Table Of Content蝶と蛾 Lepidoptera Science 67（1）: 1-6, May 2016
Pepliphorus euchylas Hübner, [1819]: a junior objective synonym of Papilio hylassus Herbst, 1800
（Lepidoptera, Lycaenidae）
Alan CASSIDY
18 Woodhurst Road, Maidenhead, Berkshire, SL6 8TF, United Kingdom
Abstract The origins of the taxa Papilio hylassus Herbst, 1800, and Pepliphorus euchylas Hübner, [1819], are
discussed. Their synonymy and the seniority of hylassus are established. A revised taxonomy/synonymy for Jamides
hylassus（ Herbst）, comb. nov. is recommended.
Key words hylassus Herbst, Jamides euchylas, Lepidoptera, Lycaenidae, priority, synonymy.
Discussion Plate 363 of Stoll, Herbst correctly quoting figures E and F,
18th/19th Century Authors while Hübner demonstrates an early typographic error by
quoting figures B and F. However, there can be no doubt that
Three Lycaenid taxa were described in the 18th Century using
they are both referring to the same taxon, which they both also
the binomial name Papilio hylas. The subject taxon of this paper,
ascribe to Pieter Cramer rather than Caspar Stoll.
authored by Stoll [1781] pp. 142-143（ Fig. 1） was the third
published, and thus a junior synonym in need of replacement. Herbstʼs plate does, however, contain one error and this may
The type locality of Papilio hylas Stoll is the Moluccan Island of
Ambon, the known distribution of this species is still restricted
to the islands of the Indonesian province of Maluku. The earlier
taxa sharing the epithet hylas were created by Denis &
Schiffermüller, 1775, and Esper, [1778]. Fig. 2 shows Stollʼs
accompanying drawing of the species.
Two such replacement names can be found in the literature:
Papilio hylassus Herbst, 1800（ Figs 3,4） and Pepliphorus
euchylas Hübner, 1819（ Fig. 5）. Both authors refer directly to
Fig. 3. Herbstʼs illustration of P. hylassus, referred erroneously
to P. hylax.
Fig. 1. Caspar Stollʼs description on pages 142-143 of Papilio
hylas.
Fig. 2. Stollʼs Plate 363: upperside（ E） and underside（ F） of Fig. 4. Herbstʼs replacement name Papilio hylassus, referred to
Papilio hylas. “Tab 363”, figs E, F of Stoll.
2 A. CASSIDY
Fig. 6. Kirbyʼs 1871 listing of “E. euchylas”, with the location
referenced to current Maluku.
Fig. 5. Hübnerʼs listing of Pepliphorus euchylas referred to Pl.
“363”, figs B（ recte E）, F of Stoll.
have been a cause for uncertainty in subsequent years. He refers
to “Cramerʼs” taxon, which he is replacing, as “hylax” rather than
“hylas”. Papilio hylax Fabricius, 1775, is a valid name for
another Polyommatine Lycaenid, currently treated as Zizula
hylax, a cosmopolitan species distributed in Africa, South and
South-East Asia and Australasia. However, the images clearly
equate to Stollʼs hylas.
William Forsell Kirby
In 1871 William Forsell Kirby produced the first version of his
synonymic catalogue of the diurnal lepidoptera and from page
345 listed 325 “species” under the generic name Cupido Schrank.
Fig. 7. Kirbyʼs introduction to his 1872 paper.
Number 9 on this list is “E. euchylas Hübn.”. This is the only
row of the 325 in which Hübner uses a capital “E” for the generic
abbreviation and I conclude that this is a printerʼs error（ Fig. 6）.
Shortly after publication of this first list, however,
correspondence with A. G. Butler makes Kirby aware of the
publication of Herbstʼs work in Volume 10 of Natursystem aller
bekannten in und auslandischen Insekten, a serial publication of Fig. 8. Kirbyʼs 1872 correction showing the priority of
which earlier volumes were authored by Carl Gustav Jablonsky. hylassus over euchylas.
So in 1872 Kirby published a paper in Transactions of the
Entomological Society of London in which he reviewed the
names published in all 10 volumes of the Jablonsky/Herbst
series.
It is apparent to Kirby that Jablonsky and Herbstʼs work has not
been widely seen. He starts his paper as shown in Fig. 7.
Kirby gets eventually to Volume 10 of Jablonsky and Herbst and
then clearly corrects his earlier list and restores the usage of
Fig. 9. Kirbyʼs 1877 Supplement entry regarding hylassus and
Herbstʼs hylassus over Hübnerʼs Pepliphorus euchylas when
euchylas.
considering the replacement of hylas St[oll]（ Fig. 8）.
20th Century Considerations
Kirbyʼs synonymic list is published again in 1877, this time with
Charles Swinhoe
a Supplement containing new taxa and corrections made since
1871. Here he again lists hylassus Herbst, 1800, combined this 1916 saw the publication by Charles Swinhoe of a new taxon
time with the genus Plebeius, but still clearly placing Hübnerʼs Jamides gamblea, from Maluku, Ceram, which he compares to
euchylas as a junior synonym（ Fig. 9）. two related taxa aleuas（ mis-spelt “alenas”）, Felder, and
hylassus Herbst. This appears to be the last time that Herbstʼs
taxon is mentioned in the literature with any kind of assurance
Papilio hylassus and Pepliphorus euchylas 3
（Fig. 10）.
Charles Bridges, 1988 and 1994, in his Catalogue of
Lycaenidae... lists hylassus Herbst, 1800, as a species-group
taxon but is uncertain about its generic placement. He also notes
Kirbyʼs association of hylassus with euchylas but makes no
further comment（ Fig. 11）.
Furthermore, Bridges lists the synonymy of Jamides euchylas Fig. 11. Bridges, 1994, pages VIII.214, IX.59, listing of
without any mention of hylassus: hylassus Herbst.
euchylas（ Hübner）, [1819], 1816-1826; Pepliphorus [original
genus]
- - - - macromarginata（ Verity）, 1926, 1926-1933.（ RA）;
Agriades
- - - - micromarginata（ Verity）, 1926, 1926-1933.（ RA）;
Agriades
- - - - ab. plumbeus（ Rothschild）, 1915; Pepliphorus Fig. 12. Waterhouse and Lyell, 1914, listing of Pepliophorus
[sic] hylas Stoll [Cramer].
- - - - postmargarita（ Verity）, 1943,（ IFS）; Lysandra
- - - - praepanoptes（ Verity）, 1928.（ RA）; Turania Australia and included taxa from the New Guinea region.
- - - hylas（ Stoll）, [1781], 1780-1790（ H）; Papilio Among these was included Pepliophorus [sic] hylas Hübner,
showing ignorance of Herbstʼs hylassus and probable
- - gamblea Swinhoe, 1916 misunderstanding of Hübnerʼs erection of euchylas（ Fig. 12）.
- - hylais（ Godart）, [1824], 1819-1824; Polyommatus
Hans Fruhstorfer
- mimetica Tite, 1960
Hans Fruhstorfer was a prolific collector and author, publishing
Thus, only Kirby, of all 19th Century authors, appears to have
numerous descriptive papers in the late 19th and early 20th
noted the significance of Herbstʼs taxon hylassus. Swinhoe is
centuries. He described numerous taxa as new, although many
aware of it in passing. Bridges, “re-discovering” it more than 100
have since been shown to be synonymous with earlier
years after Kirby, leaves the treatment of hylassus open to
descriptions. Fruhstorfer was also early to appreciate the value
question. The historical bibliography of euchylas Hübner is
of the male genitalia in distinguishing species of similar
much more comprehensive.
appearance and in 1916, the same year that Swinhoe described
Jamides gamblea, published a revision of the genus he called
Australian Butterflies
Lampides, but which also included all the taxa referred at that
In 1914 Waterhouse and Lyell published The Butterflies of stage to Jamides.
Fruhstorfer lists Lampides euchylas（ Hübner） as a valid species,
along with 8 other taxa which he considers to be subordinate to
euchylas. However, he makes no mention whatsoever of
hylassus Herbst, nor of any other Herbst/Jablonsky taxa（ Fig.
13）.
Fruhstorfer wrote extensively for Lord Walter Rothschild, whose
collection, kept in England, was later bequeathed to the Natural
History Museum in London. This collection, its curation and the
many taxa described from it, became a significant resource for
subsequent authors in the remaining years of the 20th century,
compounding the error of omission of hylassus.
Fruhstorfer makes reference to the homonymy of the taxa hylas
of ESPER, 1777 and [Stoll] Cramer, 1781, and to Hübnerʼs new
Fig. 10. Swinhoeʼs 1916 mention of hylassus when describing name euchylas. His following paragraph might be translated as:
Jamides gamblea.
4 A. CASSIDY
Although the name Papilio hylas ESP[ER]. is now transferred to has fallen out of common usage, through neglect or
Lycaena, I nevertheless hereby replace Pap[ilio] hylas misunderstanding. In its place, the Hübner name euchylas has
CR[AMER] by the Hübner name [euchylas], in order to prevent been in more common usage, especially following its re-
mistakes, which might still develop, because the old Genus publication by Fruhstorfer in 1916. However, the listing of
Lycaena（ Cupido, Plebejus） has now been divided, probably hylassus Herbst, 1800, by Kirby, 1871 & 1877, Swinhoe, 1916,
finally, into separate genera. and Bridges, 1988 & 1994, support the continued use of hylassus
as the correct, senior synonym for this taxon.
Fruhstorfer is unaware of, or ignoring, Herbstʼs creation of his
replacement name hylassus and Kirbyʼs later reference to it. He
Recommendation
is, apparently, not in correspondence with Swinhoe who writes
contemporaneously about hylassus. In this respect Fruhstorfer is The synonymy of this species, as currently recognised by the
followed by all the subsequent authors listed in the next author, is:
paragraphs.
Species:
Gerald Edward Tite Jamides hylassus（ Herbst, 1800） comb. nov.
Tite was an Experimental officer in Department of Entomology =Papilio hylas Stoll, [1781] 4: 142, pl. 363, figs. E, F.（ Nom.
at NHM following World War 2. His extensive work on the
Lycaenidae is generally considered by later experts as thorough
and reliable. In 1960, he published a review of what he titled
“The Jamides euchylas Complex”（ Fig. 14）.
It is of particular note that Tite synonymised gamblea Swinhoe
with euchylas, despite the mention of hylassus by Swinhoe in his
description of gamblea. This is indicative of the blinkered usage
of euchylas since Fruhstorfer. The title of Titeʼs paper makes it
clear that he was unaware of the significance of Herbstʼs
hylassus. This error has been perpetuated by a number of
subsequent authors, at least those listed here: Fig. 13. Fruhstorfer, 1916, history of Lampides euchylas, no
mention of hylassus Herbst.
・DʼAbrera, B. 1971, 1977: 356.
・Kato, S. 1982: 93.
・Hirowatari, T. 1992: 49.
・Bridges, C., 1988: II. 53; 1994: IX. 59.
・Rawlins, Cassidy, Müller, Schröder and Tennent, 2014: 34.
Conclusion
Since the early 20th century, the name hylassus Herbst, 1800,
Fig. 14. Tite, 1960, synonymy of Jamides euchylas.
Fig. 15. Jamides hylassus Figs: 1a-c ♂, 2a-c ♀ ssp mimetica, Buru, 3a-b ssp hylassus ♀,
♂, Seram.
Papilio hylassus and Pepliphorus euchylas 5
praeocc.） Polyommatini of the Oriental and Australian regions
（Lepidoptera, Lycaenidae, Polyommatinae）. Bull. Univ.
=Pepliphorus euchylas Hübner, [1819]: 71.
Osaka Pref（. B）, 44（ Suppl.）: 1-102.
=Polyommatus hylais Godart, [1823]: 654. Maluku, Ambon. Hübner, J., 1816 - 1826. Verzeichniss bekannter Schmettlinge.
Synonymised by Tite, 1960. Augsburg. 431+72 pp.: 1-16, 1816; 17-176, 1819; 177-208,
1820; 209-256, 1821; 257-304, 1823; 305-31, 1825; Anzeiger
=Cupido euchylas（ Hübner）; Kirby, 1871: 346.
1-72, 1826.
=Lampides euchylas（ Hübner）; Fruhstorfer, [1916]: 33, fig. 22. Kato, S., 1982. Notes on the Lycaenidae from Ambon Island,
Indonesia. Tyo to Ga 32（ 3,4）:193-202, 63 figs.
=Pepliophorus [sic] hylas（ Hübner）; Waterhouse & Lyell, 1914:
Kirby, W. F., 1871. A synonymic catalogue of the diurnal
102. lepidoptera. Supplement. Van Voorst, London, 8vo, 690pp.
Kirby, W. F., 1872. Notes on the Diurnal Lepidoptera described by
=Pepliphorus hylas ab. plumbeus Rothschild, 1915: 138.
Jablonsky and Herbst, in their “Natursystem aller bekannten
Synonymised by Tite, 1960.
Insekten.” Trans. Entomol. Soc. London 1872: 111-120.
=Jamides gamblea Swinhoe, 1916: 209（ ♂）. Synonymised by Kirby, W. F., 1877. A synonymic catalogue of the diurnal
Tite, 1960. lepidoptera. Van Voorst, London, 8vo, Supplement pp. 691-883.
Rawlins, A. A. Cassidy, C. Müller, S. Schröder and W. J. Tennent,
=Jamides euchylas euchylas（ Hübner）; Tite, 1960: 322.
2014. An Illustrated and annotated checklist of Jamides Hübner,
Subspecies: 1819, taxa occurring in the Indonesian Provinces of North
Maluku and Maluku（ Lepidoptera: Lycaenidae）. Nachr.
Jamides hylassus hylassus（ Herbst, 1800）. Maluku, Ambon.
entomol. Ver. Apollo, N.F. 35（ 1/2）: 5-39.
Jamides hylassus mimetica Tite, 1960. comb. nov. Maluku, Stoll, C., 1780-1790, in Cramer, P. De Uitlandsche Kapellen
Buru. voorkomende in der drie Waereld-Deelen Asia, Afrika en
America（ etc）. Amsterdam, Baalde & Utrecht, 4 vols（ 252 pp.
Images of the subspecies are shown in Fig. 15.
400 pls.） and Suppl.（ 184 pp. 42 pls.） Vol 1; Vol 2; Vol. 3; Vol.
4; Suppl. [see Appendix 3, Opinion 516, ICZN. for precise
Acknowledgments dates of publication of the various parts.]
Swinhoe, C., 1916. New Indo-Malayan lepidoptera. Ann. Mag. nat.
I am grateful to Dr Gerardo Lamas, of Lima, Peru, for
Hist（. 8）18: 209-211.
information concerning the publication of Papilio hylassus by
Tite, G. E., 1960. The Jamides euchylas complex（ Lepidoptera:
Herbst, 1800, and to NHM London for reproduction of the image
Lycaenidae） Bull. Br. Mus. nat. Hist. Ent. 9（ 5）: 321-332.
of the Type specimen of mimetica in Fig. 15.
Waterhouse, G. A. and G. Lyelle, 1914. The Butterflies of Australia.
A monograph of the Australian Rhopalocera. 238 pp., 888 figs.
References Sydney.
Bridges, C. A., 1988. Catalogue of Lycaenidae & Riodinidae
（Lepidoptera: Rhopalocera）. 816pp. Bridges, Urbana, Illinois. 摘 要
Bridges, C. A., 1994. Catalogue of the family-group, genus-group
Pepliphorus euchylas Hübner,［ 1819］はPapilio hylassus
and species-group names of the Riodinidae & Lycaenidae
Herbst, 1800の新参主観シノニムである（鱗翅目，シジミ
（Lepidoptera） of the world. 1113pp. Bridges, Urbana, Illinois.
チョウ科）（Alan CASSIDY）
DʼAbrera, B., 1977. Butterflies of the Australian Region, edn 2. 415
pp. Melbourne. 18世紀後半にPapilio hylas という学名が3種のシジミチョ
Denis, M. and I. Schiffermüller, 1775. Ankündung eines ウに与えられたため，2種については置換名を与える必要
systematischen Werkes von den Schmetterlingen der が生じた．本論文で扱うのは，そのうちモルッカ諸島のア
Wienergegend. Vienna, Bernardi. 322 pp. 2 pls. ンボン島をタイプ産地とするPapilio hylas Stoll [1781]（ 現
Esper, E. J. C., 1776–1830. Die Schmetterlinge in Abbildungen nach 在はルリウラナミシジミ属のJamides euchylas（ Hübner）と
der Natur. Europäische Schmetterlinge, Leipzig: T.O. Weigel. されるタクサ）である．
Vol ʼʼʼ1ʼʼʼ Tagfalter: Part（ 1）: 388pp; Part（ 2）: 190pp; Plates:
Papilio hylas Stollの置換名としてPapilio hylassus Herbst,
1-123.
1800 と Pepliphorus euchylas Hübner, 1819が提唱されたが，
Fruhstorfer, H., [1916]. Revision der Gattung Lampides auf Grand
前者は20世紀初頭以降ほとんど使用されなかった．一方，
anatomischer Untersuchungen. Arch. Naturgesch（. A）81（6）:
hylassus Herbst にまったく言及しなかったFruhstorfer
1-46, 1 text-fig., 4 pls.
（1916）に従う形で後者が使われることとなった．”The
Herbst, J. F. W., 1800. Natursystem aller bekannten in und
Jamides euchylas complex”を出版したTite（ 1960）以降，多
auslandischen Insecten. Der Schmetterlinge Vol. 10. Berlin.
Hirowatari, T., 1992. A generic classification of the tribe くの文献でこのタクサにはJamides euchylas（ Hübner）が使
用されてきたが，実のところ，忘れ去られたhylassus
6 A. CASSIDY
Herbstはまったく使用されなかった訳ではなかった． これらのことから，Kirby（ 1871 & 1877）などの扱いを尊
Kirby（ 1871 & 1877）は hylassus Herbstをシノニミックリ 重し，近年Jamides euchylas（ Hübner, 1819）とされていた
ストにあげており，euchylas Hübner が新参異名であること タクサをJamides hylassus（ Herbst, 1800）（新結合）の新参
を認識していた．また，Swinhoe（ 1916）はモルッカ諸島 客観異名とみなし，後者を用いることが推奨される．
のセラム島からJamides gambleaを記載した際にhylassus
［文責：広渡俊哉/Toshiya HIROWATARI］
Herbstと比較を行っていた．さらに，Bridges（ 1988 &
1994） はhylassus Herbstを”species-group taxon”として掲載
していた． （Received October 18, 2015. Accepted December 12, 2015）
