Syn. n. Allotinus unicolor dilutus Corbet; Corbet, 1956: 269; Cantlie, 1967: 27; Fleming, 1975: 21, pi. 57, fig.
L40cf.
The female holotype, figured by Corbet (19396) and mistaken by the Felders for a male, is an atypical specimen with heavy markings and a whiter than usual ground colour. Such specimens occur most often in Singapore, where they are connected by intermediates to normal phenotypes. The male figured by Corbet (19396), which matches the holotype fairly well but has a slightly greyer ground colour on the underside (as usual in males), would probably have been identified by Fruhstorfer as A. posidion myriandus because of its size and wing shape.
In general Bornean examples have the lycaenid markings smaller than in those from continental Asia, but compensate by being slightly more densely striated. This tendency is most extreme in examples from south-west Borneo, named eurytanus by Fruhstorfer, which are particularly densely striated but can be matched by occasional examples from other areas.
I designate as lectotype of eurytanus a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Sintang 19.IV.10/Fruhstorfer Coll. B.M. 1933-131/posidion eurytanus Fr. [in Fruhstorfer's hand]/. It was figured by Corbet (19396: fig. 10) in error as the holotype of A. unicolor eurytanus f. rebilus. The smaller specimen figured by Corbet at fig. 9 in error as the holotype of eurytanus is labelled /Type [red]/Type [Fruhstorfer orangeJ/Sintang 26.IV.10/aphocha rebilus Frhst. [in Fruhstorfer's hand]/GENITALIA Slide No. ASC 14 Allotinus/. Despite Fruhstorfer's identification label it cannot be accepted as lectotype of rebilus, since this taxon was described from North Borneo; it probably formed part of Fruhstorfer's original type-series of four males and one female of A. posidion eurytanus, and how it obtained its label as rebilus is a mystery.
DISTRIBUTION. This very variable subspecies is found in south Burma, where it has a zone of intergradation with subsp. continentalis between Rangoon and Tavoy, and in Peninsular Thailand, West Malaysia, Singapore, Lingga Is., Natuna Is. and throughout Borneo, except in the Kina Balu area where it merges into subsp. moorei.
Allotinus (Paragerydus) unicolor continentalis Fruhstorfer
(Fig. 29, cf genitalia)
[Paragerydus horsfieldi (Moore) sensu de Niceville, 1890: 26, pi. 36, fig. 156 cf . Misidentification.] [Allotinus horsfieldi (Moore) sensu Bingham, 1907: 287, 299, fig. 73 cf ; sensu Swinhoe, 1910: 198 (partim,
nee pi. 617, figs 1, la, Ib, Ic). Misidentifications.] Allotinus horsfieldi continentalis Fruhstorfer, 1913: 344; 1916: 812; Evans, 1932: 212; Cantlie, 1963: 27.
Holotype cf , BURMA: Bhamo (probably in ZSI). [Allotinus posidion subsp.; Fruhstorfer, 1913: 368.] Allotinus posidion atacinus Fruhstorfer, 1916: 811; Evans, 1932: 212. Holotype $. BURMA (BMNH)
[examined]. [Synonymised by Eliot, 1967: 70.]
Allotinus unicolor atacinus Fruhstorfer; Corbet, 19396: 68, pi. 1, fig. 2 $ holotype; Cantlie, 1963: 27. Allotinus continentalis continentalis Fruhstorfer; Corbet, 19396: 72. Allotinus continentalis Fruhstorfer; Eliot, 1967: 70, fig. 4 cf genitalia.
48 J. N. ELIOT
This is the most distinctive and largest subspecies of A. unicolor, with the forewing length of the male usually 18-19 mm. On the underside both sexes are pale buff. On the upperside the male brand is longer and wider than in the other subspecies, usually 2-0 mm wide and touching vein Cu} at its origin, while the swelling of vein M3 is a little over half its length. The female is distinguished by the prominently paler discal area on the upperside of the forewing, which may be sullied whitish in the dry season.
DISTRIBUTION. Assam; Burma as far south as Tavoy; north-west Thailand. Around the latitude of Rangoon it intergrades with subsp. unicolor.
Allotinus (Paragerydus) unicolor rekkia Riley & Godfrey
Allotinus posidion rekkia Riley & Godfrey, 1921: 180, pi. 6, figs 1 d", 2 $. Holotype cf , THAILAND: east
(BMNH) [examined]. Alliotinus unicolor rekkia Riley & Godfrey; Eliot, 1967: 70.
Females are brown without the reddish tint of subsp. unicolor, but otherwise the subspecies does not differ and is of doubtful validity.
DISTRIBUTION. Only known from eastern Thailand, but probably also occurs in Cambodia, Laos and Vietnam.
Allotinus (Paragerydus) unicolor moorei (H. H. Druce)
Paragerydus moorei H. H. Druce, 1895: 562, pi. 31, figs 5 cf , 6 $ . Syntypes, BORNEO: Mt Kina Balu (coll.
Staudinger, probably in MNHU).
Allotinus paetus moorei (Druce) Fruhstorfer, 1913: 369: 1916; 811. Allotinus aphocha rebilus Fruhstorfer, 1913: 370; 1916: 810. LECTOTYPE $, BORNEO (BMNH), here
designated [examined]. Syn. n. Allotinus moorei (H. H. Druce); Corbet, 19396: 68.
Druce's figure of the male shows a comparatively large specimen weakly marked beneath and with the brand on the upperside of the forewing rather short, broad and diffuse. Judging by material in BMNH such males are rare, and are connected by intermediates to smaller males which differ little, if at all, from nominate unicolor. The subspecies is therefore of doubtful validity.
Fruhstorfer's taxon rebilus is applicable to smaller examples. I designate as lectotype a female in BMNH labelled /Type [red]/Kina Balu/ex coll. H. Fruhstorfer/ Allotinus aphocha rebilus Fr. [in Corbet's hand]/.
DISTRIBUTION. The subspecies flies on Mt Kina Balu, and might conceivably be a local modification occurring at higher elevations than normal examples referable to subsp. unicolor.
Allotinus (Paragerydus) unicolor aphocha Kheil
Allotinus aphocha Kheil, 1884: 28, pi. 5, fig. 30 $. Holotype $, NIAS (probably in MNHU).
Allotinus posidion myriandus Fruhstorfer, 1913: 368; 1916: 811. Lectotype cf, SUMATRA (BMNH), by
designation (as holotype) of Corbet, 19396: 68 [examined]. Syn. n.
Allotinus aphocha aphocha Kheil; Fruhstorfer, 1913: 370 (partim); 1916: 810 (partim), pi. 141g cf 9- Allotinus unicolor myriandus Fruhstorfer; Corbet, 19396: 68, pi. 1, figs 5 cf holotype, 6 9 allotype. Allotinus unicolor aphocha Kheil; Corbet, 19396: 70, pi. 1, fig. 11 $.
Differs from subsp. unicolor only in that on the underside of the hindwing the postdiscal spot in space 6 is usually placed closer to the spot in space 7. Many specimens are inseparable from unicolor, so the subspecies is of doubtful validity.
Fruhstorfer did not designate a type of myriandus, but Corbet (19396) figured its 'holotype' male and 'allotype' female, and this action constitutes a valid lectotype selection.
DISTRIBUTION. Sumatra; Bangka I.; Batu Is; Mentawai Is; Nias I.
Allotinus (Paragerydus) unicolor posidion Fruhstorfer
Allotinus posidion posidion Fruhstorfer, 1913: 368; 1916: 811. Lectotype cf, JAVA: west (BMNH), by
designation (as holotype) of Corbet, 19396: 68 [examined]. Allotinus posidion molionides Fruhstorfer, 1913: 368; 1916: 811. Lectotype cf, BALI (BMNH), by
designation (as holotype) of Corbet, 19396: 70 [examined]. Syn. n.
THE MILETINI 49
Allotinus posidion niceratus Fruhstorfer, 1913: 368; 1916: 812. LECTOTYPE cf, SUMBAWA (BMNH),
here designated [examined]. Syn. n. Allotinus unicolor enganicus Fruhstorfer, 1913: 369; 1916: 811. Lectotype d", ENGANO I. (BMNH), by
designation (as holotype) of Corbet, 19396: 70, pi. 1, fig. 12 [examined]. Syn. n. Allotinus unicolor bajanus Fruhstorfer, 1913: 369; 1916: 811; Corbet, 19396: 70. LECTOTYPE cf,
LOMBOK (BMNH), here designated [examined]. Syn. n. Allotinus aphocha enatheus Fruhstorfer, 1913: 370; 1916: 810. LECTOTYPE $, JAVA (BMNH), here
designated [examined]. [Synonymised by Corbet, 19396: 70.]
Allotinus horsfieldii [sic] f. posidion Fruhstorfer; Piepers & Snellen, 1918: 12, pi. 19, figs 14a cf , 14b $. Allotinus suka Piepers & Snellen, 1918: 13 (partim), pi. 27, fig. 181 cf . LECTOTYPE cf , JAVA (probably in
RNH), here designated. [Aberration reduced to infrasubspecific status by Corbet, 19396: 70.] Allotinus unicolor Felder; Piepers & Snellen, 1918: 14, pi. 19, figs 13a cf , 13b ?. [Allotinus aphocha Kheil; Piepers & Snellen, 1918: 15, pi. 19, figs 15a cf , 15b $.] Allotinus unicolor posidion Fruhstorfer; Corbet, 19396: 68, pi. 1, figs 7 cf holotype, 8 $ 'allotype' of
enatheus.
Allotinus unicolor molionides Fruhstorfer; Corbet, 19396: 70, pi. 1, fig. 13 cf holotype. Allotinus unicolor bajanus Fruhstorfer; Corbet, 19396: 70.
The female lacks the reddish tint of subsp. myriandus and usually the forewing disc is paler. On the underside of the hindwing the postdiscal spot in space 6 is more often directly below, and conjoined to, the spot in space 7; this character occurs most often in examples from the Lesser Sunda Is. , which also have the postdiscal spots on average heavier than in Javanese examples.
Fruhstorfer did not designate types of his taxa, but Corbet (19396) figured the 'holotype' males of posidion, molionides and enganicus, and this action constitutes valid lectotype selections. He also figured a female which he described as A. unicolor posidion f. enatheus Fruh. $ allotype. I now designate this female as lectotype of enatheus; it is labelled: /Type [red]/Type [Fruhstorfer orange]/Java Occident. Sukabumi 2000' ex coll. H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/aphocha enatheus Fr. [in Fruhstorfer's hand]/.
I designate as lectotype of niceratus a male in BMNH labelled /Type [red]/Sumbawa/Allotinus posidion niceratus Fruh. Type [in Corbet's hand]/Adams Bequest B.M. 1912-399/. It is to be presumed that Fruhstorfer saw this specimen during his visit to BMNH prior to publication of his 1913 paper, as there are no specimens from Sumbawa from his collection in BMNH.
I designate as lectotype of bajanus a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Pringabaja April 1896 H. Fruhstorfer/unicolor bajanus Frhst. [in Fruhstorfer's hand]/.
Piepers & Snellen (1918) figured two different species as Allotinus suka: a female on pi. 19, fig. 12b to which they had previously referred as Allotinus horsfieldii and a male on pi. 27, fig. 181. When relegating suka to the status of a form of A. unicolor posidion, in which the usual striations of the underside are absent, Corbet (19396: 70) referred only to the latter figure which he said, in error, represented a female. Corbet's action could, perhaps, be taken to restrict the name suka to the species represented in fig. 181 , but to make absolutely certain that there should be no confusion in the application of the name I now designate this male as lectotype. It should be in RNH.
DISTRIBUTION. Java; Bali; Lombok; Sumbawa; Engano I.
Allotinus (Paragerydus) unicolor georgius Fruhstorfer
[Paragerydus horsfieldi (Moore) sensu Semper, 1889: 164, partim. pi. 31, fig. 18 cf •]
Allotinus posidion georgius Fruhstorfer, 1913: 368, partim; 1916: 812. Holotype cf, PHILIPPINES: Bohol
(SM) [examined]. Allotinus unicolor leitus Fruhstorfer, 1916: 811. Holotype $, PHILIPPINES: Mindoro (coll. Staudinger,
probably in MNHU).
Fruhstorfer named georgius from Semper's records and figures of 'horsfieldi' from Bohol and Mindanao, which he subsequently (1916) realised represented two species. Thereupon he restricted georgius to Bohol, so that this name applies to the male figured by Semper at fig. 18, which automatically becomes the holotype; it is labelled /18/213/Coll. C. Semper/reverdini/original of Semper PI. 31, fig. 18 cf from Bohol. Holotype of Allotinus posidion georgius Fruh. det. J. N. Eliot ix.1982/. It is heavily marked below much as in the holotype of unicolor, but on the upperside the brand is a little shorter and narrower than in that subspecies. Fruhstorfer named leitus from a single female with a yellowish discal area on the forewing and heavy
50
J. N. ELIOT
markings on the underside. The latter character suggests that it pertains to the same subspecies as georgius, but as I have seen no examples from Mindoro it is provisionally placed in synonymy.
DISTRIBUTION. A unicolor must be very rare in the Philippines, as it is not represented in coll. Treadaway, and in BMNH there are only two females, one of which is labelled 'Philippines Pryer'. This appears to be an aberration; it is lightly striated on the underside, as in examples from north Borneo, and the postdiscal markings are elongated into longitudinal streaks. It may have come from the Sulu Is. , which were visited by Pryer, or may be a mislabelled example from Borneo, where Pryer did nearly all his collecting.
Allotinus (Paragerydus) unicolor zitema Fruhstorfer (Fig. 30, cf genitalia)
Allotinus aphocha zitema Fruhstorfer, 1916: 810. LECTOTYPE $, SULAWESI (BMNH), here designated [examined].
The subspecies is distinguished by the male brand, which is a little narrower and more obscure than in other subspecies.
I designate as lectotype a female in BMNH labelled: Type [red]/Type [Fruhstorfer orange]/Nord- Celebes Toli Toli Nov. - Dez. 1895 H. Fruhstorfer/aphocha zitema Fr. [in Fruhstorfer's hand]/Fruhstorfer Coll. B.M. 1933-131/. I have seen no males ex Fruhstorfer coll.
DISTRIBUTION. Sulawesi and the Sula Is.
Allotinus (Paragerydus) paetus (de Niceville) (Fig. 31 cf genitalia)
Paragerydus paetus de Niceville, 1895: 269, pi. O, fig. 12 cf . Syntypes, SUMATRA: north-east (probably in
ZSI).
Allotinus paetus paetus (de Niceville) Fruhstorfer, 1913: 369; 1916: 811, pi. 141i <j>. Allotinus paetus (de Niceville); Corbet, 19396: 68, pi. 1, fig. 1 cf .
This species bears a fairly close resemblance to examples of A. unicolor which are strongly marked beneath, as in the holotype of unicolor. But it is larger, with forewing length averaging 18-19 mm in males, the forewing brand and swelling of vein A/3 are slightly longer than in all unicolor subspecies apart from subsp. continental, being just over half the length of the vein, and on the underside the ground colour is more chalky whitish. The male genitalia are rather similar to those of A. unicolor, but the phallus is stouter and the distal portion of the valva narrower, with the terminal process not curved up so strongly towards the centre line.
DISTRIBUTION. The species is only known from Sumatra, where it appears to fly in the Barisan Range from the Battak mountains in the north to the extreme south, where it was taken in numbers by Doherty.
Fig. 31 Allotinus (Paragerydus) paetus (de Niceville); Sumatra. Male genitalia.
THE MILETINI
51
Fig. 32 Allotinus (Paragerydus) parapus Fruhstorfer; Borneo. Male genitalia. Lower right, ventral view of valvae and phallus.
Allotinus (Paragerydus) parapus Fruhstorfer
(Fig. 32, cf genitalia)
Allotinus parapus Fruhstorfer, 1913: 343; 1916: 809, pi. 141h cf; Corbet, 19396: 66, fig. 8 cf genitalia. LECTOTYPE cf , BORNEO (BMNH), here designated [examined].
The sexes are alike in wing shape and in having a rounded hindwing termen with the cilia inconspicuously elongated at the vein endings. In the male vein M3 is swollen for just under half its length and clothed with the usual specialised scales. The white forewing b< >rdered with dark brown and the paler brown hindwing render the species unmistakable.
Both Fruhstorfer and Corbet, whose figure of the male valva is completely misleading, stated that vein M3 was not swollen in the male, and on this account placed the species in the /a//ajt-group (Artengruppe Allotinus). In fact the swelling is as well developed as in some other species of the subgenus, but because the surrounding area is white no visual brand is apparent.
I designate as lectotype a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/Fruhstorfer Coll. B.M. 1933-131/Nord-Borneo/ Allotinus parapus Frhst. [in Fruhstorfer's hand]/. A female, similarly labelled, is a paralectotype.
DISTRIBUTION. The species is montane and is known only from Mt Kinabalu in Sabah.
Allotinus (Paragerydus) nivalis (H. Druce)
(Figs 33, 34, cf genitalia; 75 cf ) Miletus nivalis H. Druce, 1873: 348.
This and the next two taxa form a natural group of small species distinguished by the possession of a white fleck at the end of vein R5 at the apex of the forewing in addition to the usual white flecks at the ends of veins Sc, RI, R2 and R3. In males the swelling of vein M3 is weak and short, and there is no visual brand.
In A. nivalis the forewing termen is almost regular in both sexes, and on the underside of the hindwing the central spot in space 7 is not, or only very little darkened. On average it is larger than the allied A. substrigosus and A. davidis.
The species is confined to Borneo and the Philippines.
Key to the subspecies of A. (P.) nivalis
1 Underside of forewing with a more or less developed submarginal brownish blotch about 1 -0-1 -5 mm wide astride vein M3; postdiscal series strongly dislocated at veins M3 and Cu2
nivalis nivalis (p. 52)
52
J. N. ELIOT
Fig. 33 Allotinus (Paragerydus) nivalis nivalis (H. Druce); Borneo. Male genitalia.
- Underside without a brownish blotch astride vein M3; postdiscal series placed nearer the termen
and only a little dislocated at veins A/3 and Cu2 nivalis felderi (p. 52)
Allotinus (Paragerydus) nivalis nivalis (H. Druce)
(Fig. 33, cf genitalia)
Miletus nivalis H. Druce, 1873: 348, 'cf' recte <j>. Holotype $, BORNEO (BMNH) [examined]. Allotinus nivalis nivalis (Druce) Fruhstorfer, 1913: 370; 1916: 810, pi. 141g. Allotinus nivalis (H. Druce); Eliot, 1967: 71.
The characters of the subspecies are given in the key. The figure by Fruhstorfer (1916: 141g) is very poor and shows an undersized specimen.
DISTRIBUTION. Throughout Borneo, including Pulo Laut.
Allotinus (Paragerydus) nivalis felderi Semper
(Fig. 34 cf genitalia; 75 cf )
Allotinus felderi Semper, 1889: 163, pi. 31, fig. 22 $ ; Fruhstorfer, 1913: 370; 1916: 810. LECTOTYPE cf , PHILIPPINES (SM), here designated [examined].
Fig. 34 Allotinus (Paragerydus) nivalis felderi Semper; Mindanao. Male genitalia. Lower left, ventral view of valvae and phallus of another specimen.
THE MILETINI 53
Allotinus felderi felderi Semper; Eliot, 1967: 70, partim. Allotinus niv alls felderi Semper; Eliot, 1984: 100.
The differences from the nominate subspecies are given in the key.
Semper described the taxon from two males from Mindanao and two females from Luzon. I designate as lectotype one of the males in SM, labelled /Coll. C. Semper/Ost Mind./212/All. felderi typ. Semper/.
Eliot (1967) confused felderi with A. corbeti from a pair oicorbeti and one exceptionally small female of true felderi from Mindanao, all of which had been placed under the latter name in coll. BMNH.
DISTRIBUTION. Throughout the Philippines. In addition to examples from Luzon, Sibuyan and Mindanao in BMNH I have examined examples in coll. Treadaway from Marinduque, Negros and Samar.
Allotinus (Paragerydus) substrigosus (Moore)
(Figs 35, cf genitalia; 76, 77 cT) Logania substrigosa Moore, 1884: 22.
Until separated by Eliot (1967) all authors treated this species as, or as a subspecies of, A. nivalis, from which it differs as follows. On average it is smaller, with forewing length occasionally as little as 9-0 mm. In both sexes the forewing termen is distinctly crenulate, becoming more exaggerated in the dry season form from Burma. On the underside of the hindwing the central spot in space 7 is at least partially blackened and there is often a black subbasal spot in space Ib, the corresponding spots in nivalis not being blackened. The male genitalia are very similar, but in substrigosus the phallus is narrower than in sympatric nivalis.
The species ranges from central Burma and Thailand to Sundaland and Mindanao. In Borneo it appears to be rarer than A. nivalis.
Key to the subspecies of A. (P.) substrigosus
1 Underside of hindwing with the spot mid-space 7 blackened only in upper half of space, and not
more than 1 -0 mm wide 2
- Underside of hindwing with the spot mid-space 7 solidly black, nearly 2-0 mm wide and
extending right across space substrigosus ballantinei (p. 55)
2 Underside greyish- white. Forewing crenulate 3
- Underside pale buff. Forewing only very weakly crenulate substrigosus yusukei (p. 55)
3 On underside of forewing the white fleck at end of vein 7 barely enters space 6 4
- On underside of forewing the white fleck at the end of vein R5 is continued as an oblique white
streak half-way across space 6 substrigosus substrigosus (p. 53)
4 cf with vein M3 swollen for only one-quarter of its length substrigosus lenaia (p . 54)
- cf with vein M3 swollen for one-third of its length, as in subsp. substrigosus
substrigosus sibyllinus (p. 54)
Allotinus (Paragerydus) substrigosus substrigosus (Moore)
(Fig. 35, cf genitalia)
Logania substrigosa Moore, 1884: 22; 1886; 39, pi. 3, fig. 8 $. Holotype $, BURMA: Mergui Archipelago
(probably in ZSI).
[Paragerydus nivalis (H. Druce) sensu Distant, 1884: 207, pi. 22, fig. 11 $. Misidentification.] [Allotinus nivalis (H. Druce) sensu de Niceville, 1890: 30 partim, pi. 36, fig. 159 $ holotype of substrigosa;
sensu Bingham, 1907: 301; sensu Swinhoe, 1910: 197, pi. 616, figs 2, 2b cf, 2a $; sensu Piepers &
Snellen, 1918: 16, pi. 20, fig. 19 cf . Misidentifications.] Allotinus nivalis magaris Fruhstorfer, 1913: 370; 1916: 810. LECTOTYPE cf , SUMATRA (BMNH), here
designated [examined]. [Synonymised by Eliot, 1967: 71.] Allotinus nivalis substrigosa (Moore) Fruhstorfer, 1916: 810; Evans, 1932: 212; Corbet, 19396: 68; Cantlie,
1963: 28.
Allotinus substrigosa substrigosa (Moore); Eliot, 1967: 71; 1978: 240. Allotinus substrigosus substrigosus (Moore); Fleming, 1975: 21, pi. 57, fig. L38 $ •
In the male the swelling of vein M3 extends to one-third of its length. On the underside there is a prominent white streak at the forewing apex, and on the hindwing the central spot in space 7 is strongly blackened. From central Burma as far south as Tavoy a distinct dry season form occurs. The forewing termen is more
54
J. N. ELIOT
Fig. 35 Allotinus (Paragerydus) substrigosus yusukeisubsp. n.; Mindanao. Male genitalia. Lower left, A. substrigosus substrigosus (Moore), Malay Peninsula; right valva and phallus.
strongly crenulate; on the underside the ground colour becomes more greyish; on the forewing the apex and margin to a depth of 2-0-3-0 mm are shaded with brown, so that the oblique apical white streak stands out more conspicuously; and on the hindwing there is a similar brown area in spaces 3, 4 and 5, and the outer postdiscal spot in space 7, and sometimes that in space 6 also, may be blackened on their inner edges - a feature otherwise only found in A. davidis. This dry form, with its mottled appearance and crenulate wings, gives the impression of a Logania species.
I designate as lectotype of magaris a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Selesseh 15.VII.94/Fruhstorfer Coll. B.M. 1933-131/Sumatra Monies Battak ex coll. Fruhstorfer/nivalis magaris Frhst. [in Fruhstorfer's hand]/. The locality Selesseh lies a little above sea level in north-east Sumatra and is more likely to be the correct locality than the Battak Mts.
DISTRIBUTION. Burma, as far north as east Pegu; Thailand; West Malaysia; Sumatra; Borneo; Java (Piepers & Snellen, 1918).
Allotinus (Paragerydus) substrigosus lenaia Fruhstorfer
Allotinus nivalis lenaia Fruhstorfer, 1913: 370; 1916: 810. LECTOTYPE cf, NIAS (BMNH), here
designated [examined]. Allotinus substrigosa lenaia Fruhstorfer; Eliot, 1967: 72.
In the single male which I have seen the swelling of vein M3 is confined to the basal quarter of the vein. On the underside of the forewing the white streak at the end of vein R5 at most barely enters space 6, and in females the markings are on average lighter than in the nominate subspecies.
I designate as lectotype a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/Nias ex coll. Fruhstorfer/nivalis lenaia Frhst. [in Fruhstorfer's hand]/.
DISTRIBUTION. Nias I.
Allotinus (Paragerydus) substrigosus sibyllinus Riley
Allotinus nivalis sibyllina Riley, 1944: 254, pi. 2, fig. 27 cf . Holotype cf , MENTAWAI Is.: Sipora (BMNH) [examined].
The upperside agrees with the nominate subspecies. On the underside the white fleck at the forewing apex is as redu'ced as in subsp. lenaia. On the hindwing the central spot in space 7 may be heavily blackened, as in the nominate subspecies, or only lightly blackened. In his original description Riley wrote that this spot is not blackened, but in seven out of the eight specimens in the type-series it is blackened to a greater or less degree. The eighth specimen, a worn male, is so rubbed that it is possible some black scales were originally present. The subspecies seems to be only doubtfully separable from subsp. lenaia.
DISTRIBUTION. Mentawai Is.
THE MILETINI 55
Allotinus (Paragerydus) substrigosus ballantinei subsp. n.
(Fig. 76 cf )
Cf forewing 13-5 mm. Wing shape, as in subsp. substrigosus, with forewing termen crenulate and apex rather pointed. Swelling of vein M3 inconspicuous, extending a little under one-third of its length. Underside very pale buff, rather densely striated but with postdiscal spots small or evanescent, as in some individuals of subsp. substrigosus. Forewing with white speck at end of vein R5 not extending into space 6. Hindwing with spot in mid-space 7 exceptionally large, rather quadrate, nearly 2-0 mm wide and extending right across space.
MATERIAL EXAMINED Holotype cf , Philippines: Palawan, Port Barton, ix.1983 (A Ballantine) (BMNH).
The subspecies is named after Dr Alistair Ballantine, who caught the unique holotype and presented it to BMNH.
Allotinus (Paragerydus) substrigosus yusukei subsp. n.
(Figs 35 cf genitalia; 77 cf )
Cf forewing 11-5 mm. Termen more rounded and apex less pointed than in foregoing subspecies. Swelling of vein M3 short, extending over one-quarter of vein, overlying specialised scales pale so that it stands out prominently. Underside ground colour pale buff; forewing with white fleck at end of vein R5 extending as an oblique white streak half-way across space 6. Hindwing with central black spot in space 7 narrow but well-defined; a small subbasal black spot in space la.
MATERIAL EXAMINED Holotype cf , Philippines: Mindanao, Tandag, Surigao, xii.1982 (ex Takanami coll.) (BMNH).
The subspecies is named after Mr Yusuke Takanami, who generously presented the holotype and many other rare Philippine lycaenids to BMNH.
Allotinus (Paragerydus) davidis Eliot (Fig. 36, cf genitalia)
Allotinus davidis Eliot, 1959: 377, pi. 10, figs 6 cf , 7 $ , text-fig, cf valva; 1978: 240; Fleming, 1975: 21, pi. 57, fig. L39 $. Holotype cf , SINGAPORE (BMNH) [examined].
The wing shape is that of A. nivalis, with the forewing termen barely perceptibly crenulate. On the upperside both sexes are reddish brown. In the male vein M3 of the forewing is swollen only in its basal quarter. On the underside the ground colour is pale buff; on the forewing the white fleck at the apex does not cross vein R5; on the hindwing the central spot in space 7 is blackened and the postdiscal spot is also narrowly blackened on its inner edge.
In the male genitalia the valva differs from all others in the subgenus by not having the costa truncate before the terminal process, which curves upwards in the manner of subgenus Allotinus. In all other
Fig. 36 Allotinus (Paragerydus) davidis Eliot; Singapore. Male genitalia.
56 J. N. ELIOT
respects the genitalia are typical of Paragerydus, and the absence of a humeral vein on the hindwing helps to confirm its position in this subgenus.
DISTRIBUTION. West Malaysia; Singapore; Sumatra (coll. Diehl); Peninsular Thailand (Pinratana). The species appears to be extremely rare, but is perhaps sometimes overlooked because of its small size and resemblance to A. substrigosus .
Allotinus (Paragerydus) drumila (Moore)
(Fig. 37, cf genitalia) Miletus drumila Moore, [1866]: 777.
This large species stands rather far apart from the remainder of the subgenus in its markings and in its pronounced seasonal variation. On the other hand the male genitalia, presence on the forewing of a swollen vein M3 and absence of a humeral vein on the hindwing, are characteristic of Paragerydus, so that it seems unnecessary to retain for it the separate subgenus Miletographa.
On the underside the appearance is of a Miletus species, with the catenulate markings characteristic of that genus; the difference is that in A. drumila the ground between the usual lycaenid markings is speckled with small, irregular pale-edged spots. The upperside and especially the seasonally dimorphic wing shape recall the smaller, sympatric Miletus chinensis. The wet season form is brown, with a similar curved series of postdiscal spots on the forewing, paler brown in the male, larger and sullied white in the female. In the dry season the forewing acquires a sharp point at the apex and prominent lobe at the tornus, and the mainly white female is broadly similar to the dry season M. chinensis longeana. The main point of dissimilarity is that in A. drumila the most extreme dry form occurs in the Himalayas and Assam, whereas in M. chinensis the most extreme dry form occurs in Burma.
The species is Indo-Burmese, submontane and apparently not rare where it occurs, but with a restricted distribution. There are two rather doubtfully valid subspecies.
Key to the subspecies of A. (P.) drumila
1 $ dry season form white with black costal and marginal border on the forewing and blackish
costal area on the hindwing drumila drumila (p. 56)
$ dry season form with the white areas sullied with buff scales drumila aphthonius (p. 57)
Allotinus (Paragerydus) drumila drumila (Moore)
(Fig. 37, cf genitalia)
Miletus drumila Moore, [1866]: 777, pi. 41, fig. 12 $. Holotype 'cf ' recte <j>, INDIA: Sikkim (not located).
[Dry season form.] Gerydus drumila (Moore) Moore, 1883: 521. [Wet season form.]
Fig. 37 Allotinus (Paragerydus) drumila drumila (Moore) wet season form multistrigatus de Niceville; Sikkim. Male genitalia.
THE MILETINI 57
Allotinus multistrigatus de Niceville, 1886: 253, pi. 11, figs 11 o" , 2 $ ; de Niceville, 1890: 29, pi. 26, figs 157 Cf, 158 $; Bingham, 1907: 298; Swinhoe, 1910: 195, pi. 615, figs 2, 2b cf , 2a, 2c $ ; Fruhstorfer, 1913: 371; 1916: 815, pi. 141h cf $. LECTOTYPE cf , INDIA: Sikkim (BMNH), here designated [examined] [Wet season form.] [Synonymised by Cantlie, 1963: 26.]
Miletus insignis Staudinger, 1888: 269, pi. 94 'cf' recte $. Holotype $, INDIA: Sikkim (probably MNHU). [Synonymised by de Niceville, 1890: 28.]
Allotinus drumila (Moore) de Niceville, 1890: 28; Bingham, 1907: 297; Swinhoe, 1910: 194, pi. 615, figs 1, Ib Cf, la, Ic $; Fruhstorfer, 1913: 371; 1916: 815, pi. 141i $; Evans, 1932: 211.
Miletographa drumila (Moore) Rober, 1892: 277.
Allotinus multistrigatus multistrigatus de Niceville; Evans, 1932: 242.
Allotinus drumila drumila (Moore) with wet season form multistrigatus de Niceville; Cantlie, 1963: 26, pi. 26, fig. H. 6. 2.
Moore originally described the dry season female in error as the male. Later (1883) he realised his mistake and correctly described the male from the wet season form, which de Niceville later redescribed as a separate species, A. multistrigatus.
Judging by material in BMNH, intermediate forms, with the wing shape of the dry form drumila, fly in the autumn and winter, while drumila, with its mostly white female, is really a spring form flying from February to May (one female ex Fruhstorfer coll. is labelled June, but I feel sure this is an error, as the type-series of multistrigatus was caught in this month).
Moore's type of drumila cannot be traced, but there are two wet season males in BMNH from Darjiling and the Khasi Hills from which Moore described the male, both of which bear BMNH type-labels. But as they did not form part of the original type-series they cannot be accepted as types.
The figures accompanying de Niceville's original description of multistrigatus were taken from examples in coll. Moller, and I designate as lectotype a male in BMNH, presumably from the original type-series, labelled /Type [red]/Sikkim Moller/ Allotinus multistrigatus de Niceville cf TYPE/Rothschild Bequest B.M. 1939-1/.
DISTRIBUTION. Kumaon, in the central Himalayas, to Assam.
Allotinus (Paragerydus) drumila aphthonius Fruhstorfer
Allotinus aphthonius Fruhstorfer, 1913: 371; 1916: 815. LECTOTYPE cf, BURMA (BMNH), here
designated [examined]. [Intermediate form.] Allotinus drumila grisea Riley & Godfrey, 1921: 180. Holotype $, THAILAND (BMNH) [examined]. [Dry
season form.] [Synonymised by Pinratana, 1981: 31.] Allotinus multistrigatus apthonius [sic] Fruhstorfer; Evans, 1932: 242 Allotinus drumila apthonius [sic] Fruhstorfer; Cantlie, 1963: 26; Pinratana, 1981: 31, pi. 4, fig. 19 cf , pi. 5,
figs 10 Cf, 11 $.
The subspecies is doubtfully valid, differing only in that the most extreme dry form of female has the white areas always sullied with buff scales, as in the holotype of grisea and as in Pinratana's figure of the female. The wet season and intermediate forms do not differ from the corresponding forms of subsp. drumila.
In BMNH there are one male and two females labelled as types of aphthonius. I designate as lectotype the male labelled /Type [red]/Type [Fruhstorfer orangeJ/Tenasserim Tandong 4000' Mai Fruhstorfer leg. /Fruhstorfer Coll. B.M. 1933-1/Allotinus aphthonius Frhst. [in Fruhstorfer's hand]/. Two females, similarly labelled, except that one bears a final label /Allotinus aphthonius Fruh. [in Corbet's hand]/ in place of the label in Fruhstorfer's hand, are paralectotypes. The male has a pointed forewing and rather dentate hindwing, and is marked as in the wet season form. Fruhstorfer stated that it was a dry season form, but in fact it is an intermediate form nearer to the wet than to the dry season form. Of the two females, one is intermediate, as in the male, and the other of the normal wet season form.
DISTRIBUTION. Throughout Burma, except in the extreme south of Tenasserim; Thailand.
Genus LOGANIA Distant
Logania Distant, 1884: 197, 208. Type-species: Logania malayica Distant, 1884: 208, pi. 22, fig. 21 $, by
monotypy. Gender feminine. Malais Doherty, 1889: 414, 415, 436. Type-species: Loganiasriwa Distant sensu Doherty, 1889 [= Logania
marmorata Moore, 1884], by designation of Corbet, 1940a: 111. Gender feminine. [Synonymised by de
Niceville, 1890: 32.]
Eyes smooth. Antennae half the length of the forewing costa, with about 36 segments in the type-species;
58 J. N. ELIOT
shorter, with under 30 segments in L. waltraudae; rather longer, with 40-45 segments in the marmorata- group (= Malais}. Nudum extending widely to the base of the shaft. Legs much shorter than in Allotinus, with the tibiae outwardly swollen and the fore-tarsi, except in L. waltraudae, gradually incrassate. In L. malayica and L. waltraudae the male fore-tarsus ends in a rather long, tapered, down-curved point, but in the marmorata-group the fore-tarsus ends abruptly in a rounded pad from which a minute point is directed downwards, as in Allotinus. Labial palpi shorter than in Allotinus, with the third segment usually shorter than one-half of the second segment in males, but may be slightly longer than half in females. Males of all species have a small double hair tuft on the sternum of the eighth abdominal segment. The type-species has the forewing apex produced to a sharp point and veins MI and R5 have a long common stalk; all the remaining species have a rounded or square apex and, except in L. nehalemia, the stalk of veins Mj and R5 is absent or short. There is no trace of a humeral vein on the hindwing. Males of the ma/ay/ca-group have vein A/3 of the forewing unswollen; in the marmorata-group the basal portion of vein M3 is briefly swollen and clothed with small, specialised scales which are about the same size as those of the nivalis-group of Paragerydus. In L. regina, and probably also in L. paluana, the swelling is inconspicuous and partly hidden by normal cover scales. The underside pattern is generally similar to that of Allotinus, but the ground colour is seldom uniform, being mottled in shades of black, brown and white, for which reason Evans coined the popular name 'Mottles' for the genus. On the forewing the usual lycaenid markings may be difficult to make out, but on the hindwing they are usually apparent and the postdiscal series may be catenulate. In some species there is very great individual variation in the extent of whitish scaling on the upperside of the hindwing, which may be absent or cover almost the whole of the wing. The male genitalia are of the usual miletine type and are rather constant in appearance, except that L. waltraudae shows some characters suggestive of Allotinus.
Doherty (1889: 414, 415) erected the generic name Malais and wrote that 'it will include L. marmorata and L. sriwa (probably the same species) and one or two rare kinds undescribed'. He had before him only a single female fronrMergui which he treated with some doubt as the same as the taxon named Logania sriwa Distant, 1886. De Niceville (1890: 33), who had custody of the type of L. marmorata Moore, 1884, stated that he had examined Doherty's female of 'sriwa' and found that 'it appears to differ from L. marmorata only by the greater prominence of all the markings of the underside'. As the females of L. sriwa and L. marmorata differ so strongly it is inconceivable that Doherty's female was really sriwa, and highly probable that de Niceville was correct in assuming that it was L. marmorata. I can find no convincing evidence that L. sriwa has ever been taken in Burma, although it was recorded by Evans (1932) and Cantlie (1963) from Mergui, probably on the basis of Doherty's misidentified female.
Soon afterwards Doherty (1891o: 29) doubted if Malais was distinct from Logania; and though he did not formally synonymise the two he thereafter used Logania for species which would fall naturally into Malais. Bingham (1907: 302), Swinhoe (1910: 200), Evans (1932: 199) and Cantlie (1963: 2) all treated Malais as a subjective synonym of Logania, as did Corbet (1940a: 111) who stated that L. sriwa was its type-species. However, as Doherty's 'sriwa' was misidentified, Corbet's statement cannot, under Article 70 of the Code, be held to constitute a valid type selection. Fruhstorfer (1914; 1915), whilst using Logania as the generic name, employed Malais in a subgeneric sense for the 'Artengruppe' with banded legs and rounded forewing apex which includes L. marmorata and L. sriwa. Hemming (1960: 11), apparently unaware of Corbet's action, designated Logania malayica Distant, 1884, as type-species of Malais. His action is invalid, since malayica was not one of the species originally included in Malais by Doherty; indeed, the latter (1889: 437) had specifically excluded malayica from his new genus, stressing that it was a true Logania. Malais remains, therefore, a valid and available genus-group name, which can be used as a subgenus, largely in the sense in which it was employed by Fruhstorfer, by those authors who consider that the differences between Logania and Malais are greater than those between species-groups. Malais is, however, still without a properly established type-species, and any author intending to use it will have to refer the matter to the International Commission on Zoological Nomenclature, as required under Article 70 of the Code, with a recommendation that Logania marmorata Moore, 1884, be designated as type-species.
The genus ranges from Peninsular India through the Archipelago to New Guinea and the Bismarcks, and comprises 10 species.
Key to the species of Logania
1 cf upperside of forewing with vein M3 unswollen and clothed with normal cover scales, cf
fore-tarsus, so far as known, ending in a tapered, down-curved point 2
Cf upperside of forewing with basal part of vein M3 swollen and clothed with specialised scales.
Cf fore-tarsus ending abruptly , but with a small point directed downwards from its lower edge 4
2 Forewing apex not produced 3
THE MILETINI
59
Forewing apex produced to a sharp point malayica (p. 59)
3 Upperside of hindwing all white waltraudae (p. 61)
Upperside of hindwing with outer half black and inner half white nehalemia (p. 60)
4 Legs banded, with a specially prominent, broad, brown band on outer half of tibiae 5
- Legs freckled, not clearly banded 8
5 $ upperside with grey to white areas except in marmomta diehli 6
- $ upperside entirely brown obscura (p. 68)
6 Forewing very weakly crenulate . Underside of hindwing with a white streak or patch bearing few
if any striae 7
- Forewing termen crenulate. Underside of hindwing without an unstriated area... marmorata (p. 64)
7 Upperside of hindwing pale grey to whitish regina (p. 62)
- Upperside of hindwing white with a broad, black border paluana (p. 63)
8 Forewing termen only very weakly crenulate 9
- Forewing termen crenulate watsoniana (p. 73)
9 Underside mottling more or less ochreous distant! (p. 69)
- Underside mottling dark brown; no ochreous tinge hampsoni (p. 72)
Logania malayica Distant (Figs 38, 39, cf genitalia) Logania malayica Distant, 1884: 208.
The species is instantly recognisable by the pointed and produced forewing apex. The sexes are alike, above white with a blackish forewing border expanding from less than 1-0 mm at the tornus to nearly mid-costa. Underside white densely mottled with reddish brown striae. The legs are buff-brown freckled with whitish scales, and the male fore-tarsus ends in a comparatively long, tapered, down-curved point.
The species occurs from Peninsular Thailand to Malaya, Sumatra, Borneo and the Philippines. There are two subspecies with somewhat different male genitalia.
Key to the subspecies of L. malayica
1 Underside irregularly mottled and blotchy, with, some of the usual lycaenid markings apparent
malayica malayica (p. 60)
- Underside regularly and densely mott; , with none of the lycaenid markings apparent
malayica subura (p. 60)
Fig. 38 Logania malayica malayica Distant; Malay Peninsula. Male genitalia.
60 J. N. ELIOT
Logania malayica malayica Distant
(Fig. 38, cf genitalia)
Logania malayica Distant, 1884: 208, pi. 22, fig. 21 $, text-fig. 61 hind-leg. Holotype $, WEST MALAYSIA:
Sungei Ujong (not located). Logania malayica malayica Distant; Fruhstorfer, 1914: 23; 1916: 805, pi. 141f cf ; Corbet, 1940a: 111, fig. 1
Cf valva; Fleming, 1975: 22, pi. 58, fig. L51 $; Eliot, 1978: 241, pi. 20, fig. 8 cf .
On the underside the mottling is irregular and coalesced into blotches in places, and covers both wings except for a white area above the forewing dorsum in spaces la and Ib. The usual lycaenid markings can be partly made out with difficulty.
DISTRIBUTION. Peninsular Thailand; West Malaysia; Sumatra; Borneo, including Pulo Laut.
Logania malayica subura Fruhstorfer (Fig. 39, cf genitalia)
Logania malayica Distant; Semper, 1889: 160, pi. 31, fig. 3 cf.
Logania malayica subura Fruhstorfer, 1914: 23; 1916: 805. Holotype cf , PHILIPPINES: Mindanao (SM).
On the underside the mottling of striae is regular and without blotches; none of the usual lycaenid markings can be made out. On the forewing the white area above the dorsum extends into space 2. The male genitalia differ more than usual in subspecies, and it may be that subura has achieved species status.
Fruhstorfer named subura from Semper's figure, so the male depicted therein is automatically the holotype.
DISTRIBUTION. Probably throughout the southern Philippines, but I have only seen examples from Mindanao and Samar.
Fig. 39 Logania malayica subura Fruhstorfer; Samar. Male genitalia.
Logania nehalemia Fruhstorfer stat. rev. (Figs 40, cf genitalia; 78 cf )
Logania nehalemia Fruhstorfer, 1914: 25; 1916: 808. Holotype '$' recte cf, NEW GUINEA (BMNH)
[examined]. Logania hampsoni nehalemia Fruhstorfer; D'Abrera, 1971: 384.
Above, the forewing is white with a black border which curves in above the cell to the wing base; the hindwing has the basal half white and the outer half black. The underside is white, with dense, dark brown striae more or less corresponding with the areas which are black on the upperside. In the unique male holotype, which has hitherto been regarded as a female, the legs are missing except
THE MILETINI
61
Fig. 40 Logania nehalemia Fruhstorfer; New Guinea. Male genitalia.
for the femur and tibia of one hindleg with the scales rubbed off; these hardly differ from the femur and tibia of L. malayica. The male genitalia are chiefly distinguished by the phallus, which is considerably stouter than that of its congeners except for L. waltraudae.
The holotype is labelled /Type [red]/New Guinea. Hewitson Coll. 79-69 Miletus 1/9 holotype Logania nehalemia Fruhst. [in Corbet's hand]/. Given its early date of capture, the type-locality is likely to lie in the north-western part of Irian Jay a.
DISTRIBUTION. New Guinea.
Logania waltraudae sp. n.
(Figs 41, cT genitalia; 107 cf )
C? forewing 10-0 mm. Generally similar in appearance to the sympatric L. malayica subura, and like it with the basal portion of vein M3 unswollen and clothed with normal cover scales; but differing in having the forewing apex rounded and veins MI and R5 connate.
Fig. 41 Logania waltraudae sp. n. ; Samar. Male genitalia.
62 J. N. ELIOT
Upperside white; forewing with a blackish brown apical border running from just below vein Cu2 on the termen to just above the cell apex on the costa; hindwing with a dark brown marginal hairline. Cilia dark brown. Underside pale brown very densely mottled with darker brown striae except on the forewing in most of spaces Ib and 2 and in the basal part of space 3, which are white. A dislocated series of postdiscal spots can just be made out on the forewing.
Antennae just under half the length of the forewing costa, thinner than in L. malayica, with probably 28-30 segments (both antennal clubs are broken off just before the tip after 26 segments). The middle shaft segments are just over twice as long as wide (in the remaining Logania species these segments are nearly as wide as long). The nudum extends widely to the base of the shaft, which is brown on the upper surface with a central buff patch on each segment. The palpi are clothed with brown and a few buff adpressed scales and are exceptionally short, not protruding beyond the head, with the third segment comparatively stout and only a quarter the length of the second segment. The legs are about as long as those of L. malayica, which they resemble in having the fore-tarsus ending in a tapered, down-curved point; but they differ in several respects. The femora, tibiae and tarsi are subequal; the femora are broader and somewhat flattened; the foretibiae are narrow and cylindrical, and the middle tibiae are slightly swollen, the swelling being greatest in the upper half. The hind legs are missing. Body dark brown, slightly paler beneath; the abdominal hair tufts on the eighth sternum are smaller than in the other species of Logania. The male genitalia are broadly of Logania type, but the phallus is distinctive, while the triangular vinculum flap and strut running parallel to the lower edge of the uncus plate recall those structures in Allotinus.
$ unknown.
MATERIAL EXAMINED
Holotype cf , Philippines: Samar, 18.viii.l980(C. G. Treadaway) (coll. Treadaway, but will be deposited in due course in SM).
The species, which is named after Mrs Treadaway, occupies an isolated position in the genus, differing from the remainder in palpi, antennae, legs and male genitalia, and possibly deserves to be placed in a separate subgenus.
Logania regina (H. Druce)
(Fig. 42, cf genitalia) Miletus regina H. Druce, 1873: 348.
This and subsequent species differ from the foregoing species in the male fore-tarsus, which is not tapered but ends abruptly in a rounded pad from the lower side of which a small, short point is directed downwards, as in Allotinus. The legs, in this and the next three species, are banded, most prominently on the tibiae, with whitish and brown, while in the remaining species they are freckled and sometimes have obscure longitudinal streaks.
A distinguishing character of L. regina is the white ground colour of the underside of the hindwing, which is almost devoid of striae in a streak-like area running from the wing base to the termen through the upper part of the cell and space 6.
On the upperside the male has a fuscous border on the forewing tending to run narrowly along the costa to the wing base, and a fuscous costal area above vein 6 on the hindwing. The rest of the wings are whitish to bluish grey. The female has a narrower forewing border and the pale areas on both wings are whiter than in the male.
In the female of this species and of L. paluana (infra) the abdomen is longer than in the other Logania species, and extends just beyond the hindwing tornus.
The species has a restricted distribution in Sundaland, excluding Java, Palawan and the islands off the west coast of Sumatra, but has reached the Sulu Is. where it must be a recent immigrant. There are two subspecies.
Key to the subspecies of L. regina
1 cf upperside of forewing with black border comparatively wide, filling whole of space 5. $ forewing border reaching dorsum; underside of forewing with a white area above dorsum usually reaching vein M2 regina regina (p. 63)
- cf upperside of forewing with black border narrower, not reaching base of space 5. $ forewing border fades out at, or just before, tornus; underside of forewing without a white area
regina sriwa (p. 63)
THE MILETINI 63
Fig. 42 Logania regina regina (H. Druce); Borneo. Male genitalia.
Logania regina regina (H. Druce)
(Fig. 42, cf genitalia)
Miletus regina H. Druce, 1873: 348, pi. 32, fig. 4 cT. Holotype cT, BORNEO (BMNH) [examined]. Logania regina regina (H. Druce); Fruhstorfer, 1914: 23; 1916: 806.
Logania evora Fruhstorfer, 1916: 806. Holotype $, PHILIPPINES: 'Sula Is.' recte Sulu Is (SM) [examined] Syn. n.
In the male most, if not all, the pale areas on the upperside are clothed with bluish grey scales. The female is much whiter, with a narrower forewing border which is usually about 1-5 mm wide at the tornus. On the underside of the forewing there is sometimes a narrow whitish area along the dorsum, the corresponding area in the female being much wider and usually extending into the basal half of space 4.
DISTRIBUTION. Borneo; Sulu Is (only known from unique holotype). In BMNH there is a single male from Pulo Laut, in which the fore wing border is even narrower than in subsp. sriwa, while the pale areas are white, so that the general appearance is of a female; it probably represents a further subspecies.
Logania regina sriwa Distant
Logania sriwa Distant, 1886a: 531; 18866: 452, pi. 44, fig. 16 $; Evans, 1932: 212. Holotype $, WEST
MALAYSIA (not located). Logania regina sriwa Distant; Fruhstorfer, 1914: 23; 1916: 805, pi. 141f $; Corbet, 1940«: 112, fig. 2 cf
valva; Cantlie, 1967: 28; Fleming, 1975: 22, pi. 58, fig. L52 ?; Eliot, 1978: 241, pi. 20, fig. 9 cf .
The male is usually whiter, with a narrower border, than nominate regina. In the female the forewing border usually fades out at or before the tornus. On the underside both sexes lack a white area above the forewing dorsum, and on the hindwing the white streak is narrower.
DISTRIBUTION. West Malaysia; Sumatra; peninsular Thailand (Pinratana, 1981).
Logania paluanasp. n.
(Fig- 79 9)
This taxon, at present only known from two females, appears to replace L. regina, of which it may be a subspecies, in Sulawesi. It is larger, with forewing length 17-0 mm, compared with an average of 14-0 mm in regina, and differs additionally as follows. On the upperside the forewing border is narrower, ending beyond the middle of the costa. On the hindwing there is a black border measuring nearly 4-0 mm at the dorsum, expanding to 4-5 mm at vein Cu2 and thence decreasing to 1-0 mm at the apex whence it is continued as a blackish line along the costa to the wing base. The underside is generally marked as in
64 J. N. ELIOT
regina, but on the hindwing the comparatively well-defined white streak ofregina is replaced by a wider and more obscure, because more heavily striated, white discal patch extending to vein Cu\ and the lower edge of the cell. The postdiscal markings are olive-brown and there is a well-defined marginal olive-brown line 0-5 mm wide on the forewing and between veins Cu2 and M2 on the hindwing.
MATERIAL EXAMINED
Holotype 9, Sulawesi: labelled /G. Rangkoean, Paloe, West Celebes, 900', Nov. 1936 (J. P. A. Kalis) (BMNH).
Paratype. 1 9 , data as holotype (BMNH).
Logania marmorata Moore
(Fig. 43, cf genitalia) Logania marmorata Moore, 1884: 22.
The species can be recognised by its crenulate forewing termen and banded legs, which are shorter and stouter than in any other Logania species. Except in the subspecies from Simeulue, both sexes always have a pale area on the forewing, which is more extensive in the female, but in the male of some subspecies it is reduced to a small discal patch. The hindwing of the male is normally brown, at least in the wet season, while that of the female bears some grey scales; but in Nias and south Sumatra the hindwing is partly grey in the male and nearly all white in the female.
The species ranges from central Burma to Vietnam and throughout Sundaland, the Lesser Sunda Is. and the Philippines into north Sulawesi. The dividing line between subspecies is difficult to draw because of the high degree of individual variation; 10 are provisionally recognised.
Key to the subspecies of L. marmorata
1 9 upperside of forewing with basal half whitish 2
- 9 upperside of forewing unmarked brown marmorata diehli (p. 66)
2 (if upperside of hindwing plain brown 3
- cf upperside of hindwing partly white or grey 11
3 cf upperside of forewing with some bluish grey scaling reaching wing base 4
- cf upperside of forewing with brown wing base 6
4 cf upperside of forewing with pale area outwardly white but wing base and space la rather
dark, having many brown scales intermixed with the grey. 9 upperside of hindwing sometimes plain brown 5
- cf upperside of forewing with pale area outwardly white becoming light bluish grey at wing
base and in space la. $ upperside of hindwing with at least some grey scaling below vein 6
marmorata damis (p. 65)
5 cf upperside of forewing with white extending fully across space Ib. $ underside of forewing
without a whitish area beyond cell marmorata hilaeira (p. 65)
- cf upperside of forewing with white area not below midspace Ib. $ underside of forewing with
a whitish area beyond cell extending from vein A\ to veins M2 or MI
dry season form of marmorata javanica (p. 67)
6 9 underside of forewing without a prominent whitish area beyond the cell 7
9 underside of forewing with a broad whitish area beyond the cell stretching from vein A\ to
vein M2.
Cf upperside of forewing with an ovate discal white patch about 2-5 mm wide at base of spaces 4,3,2 and sometimes just entering space Ib marmorata munichya (p. 66)
7 cf upperside of forewing with discal white patch at least 2-0 mm wide, reaching and often
crossing vein Cu2. $ upperside of forewing with base more or less grey-scaled 8
- cf upperside of forewing with discal patch about 1-25 mm wide and not below mid-space 2. 9
upperside of forewing with base brown, bearing only a trace of grey scaling
wet season form of marmorata javanica (p. 67)
8 Cf 9 underside comparatively paler and browner ; base and disc of forewing not blackish 9
Cf 9 underside dark; base and disc of forewing blackish 10
9 cf upperside of forewing with white patch usually not below vein Cu2. Underside comparative-
ly dark. Continental wet season form of marmorata marmorata (p. 65)
Cf upperside of forewing with white patch usually crossing vein Cu2. Underside comparatively light. Palawan marmorata palawana (p. 67)
THE MILETINI 65
10 9 upperside of hindwing with some grey scaling marmorata samosata (p. 67)
- 9 upperside of hindwing brown marmorata Faustina (p. 68)
11 cf upperside of hindwing brown streaked with grey below vein MI. 9 upperside of hindwing
mostly dirty whitish below vein MI dry season form of marmorata marmorata (p. 65)
- cf upperside of hindwing white below vein M\ except for an inwardly diffuse brown border
1 '0-1 -5 mm wide. 9 upperside of hindwing nearly all white marmorata lahomius (p. 66)
Logania marmorata marmorata Moore
Logania marmorata Moore, 1884: 22; 1886: 39, pi. 3, fig. 7; de Niceville, 1890: 33, frontispiece, fig. 128 $ holotype; Bingham, 1907: 303; Swinhoe, 1910: 200, pi. 618, figs 1, la 'cf' recte <j>; Evans, 1932: 213. Holotype $, BURMA: Mergui (ZSI).
[Malaissriwa (Distant) sensu Doherty, 1889: 436. Misidentification.]
Logania marmorata marmorataMoore; Fruhstorfer, 1914: 23; 1916: 806, pi. 141f cf $ ; Corbet, 19400: 112; Cantlie, 1963: 29.
The subspecies occurs in two seasonal forms, at least in the northern part of its range. In the wet season form both wings are brown, with a small white patch on the forewing above vein Cu2 in the male and a much larger white area which is greyish basally in the female. In the dry season the white areas of the forewing are much enlarged in both sexes, the hindwing is more or less overlaid below vein MI with grey or whitish scales, especially in the female, and on the underside of the forewing there is no whitish area beyond the cell. Intermediate season examples may be indistinguishable from subsp. damis.
DISTRIBUTION. Burma, from Karen Hills to Mergui; Thailand; Vietnam.
Logania marmorata damis Fruhstorfer
Logania massalia damis Fruhstorfer, 1914: 24; 1916: 807. LECTOTYPE cf , SINGAPORE (BMNH), here
designated [examined]. Logania marmorata damis Fruhstorfer; Corbet, 1940a: 111; Fleming, 1975: 22, pi. 58, fig. L53 9; Eliot,
1978: 241, pi. 20, fig. 10 cf.
In the wet and only seasonal form the pale areas on the forewing are whiter and more extensive than in any other subspecies except lahomius. In the male the pale area is outwardly white, inwardly rather pale bluish grey, reaches the dorsum and fills the cell; the hindwing is brown. In the female the pale area is whiter and broader, and the hindwing always bears at least some grey scales below vein M\.
I designate as lectotype of damis a male in BMNH labelled /Type [red]/Singapora II. 95/almost certainly type of Logania massalia damis Fruhst. [in Corbet's hand]/.
DISTRIBUTION. Peninsular Thailand; West Malaysia; Singapore; east coastal region of Sumatra.
Logania marmorata hilaeira Fruhstorfer
Logania obscura Distant & Pryer, 1887: 266. Syntypes, BORNEO: Sandakan (not located). [Secondary
homonym of Logania obscura (Rober, 1886).] Logania marmorata hilaeira Fruhstorfer, 1914: 23; 1916: 806; Corbet, 1940a: 112. LECTOTYPE cf,
SUMATRA (BMNH), here designated [examined]. Logania marmorata stenosa Fruhstorfer, 1914: 23 (nomen nudum); 1916: 806; Corbet, 1940a: 112.
LECTOTYPE 9, BORNEO (BMNH), here designated [examined]. Syn. n. Logania marmorata obscura Distant & Pryer; Fruhstorfer, 1914: 23. Logania massalia nada Fruhstorfer, 1914: 24; 1916: 807. LECTOTYPE cf, SUMATRA (BMNH), here
designated [examined]. [Synonymised by Corbet, 1940a: 112.] Logania marmorata cineraria Fruhstorfer, 1916: 806; Corbet, 1940a: 112. [Replacement name for Logania
obscura Distant & Pryer, 1887.] Syn. n. Logania massalia sora Fruhstorfer, 1916: 807. LECTOTYPE cf, BORNEO (BMNH), here designated
[examined]. Syn. n. Logania marmorata sora Fruhstorfer; Corbet, 1940a: 112.
Above, the male has a slightly wider forewing border than subsp. damis, and the base of the wing is darker with brown scales intermixed with the grey. The female generally has a darker hindwing which is often without any grey scales.
I designate as lectotype of hilaeira a male in BMNH labelled /Type [red]/CMB IV.94/Fruhstorfer Coll.
66 J. N. ELIOT
B.M. 1933-131/marmorata hilaeira Frhst. [in Fruhstorfer's hand]/. A female labelled /Type [red]/Type [Fruhstorfer orange]/Sumatra Montes Battak ex coll. Fruhstorfer/marmorata Selesseh 15.VII.94/Fruh- storfer Coll. B.M. 1933-131/$ Allotype of Logania marmorata hilaeira Fruh. [in Corbet's hand]/ is a paralectotype.
In his earlier work (1914) Fruhstorfer did not list or describe Logania marmorata stenosa, either through an oversight or lapsus calami, but he mentioned stenosa twice by comparison with his sub-spp. javanica and samosata. In 1916 he gave a brief description and type-locality, so that the name dates from 1916. I designate as lectotype of stenosa a female in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Sintang Dr. Martin H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/Marmorata stenosa Fr. [in Fruhstor- fer's hand]/.
I designate as lectotype of nada a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/CMB X.94/Fruhstorfer Coll. B.M. 1933-131/massalia nada Frhst. [in Fruhstorfer's hand]/.
I designate as lectotype of sora a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Nord-Borneo ex coll. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/massalia sora Fr. [in Fruhstorfer's hand]/.
DISTRIBUTION. Northern and western Sumatra at least as far south as Padang; Borneo, including Pulo Laut; Cagayan Sulu.
Logania marmorata lahomius (Kheil)
Miletus lahomius Kheil, 1884: 27, pi. 5, figs 28, 29 cf . Syntypes, NIAS (not located). Logania regina lahomius (Kheil) Fruhstorfer, 1914: 23. Logania lahomius (Kheil); Fruhstorfer, 1916: 806. Logania marmorata lahomius (Kheil); Corbet, 1940a: 113.
Above, both sexes are greyish white with the forewing border narrower than in the preceding subspecies. The hindwing of the male has a diffuse fuscous margin about 1-5 mm wide and the costal area is fuscous above vein MI, but in the female the margin is vestigial or absent and there is only a little fuscous dusting below the costa.
DISTRIBUTION. Nias I. There is a pair in BMNH taken by Doherty at Liwa, in the extreme south-west of Sumatra, which differ only that in the female the fuscous scaling below the hindwing costa is solid as far as vein MI; they are provisionally placed under lahomius.
Logania marmorata die/i/i subsp. n.
Cf upperside brown; forewing with a small, sullied, circular whitish patch 2-0 mm wide surrounding swollen portion of vein M3 in spaces 2, 3 and 4. Underside generally pale, with postdiscal markings ill-defined; forewing without a trace of a white or paler discal area, as in the otherwise rather similar subspecies from Java.
$ upperside entirely brown. Underside like male.
MATERIAL EXAMINED
Holotype cf , Simeulue (Simalur): 16-17. ii. 1984 (E. Dieht) (BMNH). Paratype. 1 $ (allotype), data as holotype (BMNH).
The subspecies is named in honour of the captor, Dr Edvard Diehl.
It is instructive that the two extremes of geographical variation in this species are found in the neighbouring islands of Nias and Simeulue, the former having the most extensive white markings, the latter the least; a good example of the haphazard course of evolution in small, isolated populations.
Logania marmorata munichya Fruhstorfer
Logania massalia munichya Fruhstorfer, 1914: 24; 1916: 807. LECTOTYPE cf , JAVA (BMNH), here
designated [examined]. Logania marmorata javanica Fruhstorfer; Corbet, 1940a: 112, partim.
In this and the remaining subspecies the males, at least in the dry season, have a brown forewing bearing a white discal patch and without grey scales at the wing base. In munichya the white patch is 2-5 mm wide and lies at the bases of spaces 4, 3 and 2 and just enters space Ib. The female resembles subsp. hilaeira on the upperside but is distinctive in possessing, on the underside of the forewing, a white area 3-0-4-0 mm wide beyond the cell stretching from vein A± to vein M2.
THE MILETINI 67
It seems likely that this subspecies and the east Javanese subsp. javanica may represent the ends of a cline, in which case Corbet's action in synonymising munichya with javanica would be justified.
I designate as lectotype of munichya a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Java Occident. Sukabumi 2000' ex coll. Fruhstorfer/massalia munichya Frhst. [in Fruhstorfer's hand]/.
DISTRIBUTION. West Java.
Logania marmorata javanica Fruhstorfer
Logania marmorata javanica Fruhstorfer, 1914: 23; 1916: 806; Corbet, 1940a: 112. LECTOTYPE cf , JAVA
(BMNH), here designated [examined]. Logania massalia glypha Fruhstorfer, 1914: 23; 1916: 807. LECTOTYPE cf, JAVA (BMNH), here
designated [examined]. [Synonymised by Corbet, 1940«: 112.] Logania marmorata Moore; Piepers & Snellen, 1918: 18, pi. 20, fig. 21 cf • [Logania massalia Doherty; Piepers & Snellen, 1918: 18, pi. 20, figs 22a cf , 22b $. Misidentification.]
The male differs from subsp. munichya in the smaller white patch on the forewing, which is 1-25 mm wide and does not descend below mid-space 2. The female differs in having the base of the forewing darker and, on the underside, in lacking the prominent white patch beyond the cell, this area being only slightly paler than the rest of the wing.
I designate as lectotype of javanica a male in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/ Ostjava Lawang 1897 ex coll. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/Logania javanica Frhst. [in Fruhstorfer's hand]/. Its left forewing is much discoloured, the basal two-thirds of the wing having an oily bluish sheen; the right forewing is undamaged. The figure in Piepers & Snellen (1918), showing only the left half of a male with a basally bluish forewing, was obviously made from the lectotype; Piepers said that he had himself seen no Javanese examples of L. marmorata. There are also in BMNH two females of the original type-series labelled /Logania javanica Frhst. [in Fruhstorfer's hand]/ which are paralectotypes.
I designate as lectotype of glypha a male in BMNH labelled /Type [red]/Ostjava Lawang 1897 ex coll. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/massalia glypha Frhst. [in Fruhstorfer's hand]/.
All the above lectotypes and paralectotypes appear to be wet season forms. There is in BMNH a series of 4 cf , 7 $ from Sumbawa labelled /Sumbawa Doherty Sept '91/ which must represent the dry season form and which I provisionally place under subsp. javanica. The males have a white patch on the forewing similar to that of subsp. munichya, but the base of the wing is covered with rather dark bluish grey scales. The females also are rather similar on the upperside to that sex of subsp. munichya, but the hindwing is paler brown and may bear some grey scales, while on the underside of the forewing the whitish area beyond the cell is narrower and more sullied. There is also in BMNH a single female labelled /S. Flores xi.96. Dry s. Everett/ which, despite the label appears to be a wet season form as might be anticipated from the date of capture at the change of seasons. Above, it differs only slightly from females of subsp. javanica in having the base of the forewing browner, with only a very few overlying grey scales, while on the underside of the forewing the whitish patch beyond the cell is vestigial. Provisionally I attach it also to subsp. javanica.
DISTRIBUTION. East Java; Lesser Sunda Is. (Sumbawa, Flores).
Logania marmorata palawana Fruhstorfer
Allotinus (Logania) distanti Staudinger, 1889: 93, pi. 1, fig. 3 $. Synt'ypes, PALAWAN (? MNHU).
[Secondary homonym of Logania distanti Semper, 1889.] Logania marmorata palawana Fruhstorfer, 1914: 23; 1916: 806. [Replacement name for Logania distanti
(Staudinger, 1889).]
The male resembles subsp. munichya, but the forewing patch is a little larger and faintly bluish grey. The female is most like subspp. marmorata and hilaeira, especially on the underside, but on the upperside of the forewing the border is a little wider at the tornus.
Single females from Luzon in BMNH and Marinduque (coll. Treadaway) appear to belong to this subspecies.
DISTRIBUTION. Palawan; Balabac I. (coll. Treadaway); Luzon; Marinduque.
Logania marmorata samosata Fruhstorfer
[Logania obscura Distant & Pryer sensu Semper, 1889: 160 partim, pi. 31, fig. 4 $. Misidentification.] Logania marmorata samosata Fruhstorfer, 1914: 23; 1916: 806. Holotype cf , PHILIPPINES: Cebu (SM), [examined].
68 J. N. ELIOT
I have seen only two females and no males from Cebu. The former, which have the lower part of the hindwing lightly grey-scaled, differ additionally from subsp. palawana by a much darker underside with the forewing mostly blackish. A female from Mindoro is similar.
Fruhstorfer named the subspecies from Semper's fig. 4, so the specimen depicted therein is automatically the holotype.
DISTRIBUTION. Cebu; probably Mindoro.
Logania marmorata Faustina Fruhstorfer (Fig. 43, cf genitalia)
[Logania obscura Distant & Pryer sensu Semper, 1889: 160 partim, pi. 31, fig. 5 $. Misidentification.] Logania marmorata faustina Fruhstorfer, 1914: 23; 1916: 806. Holotype 9, PHILIPPINES: Mindanao (SM), [examined].
All the females I have seen from Mindanao, Samar and Leyte have the upperside of the hindwing plain blackish brown; otherwise they do not differ from subsp. samosata. The males resemble subsp. palawana on the upperside, but are readily separable by their much darker blackish undersides.
Fruhstorfer described the subspecies from Semper's fig. 5, so the female from Mindanao depicted therein is automatically the holotype.
DISTRIBUTION. Mindanao; Leyte; Samar (coll. Treadaway); Sulu Is.: Tawi Tawi (Tite, 1969).
Fig. 43 Logania marmorata faustina Fruhstorfer; Mindanao. Male genitalia.
Logania obscura (Rober) (Fig. 44, cf genitalia; 80 cT)
Allotinus obscurus Rober, 1886: 52, pi. 4, fig. 8 cf . Syntypes, SULAWESI (? SMT).
Allotinus martinus Fruhstorfer, 1913: 371; 1916: 814, pi. 141h $. Holotype $, SULAWESI: Buton I. (BMNH) [examined]. Syn. n.
Logania donussa Fruhstorfer, 1914: 24. LECTOTYPE $, SULAWESI (BMNH), here designated [ex- amined]. Syn. n.
Logania distanti donussa Fruhstorfer; Fruhstorfer, 1916: 807.
Logania obscura (Rober) Fruhstorfer, 1916: 807.
Logania marmorata obscurus (Rober); Corbet, 1940a: 112.
In the past L. obscura has been thought to replace L. marmorata in Sulawesi and its satellite islands, and it was treated as a subspecies thereof by Corbet. However, the very recent discovery of L. marmorata in north Sulawesi in a still undescribed subspecies indicates that L. obscura is a distinct species. Its status as such is further confirmed by the following characters. The female is all brown, whereas the male has a white forewing patch - a reversal of the usual sexual differences in Logania marmorata wherein the female has
THE MILETINI
69
Fig. 44 Logania obscura Rober; Sulawesi. Male genitalia.
more extensive white areas than the male (except in Simeulue). The forewing termen and apex are more rounded and the crenulations are weaker. The legs are longer and thinner. The wing span is considerably larger. Finally, there is a small difference in the male valva.
The holotype of martinus has no head, legs nor right forewing. It bears a label /damaged in shelter by burst pipe, G.E.T./. The remaining wings show no significant difference to normal L. obscura.
I designate as lectotype ofdonussa a female in BMNH labelled /Type [red]/Type [Fruhstorfer orange]/S. Celebes Bua-Kraeng 5000' Febr. 1896 H. Fruhstorfer/Fruhstorfer Coll. B.M. 1933-131/Logania donussa Fr. [in Fruhstorfer's hand]/.
A single male from Banggai I. in coll. Bedford Russell (Fig. 80) has the forewing patch sullied all over with fuscous scales, so that it hardly shows up, and the underside is darker and more densely striated than in typical examples. A male from the Toekan Besi Is. in BMNH resembles the Banggai male on the upperside, but on the underside is paler than typical examples. These two males probably represent further minor subspecies.
DISTRIBUTION. Sulawesi.
Logania distant! Semper (Figs 45, 46, cf genitalia; 81 cf ) Logania distanti Semper, 1889: 161.
In this species I combine the subspecies which have hitherto been treated as pertaining to two distinct species, L. distanti and L. massalia. Fruhstorfer, and also Corbet in part, confused the massalia subspecies-group with L. marmorata, but the two species are easily distinguished by several characters. Firstly, in L. distanti the legs are longer, thinner and not banded, but speckled and streaked with brown on a whitish ground. Secondly, the forewing termen is barely crenulate. Thirdly, the underside is more ochreous and the striations are usually denser. In the male genitalia the phallus is much less strongly bent.
The male is brown above, normally with a whitish or bluish grey discal patch on the forewing, but in the Philippines and Malay Peninsula (and probably elsewhere) the upperside is sometimes unmarked. On the forewing vein M3 is briefly swollen and clothed with specialised scales. Females show very great individual variation, the forewing white or very pale greyish blue with a broad blackish border, while the hindwing, except in subsp. staudingeri, varies from brown to mainly whitish in any one area.
The species flies from India to the Malay Peninsula, Sumatra, Borneo and the Philippines. It is apparently absent from the islands off the west coast of Sumatra and from Java, records of L. massalia from the latter island applying to misidentified L. marmorata.
Key to the subspecies of L. distanti
1 cf with upperside of forewing brown to brownish at base. $ with upperside of hindwing variable, but in the most lightly marked examples always with a fuscous border 1-0-2-0 mm wide .. 2
70 J. N. ELIOT
Cf upperside of fo rawing with pale bluish grey area reaching wing base . $ upperside of hindwing all white except for sparse fuscous dusting above vein A/i distant! staudingeri (p. 72)
2 Cf upperside of forewing with discal patch, if present, pale bluish grey 3
Cf upperside of forewing with discal patch , if present , white distant! massalia (p . 70)
3 Philippine distant! distanti (p. 70)
Bornean distanti drucei (p. 71)
Logania distanti distanti Semper
(Fig. 81 Cf )
Logania distanti Semper, 1889: 161, pi. 31, figs 6, 7 $, 21 cf. Holotype <j>, PHILIPPINES: Cebu (SM)
[examined].
Logania distanti distanti Semper; Fruhstorfer, 1914: 24; 1916: 807. Logania distanti apsines Fruhstorfer, 1914: 24; 1916: 807. LECTOTYPE cf , PHILIPPINES: Mindanao (SM),
here designated [examined]. Logania turdeta Fruhstorfer, 1916: 807. Holotype 'cf' recte $, PHILIPPINES: Cebu (SM) [examined].
Syn. n.
Semper originally described L. distanti from three examples: a male and female of large size from Mindanao, which he figured at figs 21 and 7 respectively, and a small female from Cebu figured at fig. 6. Although he labelled the male as type he unfortunately did not specify it as such, and this left the way open for Fruhstorfer later to restrict nominate distanti to Cebu and to name the Mindanao pair as subsp. apsines. Semper's female from Cebu is, therefore, automatically the holotype of distanti.
Fruhstorfer 's action, taken at a time when it is evident that he had not even seen Semper's specimens, is doubly regrettable, since a year later, after examining Semper's collection, he named the Cebu female, which he then inexplicably mistook for a male, as a new species, L. turdeta, and remarked that Semper had confused it with L. distanti. At the same time he still recorded nominate distanti from Cebu and maintained apsines for Mindanao examples. Semper's original female from Cebu, which is the holotype of both distanti and turdeta, is labelled /CEBU/Coll. C. Semper/206/1004/No 6/Typus [and on reverse] Logania turdeta (Fruh.)/. The small ticket reading 'No 6' obviously refers to Semper's fig. 6. The specimen has extensive whitish dusting on the upperside of the hindwing and agrees fairly well with the butterfly figured by de Niceville (1894: pi. 2, fig. 13) as Logania luca.
Semper's pair from Mindanao may or may not be a valid subspecies; but as size, by which alone Fruhstorfer separated them as subsp. apsines, is an unreliable character. I treat apsines provisionally as a synonym of distanti. Fruhstorfer did not designate a type, so I now designate the male as lectotype; it is labelled /Log. Distanti typ. Semper/206/1004/16/Typus [red]/. The small ticket reading '16' should refer to the figure in Semper's pi. 31; but in fact the specimen is shown at fig. 21, while