Zoosystematics and Evolution

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Zoosystematics
and Evolution
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94 2 2018
rMf/kz*
r MUSEUM FUR
9 -^NATURKUNDE
ISSN 1435-1935 Zoosyst. Evol. 94 (2) 2018, 211-566 BERLIN
Zoosystematics and Evolution
A Bulletin of Zoology since 1898
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Zoosystematics
and Evolution
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94 2 2018
MUSEUM FUR
NATURKUNDE
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Zoosystematics and Evolution
A Bulletin of Zoology since 1898
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Zoosystematics and Evolution
Mollusca - History of Science - Evolution & Biogeography
Matthias Glaubrecht - Center of Natural History (CeNak)
2017. Volume 94. 2 Issues
Hamburg
ISSN: 1435-1935 (print), 1860-0743 (online)
Arachnida - Taxonomy - Biodiversity & Conservation Abbreviated keys title: Zoosyst. Evol.
Danilo Harms - Center of Natural History (CeNak) Hamburg
Arthropoda - Taxonomy - Molecular biology -
In Focus
Biodiversity & Conservation
Martin Husemann-Center of Natural History (CeNak) Hamburg The cover picture shows a male of Hypsolebiassplendissimus
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Reptilia - Amphibia - Taxonomy - General Ecology -
Biodiversity & Conservation
See paper of Costa WJEM et al. Diversity and conservation
Johannes Penner - Museum fur Naturkunde Berlin
of seasonal killifishesof theHypsolebiasfulminantiscomplex
from a Caatinga semiarid upland plateau, Sao Francisco
Nematomorpha - Taxonomy - Marine - Systematics
River basin, northeastern Brazil (Cyprinodontiformes,
Andreas Schmidt-Rhaesa - Center of Natural History (CeNak)
Aplocheilidae)
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Zoosystematics and Evolution
A Bulletin of Zoology since 1898
Content of volume 94 (2) 2018
Imada Y, Kato M
Descriptions of new species of Issikiomartyria (Lepidoptera, Micropterigidae) and
a new genus Melinopteryx gen. n. with two new species from Japan 211
Lima POV, Simone LRL
Complementary anatomy of Actinocyclus verrucosus (Nudibranchia, Doridoidea,
Actinocyclidae) from Indo-Pacific 237
Scherz MD, Hawlitschek 0, Razafindraibe JH, Megson S, Ratsoavina FM,
Rakotoarison A, Bletz MC, Glaw F, Vences M
A distinctive new frog species (Anura, Mantellidae) supports the biogeographic
linkage of two montane rainforest massifs in northern Madagascar 247
Volcan MV, Severo-Neto F, Lanes LEK
Unrecognized biodiversity in a world’s hotspot: three new species of Melanorivulus
(Cyprinodontiformes: Rivulidae) from tributaries of the right bank of the
Rio Parana basin, Brazilian Cerrado 263
Oliver PG, Morgenroth H
Additional type and other notable specimens of Mollusca from the Montagu Collection
in the Royal Albert Memorial Museum & Art Gallery, Exeter 281
Liu T, Wang E, Wang S
Taxonomic study of the leafmining genus Liocrobyla Meyrick, 1916 from China
(Lepidoptera, Gracillariidae, Ornixolinae) with a description of one new species 305
Koch A
Discovery of a rare hybrid specimen known as Maria’s bird of paradise at the Staatliches
Naturhistorisches Museum in Braunschweig 315
Mattos JLO, Costa WJEM
Three new species of the ‘Geophagus’ brasiliensis species group from the northeast Brazil
(Cichlidae, Geophagini) 325
Sata N
Two new skink-endoparasitic species of Meteterakis (Nematoda, Heterakidae, Meteterakinae)
from East Asian islands 339
Correa C, Zepeda P, Lagos N, Salinas H, Palma RE, Vasquez D
New populations of two threatened species of Alsodes (Anura, Alsodidae) reveal the scarce
biogeographic knowledge of the genus in the Andes of central Chile 349
Costa WJEM, Amorim PF
A new miniature cryptic species of the seasonal killifish genus Spectrolebias from
the Tocantins River basin, central Brazil (Cyprinodontiformes, Aplocheilidae) 359
Abstract & Indexing Information
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Zoological Record™ (Thompson ISI)
Zoosystematics and Evolution
A Bulletin of Zoology since 1898
Content of volume 94 (2) 2018
Song Z-S, Malenovsky I, Chen J-Q, Deckert J, Liang A-P
Taxonomic review of the planthopper genus Orthopagus (Hemiptera, Fulgoromorpha,
Dictyopharidae), with descriptions of two new species 369
de Brito PS, Guimaraes EC, Katz AM, Piorski NM, Ottoni FP
Taxonomic status of Aphyocharax avary Fowler, 1913, Aphyocharax pusillus Gunther, 1868
and Chirodon alburnus Gunther, 1869 (Characiformes, Characidae) 393
Shah AA, Ahmad M, Taqweem-UI-Haq
Deciphering conserved identical sequences of mature miRNAs among
six members of great apes 401
Protzel D, Lambert SM, Andrianasolo GT, Hutter CR, Cobb KA, Scherz MD, Glaw F
The smallest ‘true chameleon’ from Madagascar: a new, distinctly colored species of
the Calumma boettgeri complex (Squamata, Chamaeleomdae) 409
Veeravechsukij N, Namchote S, Neiber MT, Glaubrecht M, Krailas D
Exploring the evolutionary potential of parasites: Larval stages of pathogen
digenic trematodes in their thiarid snail host Tarebia granifera in Thailand 425
Veeravechsukij N, Krailas D, Namchote S, Wiggering B, Neiber MT, Glaubrecht M
Molecular phylogeography and reproductive biology of the freshwater snail Tarebia granifera
in Thailand and Timor (Cerithioidea, Thiaridae): morphological disparity
versus genetic diversity 461
Costa WJEM, Amorim PF, Mattos JLO
Diversity and conservation of seasonal ki 11 if ishes of the Hypsolebias fulminantis complex
from a Caatinga semiarid upland plateau, Sao Francisco River basin,
northeastern Brazil (Cyprinodontiformes, Aplocheilidae) 495
Cunha CM, Salvador RB
Type specimens of fossil “Architectibranchia" and Cephalaspidea (Mollusca, Heterobranchia)
in the Academy of Natural Sciences of Philadelphia 505
Coleman ChO
Shadings in digital taxonomic drawings 529
Pimenta AD, Santos FN, Cunha CM
Redescription and reassignment of Ondina semicingulata to the Pyramidellidae, with review
of the occurrence of genus Evalea in the Western Atlantic (Gastropoda) 535
Wildish DJ
Neotenorchestia Wildish, 2014 is a junior synonym of Orchestia Leach, 1814 545
Serra WS, Loureiro M
Austrolebias queguay (Cyprinodontiformes, Rivulidae), a new species of annual killifish
endemic to the lower Uruguay river basin 547
Katz AM, Barbosa MA, de Oliveira Mattos JL, da Costa WJEM
Multigene analysis of the catfish genus Trichomycterus and description of a new
South American trichomycterine genus (Siluriformes, Trichomycteridae) 557
Zoosyst. Evol. 94(2) 2018, 211-235 | DOI 10.3897/zse.94.13748
Lr/jZ*
VEHSOTT
r MUSEUM
• UNHATTUIIDRKIfUl NDE
BERLIN
Descriptions of new species of Issikiomartyria (Lepidoptera,
Micropterigidae) and a new genus Melinopteryx gen. n.
with two new species from Japan
Yume Imada1’2, Makoto Kato
3
1 Graduate School of Science and Engineering, Ehime University, 2-5 Bunkyo-cho, Matsuyama, Ehime 790-8577, Japan
2 Department of Paleobiology!, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20013, USA
3 Graduate School of Human and Environmental Studies, Kyoto University’, Yoshida-Nihonmatsu-cho, Saky’o, Kyoto 606-8501, Japan
http://zoobank. org/7D111266-5F9F-4DlE-BEAC- 7E405F3 79DB9
Corresponding author: Yume Imada ([email protected])
Abstract
Received 19 May 2017
Micropterigidae is considered to be the sister group of all other extant Lepidoptera. In
Accepted 20 February 2018
Japan, 17 species of five genera have been recorded including three endemic genera,
Published 15 March 2018
Issikiomartyria Hashimoto, 2006, Kurokopteryx Hashimoto, 2006 and Neomicropteiyx
Issiki, 1931, all of which are associated with the liverwort genus Conocephalum Hill. We
Academic editor:
discovered four new species of Issikiomartyria from snowy regions in Northeastern Japan,
Michael Ohl
and two new species of a new genus Melinoptetyx gen. n. from the subalpine zone of the
Akaishi Mountain Range. All these new taxa, 1. hyperborea sp. n., 1. leptobelos sp. n.,
Key Words I. catapasta sp. n., I. trochos sp. n., M. coruscans sp. n. and M. bilobata sp. n. are
also associated with Conocephalum liverworts. Furthermore, females of I. akemiae
Sea of Japan Hashimoto, 2006 and 1. plicata Hashimoto, 2006 are described here for the first time. Our
nonglossata extensive surveys revealed that the fine-scale endemism of Issikiomartyria restricted to
Zeugloptera the fragmented area facing the Japan Sea. Keys to Issikiomartyria species based on the
bryophyte-feeding adult morphology are provided.
Introduction
Kinzelbach 2008), and more than 100 species have
been found which are still undescribed (van Nieukerken
Micropterigidae represents one of the branches in the first et al. 2011). The geographic ranges of the micropterigids
splitting event among extant Lepidoptera (Kristensen are patchy, with a concentration of endemic genera in the
1999, Kristensen et al. 2007). The fossil record of Australian ecozone and the Japanese Archipelago, where
Micropterigidae extends to the Early Cretaceous (Whalley four and three genera are known respectively (Gibbs 2010,
1978) and likely originated during or before the Late 2014, Hashimoto 2006). The global diversity and high
Triassic (Zhang et al. 2013, van Eldijk et al. 2018). The endemism of the micropterigids provide us with a wealth
Micropterigidae is a non-glossatan moth family, which of opportunities for understanding of the biogeographic
along with Agathiphagidae and Heterobathmiidae are patterns and processes suggested by primitive moths
devoid of a coilable proboscis and instead retain functional (Gibbs 1983, 2006, Gibbs and Lees 2014) whose
mandibles which in micropterigids only are functional in evolution and ecological associations largely predates the
the adult (Kristensen 1984, 1999, Kristensen et al. 2015, diversification of angiosperms (Imada et al. 2011).
Regier et al. 2015). The Micropterigids are species- Micropterigidae in Japan comprise 17 described spe¬
rich, comprising about 160 described species in 21 cies in five genera: namely, Micropterix Hiibner, 1825,
genera from all biogeographic regions (Gibbs 2010, van Paramartyria Issiki, 1931, Neomicropteiyx Issiki, 1931,
Nieukerken et al. 2011), including four fossil genera Kurokopteryx Hashimoto, 2006, and Issikiomartyria
(Whalley 1978, Kozlov 1988, Skalski 1995, Engel and Hashimoto, 2006. Micropterix is a genus distributed
Copyright Yume Imada, Makoto Kato. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which
permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
212 Imada, Y. & Kato, M.: Descriptions of new species of Micropterigidae
over the Palearctic ecozone as far as the Himalayan foot¬ series. Genitalia were then stained with 5 % chlora-
hills (Lees et al. 2010); in Japan, M. aureatella (Scopoli, zol black E for 10 min, and dehydrated in a series of
1763) patchily inhabits Hokkaido and subalpine zones 70-100 % ethanol. After washing, specimens were mount¬
of Honshu (Issiki 1953; Hashimoto 2006, Imada et al. ed and stored in 70 % glycerol. For some specimens, to
2011). Among East Asian Paramartyria, two species are examine the wing veins, wings were removed and scales
known only from Japan (Issiki 1931): P. immaculatella were removed by brushing in 70 % EtOH. Observation
Issiki, 1931 and P. semifasciella Issiki, 1931. Three gen¬ and measurements were made under an Olympus BX53F
era, Neomicropteryx, Kurokopteryx, and Issikiomartyria, microscope at 10^10x with the aid of a micrometer scale.
are endemic to Japan and the species ranges are often re¬ All the specimens examined in this study are deposited
stricted in small geographic areas in a strongly allopatric in the following collections: National Museum of Nature
fashion (Hashimoto 2006, Imada et al. 2011). and Science (NMNS), Graduate School of Human and En¬
All species of Japanese endemic genera, comprising vironmental Studies, Kyoto University (KUHE).
at least 14 species, feed exclusively on Conocephalum Terminology follows Gibbs and Kristensen (2011) and
Hill, liverworts and also seem to occupy similar ecolog¬ Davis and Landry (2012). Author’s names are abbreviat¬
ical niches (Hashimoto 2006, Imada et al. 2011). Re¬ ed: YI and MK stand for Yume Imada and Makoto Kato,
markably, the liverwort-feeding micropterigids in Japan respectively.
have experienced diversification without changing host-
plants, during or after the Miocene (Imada et al. 2011),
Taxonomy
which represent one of the largest radiations known to
have taken place on a single host-plant genus. These
Melinopteryx gen. n.
Conocephalum-feeders are widely distributed across
the major islands of Japan (i.e. Honshu, Shikoku, Ky¬
http://zoobank.org/54C3DAD7-B3D6-4EA7-8706-6DAFBCD57E63
ushu) yet the species do not co-occur with one another
Figs la, b, 2-7
(Imada et al. 2011).
Our thorough field surveys for molecular phylogenetic Type species. Melinopteryx coruscans sp. n. by present
analysis have revealed that the micropterigid fauna in the designation.
Northeastern Japan is unexpectedly diverse. We reveal a
new genus that inhabits an elevational zone of ca. 1500- Diagnosis. Aedeagus with three pairs of dorsal fins, a pair
1800 m in the Akaishi Mountain Range, which is sister of lateral triangular fins, and a ventral longitudinal fin.
to Issikiomartyria in the molecular phylogenetic analysis Genital chamber with a large genital sclerite with four
of Imada et al. (2011). In addition, we newly discovered paddle-shaped accessory sclerites at posterior end.
that four new species of Issikiomartyria were distributed
across the area ranging from the northern end of Honshu Description. The generic description is based on M. cor¬
to the northern part of Yamagata Prefecture, which has not uscans sp. n. and M. bilobata sp. n.
sufficiently been investigated. Issikiomartyria is a genus Head capsule densely covered by microtrichia; genal
consisting of five species, and each species tends to be area glossy and naked; most of clypeus, frons, and ver¬
found from geographically fragmented areas facing the tex covered with brownish yellow piliform scales. Ocelli
Japan Sea (Issiki 1953; Hashimoto 2006). Furthermore, present. Antenna moniliform, approximately as long as
we describe females of I akemiae Hashimoto, 2006 and forewing in male, longer than in female; densely covered
I plicata Hashimoto, 2006, for the first time. with fuscous piliform scales on scape and pedicel; scape
In this study, males and females of four Issikiomartyria the largest segment, three times longer than most basal
species new to science and females of two known species flagellum; pedicel small, as long as most basal flagellum.
are described. Also, a new genus is established based on SOI (Kristensen and Nielsen 1979) about 0.4. MIOI (Hi-
two species new to science. An updated account of the dis¬ rowatari 1997) about 0.5. Interocellar sulcus complete.
tribution of micropterigids in the northeastern Japan based Postinterocellar sulcus distinct. Epicranial sulcus almost
on the detailed additional sampling records is provided. absent. Temporal sulcus as a darker line. Occipital sulcus
interrupted at ventro- and dorso-lateral corner. Occipus
fan-shaped. Labrum approximately pentagonal, length
Methods
more or less twice that of clypeus. Mandible elongate
rectangular, distal edge truncated. Proximal prelabium
Adults and larvae of micropterigid moths were collected sclerite weakly sclerotized. Labial palp 2-segmented.
from temperate forests in Japan, and larvae were reared Maxillary palp 5-segmented.
in plastic cases with their host-plants. A total of 226 adult Foretibial epiphysis absent. Antero-lateral processes
pinned specimens were used for this study. For genital of pronotum present, weakly sclerotized.
dissections, the whole abdomen was removed and mac¬ Wing venation as shown in Fig. 3A, B. Fore- and hind-
erated for 30 min in 10 % KOH. Residual scales and tis¬ wings obtuse at apex, forewing with brown to purple lus¬
sues were then washed in distilled water to remove KOH, ter, cilia shining grayish brown. Forewing with Sc forked,
immersed in 50 % ethanol and dehydrated in an ethanol R1 unforked; R3 stalked with R4+5. Hind wing with a
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Zoosyst. Evol. 94(2) 2018, 211-235 213
.
Figure 1 Habitus of adult males of Melinopteryx and Issikiomartyria. A: Melinopteryx coruscans sp. n. [holotype; MC 0204];
B: M. bilobata sp. n. [holotype: MC 0221]; C: Issikiomartyria hyperborea sp. n. [holotype: MC0252]; D: I. leptobelos sp. n.
[MC 0250]; E: I. catapasta sp. n. [MC 0241]; F: I. trochos sp. n. [MC 0231]
main stem of R absent; most anterior vein of hindwing ly membranous at proximo-dorsal surface, with a proxi¬
forked near terminal end (Scl and Sc2+Rl). Abdomen mo-ventral ridge; anterior portion fused with median plate;
grayish brown, covered with piliform and lamellar scales, median plate large, roughly fan-shaped. Phallobase strong¬
scattered with dark orange piliform scales on venter and ly curved, without ventral longitudinal ridge. Aedeagus
genital segments in male. Sternum V gland present; ori¬ stout at caudal end, with three pairs of fins dorso-medially;
fice of gland a narrow slit. a pair of lateral triangular fins extending horizontally; a
Male abdomen and genitalia. Sternum VIII membra¬ pair of ventral fin extending vertically; dorsal apex of ae¬
nous. Segment IX a complete ring, well sclerotized, with deagus acute and ventral one slightly forked, longer than
a posterior expansion dorsally. Valva triangular, broad¬ dorsal one; gonopore opening horizontally; vesica with
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214 Imada, Y. & Kato, M.: Descriptions of new species of Micropterigidae
Figure 2. Adult heads of Mehnopteryx and Issikiomartyria. A: Melinoptetyx bilobata sp. n., frontal view; B: ditto, posterior view;
C: Issikiomartyria hyperborea sp. n., frontal view. Abbreviations: an = antenna; cl = clypeus; dp = distal prelabium; ess = episto-
mal sulcus; gl = galea; la = labium; Ip = labial palp; md = mandible; mp = mandibular palp; oc = ocellus; os = occipital sulcus;
pp = proximal prelabium. Scale bars: 0.5 mm.
serrate minute projections. Tergum X broader than long, large as the intergeneric COI divergence among other Japa¬
with a pair of long ventral plates (venter X plates) extend¬ nese micropterigid genera (for example, 7.8 ± 0.1% between
ing antero-ventrally at base of terminal processes. Neomicropteryx and Kurokopteryx), indicated by Imada et
Female abdomen and genitalia. Segment IX forming al. (2011). Melinopteryx has a 2-segmented labial palp, while
a complete ring, strongly sclerotized, with a dorso-lateral 1-segmented in Issikiomartyria, except for I bisegmentata
concavity, without lateral protrusion. Segment X com¬ Hashimoto, 2006 which is 2-segmented. Melinopteryx is
posed by lateral sclerites and one or two dorsal sclerotized characteristic in its aedeagus with three pairs of dorsal fins,
plates; lateral sclerites simple, broader than long, having a pair of lateral triangular fins, a ventral longitudinal fin or
digitate projections with an apical seta at terminal inner protrusion extending vertically in the male. In female, a large
margin. Corpus bursae large, globular, membranous, with sclerite with more than four accessory sclerites in genital
signa composed of three or four sclerites near caudal end. chamber is also unique in Melinopteryx. Fore- and hindwing
Genital chamber with a large sclerite (genital sclerite) and without a radial cell and large bulbous corpus bursae in fe¬
a few tiny sclerites; genital sclerite deeply furcated poste¬ male are shared with Issikiomartyria.
riorly into four paddle-shaped accessory sclerites.
Etymology. The genus name is a compound noun de¬
Remarks. The genetic distance between Melinopteryx cor- rived from the Greek words transliterated into Latin,
uscans sp. n. (labelled as ‘Issikiomartyria’ sp. in Imada et al. “melinos” (honey-color) and “pteryx” (wing), referring
2011) and its sister clade Issikiomartyria spp. (6.3 ± 0.9%, to the adult wing color of the species of this genus. The
based on the Kimura 2-parameter model) was almost as gender is feminine.
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