Description of a new species of Phassus Walker, 1856 from Costa Rica, Pallas, gen. n., with a new species from Guatemala, and taxonomic notes on Sthenopis Packard, [1865] (Lepidoptera, Hepialidae)

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December, 28, 2015 Vol. 7, No 4

113The European Entomologist, Vol. 7, No. 4
Description of a new species of Phassus Walker, 1856 from Costa Rica,
Pallas, gen. n., with a new species from Guatemala, and taxonomic
notes on Sthenopis Packard, [1865] (Lepidoptera, Hepialidae)
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Abstract
Phassus violetteae sp. n. from Costa Rica is described as new and compared to its
putatively close relatives, Phassus triangularis Edwards, 1885 and Phassus huebneri
*H\HU>@7KHKRORW\SHPDOHZLOOEHGHSRVLWHGLQ,1%,2,QVWLWXWR1DFLRQDOGH
Biodiversidad, Santo Domingo de Heredia, Costa Rica. Pallas gen. n. is erected for
Pallas reynaudi sp. nW\SHVSHFLHVIURP*XDWHPDOD,WGLIIHUVIURPLWVPRUSKRORJLFDOO\
closest genera Phassus Walker, 1856 and Schausiana9LHWWHPDLQO\E\WKHODELDO
palp segments, absence of an androconial brush in the male metatibia and genital
characters in both sexes. The holotype female of Pallas reynaudi sp. n. will be deposited
in Universidad del Valle de Guatemala, Guatemala. Hepialus auratus Grote, 1878 syn.
n. and Phassus eldorado3¿W]QHUsyn. nDUHV\QRQ\PL]HGWRSthenopis pretiosus
+HUULFK6FKDIIHU>@comb. n. The male holotype of Phassus eldorado3¿W]QHU
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Keywords:ELRJHRJUDSK\&HQWUDO$PHULFD'1$EDUFRGHPRUSKRORJ\1HDUFWLF1HRWURSLFDO
Introduction
The phylogeny and taxonomy of many groups within the Hepialidae remain unresolved.
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and recent revisions of Aepytus Herrich-Schäffer, [1856] and descriptions within &LE\UD
Walker, 1856 have been undertaken (Mielke 2014, 2015; Mielke & Casagrande 2013,
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comprise two or three broad groups: Phassus Walker, 1856/Schausiana 9LHWWH
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number of other undescribed species that may belong to either of these groups or
represent other, related groups.
The phylogenetic status of Phassus and Schausiana has not yet been comprehensively
assessed and a future phylogenetic and taxonomic analysis is highly desirable as
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114 The European Entomologist, Vol. 7, No. 4
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Phassus by wing vein characters, the fused patagia with three spines on each side, the
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Genital characteristics within Phassus may provide evidence for monophyly but these
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7KHIRUHZLQJSDWWHUQ RIPhassus/Schausiana appears to be distinctive, at least
for some species and this may prove to be phylogenetically informative.
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on external and genital characteristics. Either or both sets of features were examined
for all previously described species of Hepialidae recorded from Mexico and Central
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but his description of the wing colour as monotonous dull, greyish-brown does not
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features for 3URVXOHQWXV, but the description does not match any of the newly described
species here. In addition, the species was described from Mexico whereas our species
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notes on the taxonomy of Phassus and Sthenopis Packard, [1865], and consider some
of the biogeographic implications for the distribution of Phassus, Schausiana, and the
newly described genus.
Material & Methods
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concluded that they represented new species. While examining one of these new taxa,
several diagnostic characters could not be matched to any of the available genera, and
so a new genus is erected.
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115The European Entomologist, Vol. 7, No. 4
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projects and in GenBank (access codes for each specimen cited in the list of material
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relationships among the specimens analysed. Four nucleotide sequences (one of Pallas
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Abbreviations
The following abbreviations for collections are used in the text:
&*&0 &ROO&DUORV*&0LHONH&XULWLED3DUDQi%UD]LO
CGR Coll. Gérard Reynaud, Paris, France.
,1%,2
Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica.
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SMFL
Senckenberg-Museum, Frankfurt am Main, Lepidoptera collection, Germany.
UVG Universidad del Valle de Guatemala, Guatemala, Guatemala.
ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany.
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Taxonomy section
Pallas C. Mielke & Grehan gen. n.
Type species: Pallas reynaudi sp. n., by present designation.
Etymology: It is named for a son of Lycaon, as is PhassusDOWKRXJK:DONHUGLG
not give evidence that the genus-group name Phassus was derived in this way.
Description:
Pallas gen. n comprises the type species only. Medium to large moths wingspans of
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Venation similar to Phassus³KHSLDOLQH´SDWWHUQ'XPEOHWRQZLWK5V5V
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the wing outer margin. ƃ metatibia without androconial organ. Scutum III devoid of
scales. ƃand ƂWHUJXP9,,,VFOHURWL]HGQRWGLIIHUHQWLDWHGVWHUQXP9,,,UHGXFHG)RU
genitalia details, see description of the new species.

116 The European Entomologist, Vol. 7, No. 4
Diagnosis:
Pallas gen. nVXSHU¿FLDOO\UHVHPEOHVWKHH[WHUQDOPRUSKRORJ\RIODUJHERGLHGVSHFLHV
of Phassus and Schausiana. However, several characters are diagnostic for Pallas gen.
n.: partial fusion of the second and the third segments of the labial palps, presence of
Sc1 (at least absent in PhassusDEVHQFHRIWKHDQGURFRQLDORUJDQRQWKHƃ metatibia,
and in the ƃJHQLWDOLDWKHUHLQIRUFHG VFOHURWL]DWLRQRIWKH EDVHRIWKHYDOYD DQGWKH
pseudoteguminal plates fused dorsally and ventrally.
Remarks:
The wing pattern, wing venation, and genital characteristics of Pallas gen. n. are
VXI¿FLHQWO\GLVWLQFWIURPWKRVHRIPhassus and Schausiana to justify its description as
a new genus, although it is possibly a sister group to a Phassus/Schausiana clade.
Pallas reynaudi sp. n.
Figs. 1a, 1b, 2a, 2b, 3, 13, 14a, 14b, 15a, 15b, 16, 17, 23, 24.
Holotype ƂZLWKWKHIROORZLQJODEHOVVHSDUDWHG E\IRUZDUGVODVKHVHolotypus,
Pallas reynaudi C. Mielke & Grehan det. 2015/ Guatemala, Huehuetenango,
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Paratypes LQ WRWDO ƃƃ ƂƂ $OO Guatemala ƃƃ ƂƂ >&KLPDOWHQDQJR
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6>OHJ@=0+%ƃƂƂZLWKVDPHGDWDDVWKHKRORW\SH&*&0
Ƃ VDPHGDWDDVWKH KRORW\SH&*5Ƃ +XHKXHWHQDQJR6DQ0DWHR
,[WDWiQ  NP 1 %XOHM  P 9 &DPSRVHFR< 0RQ]yQ OHJ &*&0
>%&-;ES@
Etymology. P. reynaudi sp. n LV QDPHG IRU 0U *pUDUG 5H\QDXG 3DULV IRU KLV
motivation and enthusiasm for Lepidoptera and lepidopterists.
ƃ)LJVDEDE6HHDOVRGHVFULSWLRQRIWKHJHQXV):OHQJWKPP
ZLQJVSDQPP$QWHQQDZLWKVHJPHQWV(SLSK\VLVSUHVHQWVWRXWDQGVKRUW):
costal margin convex at the base, apex slightly falcate, outer margin curved. Thorax dark
brown with whitish collar anteriorly, metathorax light brown without scales anteriorly.
FW dorsal ground colour brown to light brownish grey to light brown; basoproximal,
basodistal, and postdiscal areas pale brownish grey; basocentral, central patch, and
premarginal areas brown reaching the costal margin darker or black; submarginal and
marginal bands merge into both patterns, marginal area represented only by rounded
and disrupted spots at each vein; basocentral area and central patch fused posteriorly
forming the typical V-shaped band highlighted by a thin black line internally as the
premarginal area proximately. Consequently, basoproximal generally fused to the
postdiscal area, the former with black dots also present at the base of the V. Stigma

117The European Entomologist, Vol. 7, No. 4
bright pale yellow as the conspicuous spots present in the basocentral, premarginal, and
submarginal areas. HW dorsally brown, lighter at the base, with some ornamentation
along costal margin distally and outer margin anteriorly. Tergosternal bar simple, well
VFOHURWL]HG QDUURZ DQG UHFWDQJXODU ZLWK FHQWUDO EUHDN DW ULJKW DQJOH WR WKH DQWHULRU
PDUJLQ)LJ$EGRPLQDOVHJPHQWVZLWKWXEHUFXODWHSODWHVZHOOPDUNHG)LJVD
E Tergum VIII rectangular, sternum VIII smaller and triangular, wider towards
SRVWHULRUPDUJLQ)LJE
ƃJHQLWDOLD )LJVD ESaccus U-shaped, slightly expanded anteriorly; postero-
dorsal edge with convex margin laterally and mesally with a deep invagination.
7HJXPHQYHUWLFDOIXVHGODWHUDOO\WRWKHSVHXGRWHJXPLQDOSODWH7HUJDOOREHVKRUL]RQWDO
IXVHG PHVDOO\ ÀDWWHQHG VRIWO\ VFOHURWL]HG SRVWHURODWHUDOO\ DQG PHVDOO\ VFOHURWL]HG
setose. Pseudoteguminal plates C-shaped, fused dorsally and ventrally, tapered antero-
ventrally with pointed edges. Fultura inferior curved, hourglass shaped. Trulleum
FRPSRXQGE\WZRSDUDOOHOEDUVVOLJKWO\VFOHURWL]HGUHFWDQJXODUDQGDWWDFKHGDQWHULRUO\
to the pseudoteguminal plate as the tegumen. Valva slightly curved, distally rounded;
EDVHHQODUJHGZLWKWKHDQWHURYHQWUDOSRUWLRQGHQVHO\VFOHURWL]HGSURGXFLQJWZRSRLQWHG
projections; internally and distally setose. Phallus membranous, tubular.
Ƃ )LJV D E  ): OHQJWK  PP ZLQJVSDQ  PP ): GRUVDOO\
ornamented as the ƃ, but ground colour paler as the stigma and its similar spots.
Ƃ JHQLWDOLD )LJV   7HUJXP 9,,, UHFWDQJXODU DQG VFOHURWL]HG VWHUQXP 9,,,
UHGXFHGUHFWDQJXODU/DPHOODDQWHYDJLQDOLVVOLJKWO\VFOHURWL]HGODWHUDOO\GRUVDOPDUJLQ
PHVDOO\ ³(VKDSHG´ VHWRVH DQG WKLFNHU VFOHURWL]DWLRQ ZLWK FHQWUDO OREH SURQRXQFHG
IROORZHGE\VPDOOGLJLWLIRUPOREHVRQHDFKVLGH)LJ&RUSXVDQGGXFWXVEXUVDHRI
WKHVDPHOHQJWK)LJ
Geographical distribution:
P. reynaudi sp. nLVFRQ¿QHGWR*XDWHPDODDWDOWLWXGHVUDQJLQJIURPWRP
)LJV
Diagnosis:
P. reynaudi sp. n. can be easily separated from other species by the presence of
a conspicuous whitish collar on the thorax, the wing ornamentation, absence of the
PHWDWLELDO DQGURFRQLDO EUXVK DQG WKH FRQ¿JXUDWLRQ RI WKH ƃ and Ƃ genitalia. These
features are so distinct, that it cannot be confused to any other described species.
Phassus violetteae sp. n.
)LJVDEDEDEDE
Holotype ƃZLWKWKHIROORZLQJODEHOVVHSDUDWHG E\IRUZDUGVODVKHVHolotypus,
Phassus violetteae &0LHONH*UHKDQGHW &RVWD5LFD &DUWDJR7DSDQWt

118 The European Entomologist, Vol. 7, No. 4
P ,9&DPSRVHFR0RQ]yQOHJ &*&0 'RQDWHGE\WKH
¿UVWDXWKRUDQGZLOOEHGHSRVLWHGLQ,1%,2±)LJVDE
ParatypesLQWRWDOƃƃƂƂ$OOCosta Rica&DUWDJRƃVDPHGDWDDVWKH
KRORW\SH &*&0   ƃ  Ƃ 3T 1DF 7DSDQWt  P ,9
:¶¶¶1¶¶¶7'HFDsQVOHJ&*&0>%&*8
ES@>%&*8ES@Ƃ3T1DF7DSDQWtP,9
: ¶¶¶ 1 ¶¶¶ 7 'HFDsQV OHJ &*&0  >%& *8
ES@
Etymology. Phassus violetteae sp. n LV QDPHG DIWHU WKH VHFRQG DXWKRUV ZLIH
&ODXGLD$9LROHWWHIRUDOOKHUVXSSRUWIRUKLVJKRVWPRWKZRUN
ƃ)LJVDED EDE):OHQJWK PPZLQJVSDQPP
$QWHQQD ¿OLIRUP ZLWK  VHJPHQWV (SLSK\VLV SUHVHQW VWRXW DQG VKRUW 3UR DQG
mesothorax dark brown, metathorax without scales anteriorly, light brown posteriorly.
FW costal margin convex at the base, apex slightly falcate and produced, outer margin
curved. FW dorsal ground colour light brown to brown; basoproximal, basodistal, and
postdiscal areas light brown, basocentral, central patch, and anterior premarginal areas
brown reaching the costal margin much darker; submarginal and marginal bands merge
into both patterns forming a mosaic. Basocentral area with distal margin straight and
fused to central patch posteriorly forming the typical V-shaped band with a thin black
internal line posteriorly. Consequently, basoproximal generally fused to the postdiscal
DUHDWKHIRUPHUZLWKEODFNGRWV7KH9VKDSHGEDQGPHUJHVLQWR DKRUL]RQWDOEURZQ
stripe between Rs4 and M1 and to a second wide branch from the stigma area ventrally
WRWKHRXWHU PDUJLQQRWUHDFKLQJ LW6WLJPDEULJKWOLJKW\HOORZJHQHUDOO\KRUL]RQWDO
Some spots similar to the stigma also present in the apex of the V-shaped band, always
conspicuous and vertical, and in the submarginal and marginal areas. HW dorsally
brown, costal and outer margins sometimes with ornamentation; veins sometimes
PDUNHGGLVWDOO\7HUJRVWHUQDOEDUVLPSOHZLWKRXWOREHDQGWDSHUHGYHQWUDOO\)LJ
7XEHUFXODWHSODWHRQWKHDEGRPLQDOVHJPHQWVIDLQWO\PDUNHG)LJVDE7HUJXP
9,,,HQODUJHGDWWKHEDVHZLWKODWHUDOFRQYH[DQGWDSHUHGSRVWHULRUO\WRQJXHVKDSHG
VWHUQXP9,,,UHFWDQJXODUZLWKWKUHHSURMHFWLRQVSRVWHULRUO\WKHFHQWUDOZLGHU)LJE
ƃJHQLWDOLD)LJ6DFFXV8VKDSHGDQWHULRUPDUJLQZLWKWRQJXHVKDSHGSURMHFWLRQ
postero-dorsal edge lobate mesally. Tegumen comma-shaped, pointed anteriorly, fused
WR WKH SVHXGRWHJXPLQDO SODWH SRVWHULRUO\ 7HUJDO OREHV IDLQWO\ VFOHURWL]HG ODWHUDOO\
VHPLOXQDWH PHVDOO\ IXVHG IRUPLQJ D WKLQ ZHOO VFOHURWL]HG DQG SURQRXQFHG EDU
Pseudoteguminal plates dorsally fused to the tergal lobes, thus the posterior side of the
phallocrypt contiguous; C-shaped, projected posteriorly, tapered antero-ventrally with
pointed edges. Fultura inferior cup-shaped; trulleum X-shaped well separated from the
pseudoteguminal plates. Valva boomerang-shaped, internal side with ventral portion
setose as the lobate distal portion. Basal portion half-length of the distal one. Phallus
membranous, tubular. Bulbus ejaculatorius long, 37 mm.

The European Entomologist, Vol. 7, No. 4
Ƃ)LJVD E):OHQJWKPP ZLQJVSDQPP ):GRUVDOO\
ornamented as the ƃEXWSDOHFRORXUHG7HUJXPDQGVWHUQXP9,,,VFOHURWL]HGZLWKRXW
GLIIHUHQWLDWLRQ)LJ
Ƃ JHQLWDOLD )LJV  /DPHOOD DQWHYDJLQDOLV VOLJKWO\ VFOHURWL]HG SRVWHULRUO\
D KRUL]RQWDO VHWRVH EDU ZLWK D VRIW LQYDJLQDWLRQ PHVDOO\ DULVLQJ DQWHULRUO\ D UREXVW
WRQJXHVKDSHGOREHSURMHFWHGSRVWHULRUO\RYHUWKHKRUL]RQWDOEDUIRUPLQJDGHSUHVVLRQ
EHWZHHQERWKVWUXFWXUHVOREHZHOOVFOHURWL]HGDQGVHWRVHGLVWDOO\DQGYHQWUDOO\6XEDQDO
SODWHVFOHURWL]HGZLWKRXWGLIIHUHQWLDWLRQ&RUSXVDQGGXFWXVEXUVDHQRWDYDLODEOHOLNHO\
due to the membrane deterioration by fungus after collecting.
Geographical distribution:
7KHQHZO\GHVFULEHGVSHFLHVLVNQRZQRQO\IURPWKHW\SHORFDOLW\)LJ
Remarks:
3YLROHWWHDH sp. nSUHVHQWVVHYHUDOIHDWXUHVWKDWVXJJHVWLWVFORVHVWDI¿QLWLHVOLHZLWKLQ
Phassus. The two ƃIRUPVDUHVRGLVWLQFWWKDWWKH\FRXOGEHPLVLGHQWL¿HGDVGLIIHUHQW
species, but no morphological differences were found other than their wingspans
DQG JURXQG FRORXU '1$ EDUFRGLQJ VKRZV QR GLYHUJHQFH 7KH SUHVHQFH RI ³GZDUI´
specimens is often seen in &LE\UDXVXDOO\LQƂƂEXWDPRQJPhassus and its relatives
WKLVUHSUHVHQWVD¿UVWUHFRUG
Diagnosis:
3 YLROHWWHDH sp. n. can be easily distinguished from its similar relatives, Phassus
triangularis Edwards, 1885 and Phassus huebneri*H\HU>@E\WKHEDVRFHQWUDO
DUHDQHYHUEHLQJH[SDQGHGGLVWDOO\VWUDLJKWPDUJLQZKLFKLQERWKWKHODWWHUWZRVSHFLHV
is produced as a convex edge, and the presence of the conspicuous bright yellow patch
at the vertex of the V-shaped band in both ƃDQG Ƃ,QDGGLWLRQ 3YLROHWWHDH sp. n.
shows a distinct shape of the saccus and the valve in the ƃ genitalia, and a distinct
ODPHOODDQWHYDJLQDOLVLQWKHƂJHQLWDOLD
Discussion
Wing pattern. The external wing patterns of both Pallas reynaudi sp. n. and Phassus
violetteae sp. nVXJJHVWDQDI¿QLW\ZLWKDWOHDVWVRPHUHFRJQL]HGPhassus species. Wing
patterns are, however, notoriously inconsistent within Hepialidae. Consistently shared
wing pattern characteristics occur within many genera, but they are not necessarily
FRLQFLGHQW ZLWK UHFRJQL]HG JHQHULF OLPLWV DQG PRVW RIWHQ FKDUDFWHUL]H D VXEVHW RI
species within a genus. In some species there is also an immense amount of variability
that includes incongruous wing patterns as illustrated in Dalaca pallens (Blanchard,
1LHOVHQ5RELQVRQ 7KHUHLVDYDULHW\RI ):SDWWHUQVLQPhassus and
DWWKLVWLPHWKHUHLV QRGH¿QHGIHDWXUHWKDWLV UHFRJQL]HGDVXQLTXHDQGVKDUHGE\DOO
species of Phassus$OOPhassus species viewed by us, other than 3 SKDOHUXVDruce,
1887, share the presence of a marginal spot located on the outer margin at the end of

120 The European Entomologist, Vol. 7, No. 4
each vein. In Hepialidae, this alignment of a marginal spot with the vein is otherwise
only known in Schausiana, an undescribed genus from Peru, 3UH\QDXGL sp. n. and an
undetermined species of Trictena 0H\ULFNIURP$XVWUDOLD-RKQ1LHOVHQSHUVRQDO
FRPPXQLFDWLRQ,QDOORWKHU+HSLDOLGDHWKHRXWHUPDUJLQDOVSRWVOLHEHWZHHQWKHYHLQV
In 3FKDPSLRQL Druce, 1887, P. KXHEQHUL3WULDQJXODULV3QVLJQDWXV:H\PHU
the spots on the outer margin are narrow, whereas in the other species, the spots broaden
to the outer margin, which is also the case for 3 UH\QDXGL sp. n. and the undescribed
Peruvian genus. In Phassus, the outer spots may be solid, or faint in the center so that
the edges are more prominent.
The basocentral area and central patch fuse posteriorly to form a V-shaped tick mark on
WKH):GDUNHUWKDQVXUURXQGLQJJURXQGFRORXUIURPWKHEDVHRI6FWRWKHDQDOYHLQDQG
then towards the apex, terminating at or near the stalk of the R vein and the stigma. In
some species, this formation is very faint, but in 3KXHEQHUL and 3WULDQJXODULV the band
is prominent and partially edged with a thin black line. There is also a narrow shaded
stripe extending between the stigma towards the outer margin between Rs4 and M1.
These features are unique within the Exoporia and therefore represent derived features
SRVVLEO\UHÀHFWLQJDFORVHSK\ORJHQHWLFUHODWLRQVKLSIRUWKHVHVSHFLHV,QWKH):RI3
QVLJQDWXV, the V-shaped band is not as contiguous, although the dash is present. With
respect to these FW characteristics, 3 YLROHWWHDH sp. n. is closer to 3 KXHEQHUL and
3WULDQJXODULV than to other Phassus species.
First abdominal segment. The structure of the tergosternal bar and its connection to
WKH SRVWHURODWHUDO PDUJLQ RI WKH ¿UVW DEGRPLQDO WHUJLWH LQFOXGHV WZR XQLTXH IHDWXUHV
that support the cibyrine clade. These are the presence of a disrupted anterior margin
and a laterally protruding lobe that varies from a rounded bump to an almost digitiform
protuberance. This structure has not yet been observed in Phassus or Schausiana,
DOWKRXJK LW LV QRW NQRZQ LI HLWKHU RU ERWK JHQHUD H[KLELW D GLIIHUHQW VSHFLDOL]HG
structure, and neither of the new species exhibit the cibyrine structure. The tergosternal
connection in 3 UH\QDXGL sp. n. is of particular interest as it shows an arrangement
QRWSUHYLRXVO\UHFRUGHG LQ+HSLDOLGDHFI*UHKDQ 7KHIRUP DQGVKDSHRIWKH
tergosternal connection in P. violetteae sp. n. conforms to that seen so far seen in other
Phassus species.
ƂJHQLWDOLD Examined species of Phassus show the tergum IX to be dorsally U-shaped
posteriorly with a slight or prominent convex edge ventrally. The subanal plate may
be tall or dorso-ventrally short. The lamella antevaginalis generally has a posterior bar
WKDWLVODWHUDOO\OREDWHDQGDZHOOVFOHURWL]HGSURQRXQFHG OREHDQWHULRUO\7KHODWHUDO
lobes vary in relative length, being shorter, longer, or equal to the anterior lobe, and are
rare in the Hepialidae, although many species have not been adequately described for
the Ƃ genitalia. There is an elongate digitiform lobe in Phassodes vitiensis (Rothschild
 IURP )LML /DQGFDUH 1HZ =HDODQG GLVVHFWLRQ / 1HZ =HDODQG $UWKURSRG
&ROOHFWLRQ$XFNODQG1HZ=HDODQGDQGDVPDOOGLJLWLIRUPOREHLQDWOHDVWRQHVSHFLHV

121The European Entomologist, Vol. 7, No. 4
of Sthenopis 3DFNDUG >@ GLVVHFWLRQ ) >-5*@ 1LHOVHQ  5RELQVRQ 
GHVFULEHG DQG LOOXVWUDWHG D ³VKRUW GLJLWLIRUP SURFHVV´ LQ Andeabatis chiliensis Ureta
 EXW WKHLU LOOXVWUDWLRQ LV QRW VXI¿FLHQWO\ GHWDLOHG WR HYDOXDWH LWV PRUSKRORJLFDO
similarity to that structure in Phassus. The genitalia of 3YLROHWWHDH sp. n. conform to
Phassus with the exception that the lateral lobes are not digitiform, but form a slight,
VHWRVHEXPSWRZDUGVWKHDQWHULRUOREH7KHVKDSHDQGUHODWLYHVL]HRIWKHDQWHULRUOREH
is most similar to that of 3WULDQJXODULV. The Ƃ genitalia of Pallas reynaudi sp. n. is
also more similar to that of Phassus than to cibyrine species that we have examined,
although the anterior lobe is absent.
ƃJHQLWDOLD. In Phassus, the valves are relatively large and the pseudoteguminal plate is
simple with a narrow apex and dorsally broad (giving a triangular shape from the ventral
SUR¿OH7KHJHQLWDOLDRIPallas reynaudi sp. n. is distinct from all Phassus species with
UHVSHFWWKHEURDGDQG WKH VKRUW DQWHULRUSURMHFWLRQRIWKH VDFFXV DQG WKHVFOHURWL]HG
posterior and concave margin of the saccus. The valves in Phassus are distally lobate,
and slightly or strongly elbowed. The valves of 3 UH\QDXGL sp. n. are also distally
lobate but this condition is widespread in the Hepialidae, and occurs in other families
of Hepialoidea, and so is not regarded as directly informative about a close relationship
with Phassus7KHVWURQJO\VFOHURWL]HGEDVHRIWKHYDOYHVDORQJZLWKWKHSDLURIVKRUW
and sharp projections are also unlike the condition in either Phassus or Schausiana. The
ƃ genitalia of 3YLROHWWHDH sp. n. are most similar to these of 3WULDQJXODULV with respect
to the presence of mesally fused tergal lobes extending vertically as a thin bar. This
feature is unique among hepialoids and therefore constitutes a shared derived character
VWDWH)XUWKHUZRUNLVUHTXLUHGWRGH¿QLWLYHO\ FRQ¿UPWKHSUHVHQFHRUDEVHQFHRIWKLV
structure in other Phassus species.
The separation of Schausiana from Phassus HPSKDVL]HVWKHKHWHURJHQHLW\ ZLWKLQWKH
ODWWHU$OWKRXJK9LHWWHGHVFULEHGSchausiana VXSHU¿FLDOO\ZLWKRXWDFRPSDULVRQ
to Phassus or other genera, it appears that the description is correct, especially regarding
some derived features found in the ƃJHQLWDOLD DW WKH EDVH RI WKH YDOYH +RZHYHU
a detailed study of other structures is needed to reinforce the generic status. In our
present taxonomic evaluation of Pallas reynaudi sp. n. and Phassus violetteae sp.
nLWZDVVXI¿FLHQWIRURXU SXUSRVHV WR DVVHVV WKHUHODWLRQVKLSZLWKUHVSHFWWR WKRVH
species currently included in Phassus and Schausiana, and particularly with Phassus
triangularis as the type species for Phassus.
Taxonomic notes. Inclusion of new species in Phassus has historically been a largely
ad hocSURFHVV UHVXOWLQJLQDQ DVVHPEODJHRIVSHFLHV ODFNLQJDQ\GH¿QLWLYHHYLGHQFH
RIVKDUHGPRQRSK\O\&RQVHTXHQWO\LWZDVGHWHUPLQHGE\ 0LHONH  *UHKDQ 
WKDWVHYHUDOVSHFLHVGLGQRW FRQIRUPRUVKRZHYLGHQFHRI FORVHDI¿QLW\ZLWKWKHW\SH
species (Phassus triangularis DQG ZHUH WKHUHIRUH SODFHG DV incertae sedis pending
further investigation. However, it has already become apparent from comparison of
illustrations and examination of the holotype of Phassus eldorado 3¿W]QHUsyn.

122 The European Entomologist, Vol. 7, No. 4
nIURP9HQH]XHOD )LJVSHFLPHQSKRWRJUDSKSXEOLVKHGIRUWKH¿UVWWLPHWKDWWKLV
taxon and Hepialus auratus*URWH)LJVsyn. n. are junior subjective
synonyms of Sthenopis pretiosus+HUULFK6FKlIIHU>@comb. nIURP%UD]LO,WLV
REYLRXVWKDWVSHFLPHQVIURP6RXWK$PHULFDDUHPLVODEHOOHGDVWKHVSHFLHVLVRQO\IRXQG
LQWKHQRUWKHDVWRIWKH1RUWK$PHULFDLQ&DQDGDDQGWKH8QLWHG6WDWHV$OWKRXJKWKH
primary type of Epilaus [sic] pretiosusKDVQRWEHHQORFDWHG0LHONH*UHKDQ
WKHLOOXVWUDWLRQLQ WKHRULJLQDOGHVFULSWLRQ )LJLV VXI¿FLHQWO\FOHDUWR HQVXUHWKDW
there is no other described taxon with which it could be confused.
Biogeographic notes. The distribution of Phassus lies between southern Mexico and
Panama (with SchausianaKDYLQJDPRUHUHVWULFWHGGLVWULEXWLRQZLWKLQWKDWUDQJH(LJKW
species are recorded from southern Mexico, including at least two with a range also
extending to Costa Rica or Panama, but the systematic validity of many of these species
has yet to be subject to detailed analysis and the sister group of the Phassus/Schausiana
group outside this region is unknown. The total range of 3KDVVXV6FKDXVLDQD lies in
a region of complex tectonic structure where eastern Mexico and the Yucatan comprise
SDUWRIWKH1RUWK$PHULFDQFUDWRQDQGZHVWHUQ0H[LFRLVPDGHXSRIWKH/DWH-XUDVVLF
Cretaceous Guerrero and associated terranes. Between these regions and extending
south to Costa Rica is another complex belt of continental margins deformed during the
0HVR]RLFDQG7HUWLDU\3RWUDHWDOLQFOXGLQJWKH&KRUWLVEORFNRIWKH+RQGXUDV
DQG*XDWHPDODWKDWVRPHJHRORJLVWVEHOLHYHPD\KDYHDFFUHWHGIURPWKHHDVWHUQ3DFL¿F
DORQJZLWKRWKHUWHUUDQHV0RUiQ=HQWHQRFRPSULVLQJ&RVWD5LFD3DQDPDDQG
WKHPDUJLQVRIQRUWKZHVWHUQ6RXWK$PHULFD,WLVSRVVLEOHWKDWWKHRULJLQRIWKHSUHVHQW
distribution range of 3KDVVXV6FKDXVLDQD evolved from a common ancestor present on
WKHWHUUDQHVEHWZHHQVRXWKHUQ0H[LFRDQGQRUWKHDVWHUQ6RXWK$PHULFD
Pallas reynaudi sp. n. is known only from higher elevations in three scattered localities
that together straddle the western end of the most prominent tectonic feature in Central
$PHULFD ± WKH 3RORFKLF DQG 0RWDJXD IUDFWXUH ]RQHV )LJ  7KHVH IDXOWV FURVV
Guatemala and represent the terrestrial end of a tectonic boundary between the North
$PHULFDQDQG&DULEEHDQSODWHV/RGROR HW DO 3LQGHOO.HQQDQ 0RVW
KLVWRULFDOUHFRQVWUXFWLRQV VXJJHVWWKLV ]RQHKDV EHHQDFWLYHVLQFHWKH0HVR]RLFDVWKH
&DULEEHDQ SODWH PRYHG HDVW UHODWLYH WR WKH 1RUWK $PHULFDQ SODWH *X]PiQ6SH]LDOH
 7KH &DULEEHDQ VHFWLRQ RI *XDWHPDOD LV PDGH XS RI WKH &KRUWLV EORFN WKDW LV
believed to be derived either from a position west of the Guerrero Terrane (Pindell
.HQQDQRUIXUWKHUZHVWLQWKHHDVWHUQ3DFL¿F0RUiQ=HQWHQR&RQWDFW
between the Chortis and Maya blocks in Guatemala has generated considerable tectonic
uplift with the highest elevation in the present landscape being the Cuchumatanes
SODWHDX\HOORZVWDULQ)LJZKHUHXSOLIWRIPLGGOH0LRFHQHSDOHRVXUIDFHUHDFKHV
nearly 4,000 m $QGHUVRQHWDO7HFWRQLFXSOLIWFDQUHVXOWLQVSHFLHVGLYHUJHQFH
as formerly lowland populations become isolated over time and altitudinal space from
RWKHU SRSXODWLRQV WKDW UHPDLQHG LQ WKH ORZODQGV +HDGV  7KH ORFDOL]DWLRQ RI
P. reynaudi sp. n. across the 3RORFKLFDQG0RWDJXDIUDFWXUH]RQHVPD\LQGLFDWHWKDWLWV

123The European Entomologist, Vol. 7, No. 4
high elevation distribution is the result of tectonic uplift of ancestral populations that
occupied lowland habitats possibly as recently as 17-10 Ma when the Cuchumatanes
SODWHDXLVHVWLPDWHGWRKDYHEHHQÀDWDQGQHDUVHDOHYHO$XWKHPD\RXHWDO7KH
precise evolutionary relationship between the high elevation range of 3UH\QDXGL sp. n.
and the history of tectonic uplift in the region will require more detailed information its
JHRJUDSKLFGLVWULEXWLRQLQWKHUHJLRQDQGLGHQWL¿FDWLRQRILWVVLVWHUJURXSUHODWLRQVKLSV
Acknowledgements
:HWKDQN-RVp0RQ]yQ*XDWHPDOD &LW\'U7KLEDXG'HFDsQV0DUVHLOOH7KLHUU\
3RULRQ-DXMDFIRUSURYLGLQJ YDOXDEOHPDWHULDOZKLFKPDGH WKLVDUWLFOHSRVVLEOH'U
:ROIUDP0H\=0+%IRUDFFHVVWKHFROOHFWLRQLQ%HUOLQ*HUPDQ\'U:ROIJDQJ1lVVLJ
60)/IRUDFFHVVWKHFROOHFWLRQ(UQVW%URFNPDQQ/LFKIRULPDJLQJWKHKRORW\SHRI
6HOGRUDGR'U-RKQ 5DZOLQV&DUQHJLH0XVHXP RI1DWXUDO+LVWRU\3LWWVEXUJKIRU
scanning the S. pretiosusLPDJH'U-DVRQ'RPEURVNLH&RUQHOO8QLYHUVLW\,WKDFDIRU
sending the photograph of a living Sthenopis; Ryan St. Laurent (Cornell University,
,WKDFDIRULPDJLQJWKHƃ of Sthenopis; Dr. /RGROR,QVWLWXWR1D]LRQDOHGL2FHDQRJUD¿D
HGL *HR¿VLFD6SHULPHQWDOH7ULHVWHIRUDOORZLQJXVWR UHSURGXFHLPDJHV DQG¿QDOO\
'U,DQ.LWFKLQJ1+08.IRUVXJJHVWLRQVDQGUHYLHZLQJWKHPDQXVFULSWFunding for
'1$EDUFRGLQJZDVSURYLGHGE\WKHJRYHUQPHQWRI&DQDGDWKURXJK*HQRPH&DQDGD
DQG WKH 2QWDULR *HQRPLFV ,QVWLWXWH LQ VXSSRUW RI WKH ,QWHUQDWLRQDO %DUFRGH RI /LIH
project, and by NSERC.

124 The European Entomologist, Vol. 7, No. 4
Fig. 1a Fig. 1b
Figs. 1–3: Pallas reynaudi sp. n37ƃGRUVDOYLHZD>&*&0@YHQWUDOYLHZ
E+7ƂGRUVDOYLHZ D >&*&0 @ YHQWUDO YLHZE37ƂGRUVDO YLHZ
>&*&0@
Fig.2a Fig. 2b
Fig. 3
6FDOHEDUVFP6FDOHEDUVFP
6FDOHEDUVFP6FDOHEDUVFP 6FDOHEDUVFP

125The European Entomologist, Vol. 7, No. 4
Figs. 4–7: Phassus violetteae sp. n+7ƃGRUVDOYLHZD>&*&0@YHQWUDO
YLHZE37ƃGRUVDOYLHZD>&*&0@YHQWUDOYLHZE37ƂGRUVDOYLHZ
D>&*&0@YHQWUDOYLHZE37ƂGRUVDOYLHZ&*&0
Fig. 4a Fig. 4b
Fig. 5a Fig. 5b
Fig. 6a Fig. 6b
Fig. 7
6FDOHEDUVFP
6FDOHEDUVFP
6FDOHEDUVFP
6FDOHEDUVFP
6FDOHEDUVFP
6FDOHEDUVFP
6FDOHEDUVFP

126 The European Entomologist, Vol. 7, No. 4
Figs. 8–12: Sthenopis pretiosus+7ƃRIPhassus eldorado3¿W]QHUGRUVDOYLHZƃ
GRUVDOYLHZ86$1<7RPSNLQV&R9,,-*)UDQFOHPRQWOHJ>&RUQHOO
8QLYHUVLW\,WKDFD86$@3KRWR E\5\DQ6W /DXUHQWƂGRUVDOYLHZ86$1<
&DPEULGJH:DVKLQJWRQ&R>&*&0%&-;ES@
ƃOLYLQJVSHFLPHQ&DQDGD2QWDULR$OJRQTXLQ3DUN%UXWRQWZSRQORJJLQJURDG
P1RI:DWVRQ/DNH9,,3KRWRJUDSKE\'U-DVRQ'RPEURVNLHƃGRUVDO
YLHZVFDQRIRULJLQDOGUDZLQJRI+HUULFK6FKlIIHU
Fig. 8
Fig. 9
Fig. 10 Fig. 12
Fig. 11
6FDOHEDUVFP 6FDOHEDUVFP6FDOHEDUVFP

127The European Entomologist, Vol. 7, No. 4
Fig. 14: Pallas reynaudi sp. n37ƃDEGRPLQDOVHJPHQWV,9D9,9,,,E
Fig. 13: Pallas reynaudi, sp. n37ƃWHUJRVWHUQDOEDU
6FDOHEDUVPP
6FDOHEDUVPP
6FDOHEDUVPP

128 The European Entomologist, Vol. 7, No. 4
Fig. 15: Pallas reynaudi sp. n37ƃJHQLWDOLDYHQWUDO YLHZDYHQWUDO YLHZZLWK
SVHXGRWHJXPLQDOSODWHVSXOOHGRXWE
Fig. 16: Pallas reynaudi sp. n+7ƂJHQLWDOLDSRVWHULRUYLHZ
6FDOHEDUVPP
6FDOHEDUVPP
6FDOHEDUVPP

The European Entomologist, Vol. 7, No. 4
Fig. 17: Pallas reynaudi, sp. n+7ƂJHQLWDOLDGXFWXVDQGFRUSXVEXUVDH
Fig. 18: Phassus violetteae, sp. n37ƃWHUJRVWHUQDOEDU
6FDOHEDUVPP
6FDOHEDUVPP

130 The European Entomologist, Vol. 7, No. 4
Fig. 19: Phassus violetteae, sp. n37ƃ
DEGRPLQDO VHJPHQWV ,,9 D 99,,,
E
Fig. 20: Phassus violetteae, sp. n37ƂDEGRPLQDOVHJPHQWV9,,9,,,
6FDOHEDUVPP6FDOHEDUVPP
6FDOHEDUVPP

131The European Entomologist, Vol. 7, No. 4
Fig. 20: Phassus violetteae, sp. n37ƂDEGRPLQDOVHJPHQWV9,,9,,,
Fig.23:
Pallas reynaudi,
sp. n. and Phassus
violetteae sp. n.
distribution within
Costa Rica and
Guatemala.
6FDOHEDUVPP
6FDOHEDUVPP

132 The European Entomologist, Vol. 7, No. 4
References
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ZHVWHUQ$OWRV&XFKXPDWDQHVQRUWKZHVWHUQ*XDWHPDOD *HRORJLFDO6RFLHW\ RI$PHULFD%XOOHWLQ 84: 805-
826.
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fault systems: Pull-XSDQG]LSSHUPRGHOVTectonics 30: 1-23.
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=RRWD[D1944: 34–52.
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133The European Entomologist, Vol. 7, No. 4
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+HSLDOLGDHDVIRRGRILQGLJHQRXVSHRSOHLQ0H[LFRJournal of Insects as Food.
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+HDGV0-Molecular Panbiogeography of the Tropics. University of California Press, Berkeley,
565 pp.
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134 The European Entomologist, Vol. 7, No. 4
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Costa Rica, 161 pp.
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