Morphology-Based Phylogeny and Biogeography of Wockia (Lepidoptera: Urodidae) with Description of a New Species from Japan and South Korea Author(s): Jae-Cheon Sohn Source: Zoological Science, 31(4):258-265. 2014. Published By: Zoological Society of Japan DOI: http://dx.doi.org/10.2108/zs130199 URL: http://www.bioone.org/doi/full/10.2108/zs130199
BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.
BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research.
¤ 2014 Zoological Society of Japan
ZOOLOGICAL SCIENCE 31: 258–265 (2014)
Morphology-Based Phylogeny and Biogeography of Wockia (Lepidoptera: Urodidae) with Description of a New Species from Japan and South Korea Jae-Cheon Sohn1,2* 1
Department of Entomology, University of Maryland, 4112 Plant Sciences Building, College Park, MD 20742, USA 2 Department of Entomology, Smithsonian Institution, National Museum of Natural History, 10th & Constitution NW, Washington, DC 20560, USA
Wockia, one of the six genera within the lepidopteran family Urodidae, currently includes nine species distributed in Holarctic, Oriental, and Neotropical regions. A new species of the genus, W. magna sp. nov., is described from Japan and South Korea. This is the first record of Urodidae from Japan. A cladistic analysis was constructed based on 27 morphological characters from five ingroup species and two outgroup taxa. A single most parsimonious tree was found (length = 38, CI = 71, RI = 70). The resulting tree failed to recover the monophyly of Wockia chewbacca and a clade Wockia sensu stricto, including four congeners from North America, temperate East Asia and Southeast Asia. Wockia sensu stricto was resolved as monophyletic and was divided into two subgroups, one including W. asperipunctella and W. magna, and the other including W. koreana and W. variata. The sister-group relationship of W. asperipunctella and W. magna was moderately supported. Synapomorphies of Wockia sensu stricto are provided from the resulting phylogeny. Systematic definitions of Wockia and other allied genera are revised. Optimal ancestral area reconstruction implemented in DIVA resulted in different hypotheses for Wockia, depending on how to constrain the maximum number of areas. The maximum area number set as two yielded a more likely scenario suggesting that Wockia sensu stricto originated in temperate East Asia and then dispersed into the Oriental region, North America, and Europe. A working hypothesis and other equally possible alternative explanations for the biogeography of Wockia are provided. Key words: biogeography, cladistic analysis, taxonomy, Urodoidea, Wockia INTRODUCTION The False Burnet moth family, Urodidae currently include six genera: Urodus Herrich-Schäffer, 1854, Wockia Heinemann, 1870, Spildarcha Meyrick, 1913, Anchimacheta Walsingham, 1914, Incawockia Heppner, 2010a, and Anomalomeuta Sohn, 2013a. Many of the included species have long been misidentified as Yponomeutidae or Plutellidae. Kyrki (1984) first suggested that they are distinguishable from Yponomeutoidea in having the tortricoid type of thoracic-abdominal articulation and hence representing close ties with Apoditrysia. Later, Kyrki (1988) assigned them to their own family Urodidae. This proposal has been corroborated by two recent molecular studies (Sohn et al., 2013; Regier et al., 2013), whereas their systematic position within Apoditrysia remains unclear. Kyrki (1988) noticed that Urodidae share some characteristics with Schreckensteiniidae, such as the elongate prolegs of the larvae and the loosely-meshed cocoon, although those similarities are not * Corresponding author. Tel. : +1-301-717-5462; Fax : +1-301-314-9290; E-mail : [email protected]; [email protected] Supplemental material for this article is available online. doi:10.2108/zs130199
exclusively unique to those two families. Among molecular phylogenetic studies, Mutanen et al. (2010) recovered Schreckensteiniidae as sister to Urodidae. On the other hand, Cho et al. (2011) found that the Urodidae are a sister taxon to Alucitidae, Choreutidae, Pterophoridae, or Tortricidae, depending on the data set. In Regier et al. (2013), Urodidae were identified as sister to the rest of apoditrysian families. None of the abovementioned studies, however, resolved the relationships with significant support. Wockia is a relatively small genus comprising gray or fuscous micro-moths with an oblique antemedian band of raised scales (Kyrki, 1986). Members in the genus are very similar to those in Blastobasidae or Phycitinae (Pyralidae) in superficial appearance (Landry, 1998). Kyrki (1988) provided generic features, based on W. asperipunctella (Bruand, 1851), the only confirmed species of the genus in his study. Since then, the species diversity of Wockia has increased with discoveries of new species from several faunal regions. Wockia currently include nine described species: W. asperipunctella from Europe and North America (Kyrki, 1988; Heppner, 1997; Landry, 1998); W. balikpapanella Kyrki, 1986 from Borneo (Kyrki, 1986); W. koreana Sohn, 2008 from South Korea (Sohn and Adamski, 2008); W. variata Sohn and Park, 2009 from North Vietnam (Sohn and Park, 2009; Heppner, 2010b); two Mexican species, W.
Phylogeny and biogeography of Wockia chewbacca Adamski, 2009 and W. mexicana Adamski, 2009 (Adamski et al., 2009); two Jamaican species, W. diabloca Sohn, 2013 and W. tetroidon Sohn, 2013 (Sohn, 2013b); and W. pelotas Heppner, 2008 from Brazil (Heppner, 2008). This increase in species diversity has prompted a reconsideration of Kyrki’s (1988) definition of the genus. Global distribution of Wockia includes at least four zoogeographic regions, in contrast to the other five urodid genera, Anchimacheta, Anomalomeuta, Incawockia, Spiladarcha, and Urodus, which are entirely or predominantly Neotropical. Such widely scattered occurrence of Wockia may not be easy to understand without phylogenetic approaches. Thus far, however, no phylogenetic study has attempted to resolve the interspecific relationships or the biogeography of Wockia. The objectives of this study are to: 1) construct a phylogeny of Wockia, based on their morphologies, 2) understand their biogeography, and 3) describe a new species from Japan and South Korea that is the second congener from the temperate East Asia. MATERIALS AND METHODS Preparation of data set A total of seven taxa, including two outgroup taxa, Spiladarcha adamskii Sohn, 2012 and Anchimacheta iodes Walsingham, 1914, were sampled. Pinned specimens were obtained from the following collections (also see Supplementary Table S1 online): Natural History Museum, London, United Kingdom (BMNH); Chungbuk
259
National University, Cheongju, South Korea (CNU); Florida Museum of Natural History, Gainesville, Florida, USA (FLMNH); Korea National Arboretum, Pocheon, Kyonggi, South Korea (KNA); United States Museum of Natural History, Washington DC, USA (USNM). External morphologies were examined with a Leica MZ APO stereoscope and photographed using a Nikon D40 digital camera. Genitalia slides were prepared following Clarke (1941), with the modifications that Chlorazol Black was used for staining and Euparal resin for permanent slide mounting. The wings were slidemounted according to Hodges (2005). Slide-mounted specimens were examined with a Leica LETTZ-DMRX microscope. Terminology follows Klots (1970) for genitalia; Wootton (1979) for wing venation; and Heppner (1998) for other body parts. Ovipositor is defined as the segments and intersegmental membranes after the abdominal segment VIII. Verbatim label data are given only for holotype. Character coding The data matrix was constructed using WinClada ver. 1.0 (Nixon, 2002). A total of 27 adult morphological characters were coded. Parsimony-uninformative characters were excluded to avoid their influence on branch support calculation (Carpenter, 1992). The morphological characters (Table 1) were retrieved from the heads (n = 3), wings (n = 3), male genitalia (n = 12), and female genitalia (n = 9). All encoded characters were binary, non-additive, and equally weighted. Missing data were encoded as “?” and inapplicable character states as “–”. The data matrix is available at the Supplementary Table S2 online.
Table1.ޓCharacters coded for cladistic analysis. Characters Head 0: Scales on the vertex in females 1: Spiniform scales on terminal 1/3 of 3rd labial palpal segment 2: Scales on dorsal and lateral areas of occiput Wings 3: Erected scales of two subbasal dots on forewing 4: Erected scales of small spot near forewing tornus 5: Transparent hyaline area on hindwing Male genitalia 6: An articulation between tegumen and uncus 7: Shape of uncus 8: Long setae on the top of uncus 9: Posterolateral bulges or projections on uncus 10: Gnathal arms 11: Corona with strong spines 12: Stick-like terminal elongation of valva 13: Terminal saccular angulation 14: Nonsetose lobe beyond saccular angulation 15: Costal process 16: Shape of juxta 17: Spiniform cornutus or spinulate cornuti Female genitalia 18: Position of ostium bursae 19: Winkles in ductus bursae near to ostium bursae 20: Antrum 21: Sclerotization of antrum 22: Length of ductus bursae with relation to corpus bursae 23: Burge at the junction of ductus seminalis 24: Entry of ductus seminalis 25: Corpus bursae 26: Two identical signa
Character States 0) piliform, 1) broad 0) absent, 1) present 0) dorsal scales longer than lateral scales, 1) dorsal and lateral scales similar in size 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) triangular, 1) rectangular 0) absent, 1) present 0) absent, 1) present 0) not continuous or reduced, 1) continuous 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) U-shaped, 1) elongate 0) absent, 1) present 0) on or after anterior margin of sternite VII, 1) before anterior margin of sternite VII 0) absent, 1) present 0) absent, 1) present 0) complete, 1) partial 0) longer or almost same length, 1) shorter 0) absent, 1) present 0) on the way of ductus bursae, 1) near to corpus bursae 0) globular or ovoid, 1) elongate-elliptical 0) absent, 1) present
J.-C. Sohn
260
Phylogenetic analysis Parsimony analyses were performed using WinClada ver. 1.0 (Nixon, 2002) which implements NONA ver. 2.0 (Goloboff, 1999) to search for the most parsimonious tree. The following commands were used: hold 1000, hold/100, mult*1000, max*, and mswap+. The resulting trees were examined with WinClada under unambiguous, fast (=ACCTRAN), and slow (=DELTRAN) character optimizations. The latter two were equally considered due to no theoretical ground of favoring one of those (Agnarsson and Miller, 2008). Confidence was estimated by two resampling plans, jackknife and nonparametric bootstrap, and Bremer support. Jackknifing and bootstrapping were estimated from 500 replications or 500 pseudoreplicates respectively. Bremer supports were calculated with commands that searched for suboptimal trees at least five steps longer than the shortest tree (command: ‘bs 5;’) . The Bremer support ‘five’ represents a threshold of high support (Wenzel, 2002) and thus the values higher than five were not calculated. I regard more than 90% of jackknife or bootstrap and more than ‘3’ of Bremer supports (Davis, 1995; DeBry, 2001) as strong nodal supports. A strict consensus tree (Sokal and Rohlf, 1981) was used to summarize results. Biogeography Cladistic biogeography of Wockia was traced using the Divergence– Vicariance analysis (DIVA ver. 1.1: Ronquist, 1996). Five general distribution areas were used: Europe, eastern Palearctic (Korea and Japan), Oriental, Neotropical, and Nearctic regions. If a species is found in more than one area, all areas were coded for its extant distribution. The most parsimonious tree from my cladistic analysis was used as a tree topology for the DIVA analysis. Optimization parameters were set as (1) default (no limitation in maximum areas) and (2) two maximum areas (“maxareas = 2”).
male genitalia; having the shorter appendix bursae; and the membranous posterior half of ductus bursae in the female genitalia. Description. Head: Vertex and frontoclypeus dark brownish gray. Antenna laminate in the male, filiform in the female, 1/2 as long as forewing; flagellomeres silvery gray dorsally, naked ventrally. Labial palpus upcurved to middle of frontoclypeus, gradually narrower apically, brownish gray, darkened ventrally; 2nd segment as long as 3rd, 2 × longer than 1st, with longer scale tuft distally. Thorax: Patagia pale brownish gray; tegula and mesonotum dark brownish gray. Foreleg with coxa dark brownish gray; femur, tibia and tarsus dark gray dorsally, pale gray ventrally; tarsomeres with a pale gray ring distally. Midleg with coxa pale brownish gray; femur dark brownish gray, paler in basal 1/3; tibia brownish gray, paler in distal 1/4; tarsomeres with a pale gray ring distally. Hindleg lustrous, yellowish gray; tibia hairy dorsally. Forewing length 7.8–8.0 mm (n = 2), elongate-elliptical, dark brownish gray, intermixed with pale gray scales (Fig. 1A); sub-basal line pale gray, with two small black round patches of raised
RESULTS Description of new species Wockia magna sp. nov. (Fig. 1A–D) Type. Holotype: ۅ, “HOLOTYPE/ Wockia/ magna Sohn/ 2014” [on red label], “[Japan] Tyubu-Nagano/ Tobiraonsen/ 13 IX 1953/ [leg] T. Kodama”, “Issiki/ Collection/ 1972”, “Genitalia slide ۅ/ By J C SOHN/ USNM 115144”, deposited in USNM. Paratype: 1 ۃ, South Korea, Seoul, Mt. Inwangsan, 23 VII 1998 (leg. J.C. Sohn), genitalia slide no. SJC-470, CNU. Diagnosis. Wockia magna is the largest of all Wockia species. Wockia magna is close to Wockia asperipunctella, which occurs in Europe and North America, in overall genital structures, but differs from the latter in having the shorter posterolateral projections on the uncus; the absence of a rectangular projection at the middle of the costa in the
Fig.1.ޓWockia magna sp. nov. (A) Habitus, dorsal view, holotype; (B) Male genital capsule, missing left costal process indicated by dotted line, holotype, USNM 115144; (C) Male phallus, holotype, USNM 115144; (D) Female abdominal segment VIII and genitalia, paratype, SJC-470. The numbers marked with line indicate the morphological characters encoded in Table 1 (character number: character state).
Phylogeny and biogeography of Wockia scales; antemedian line slightly oblique, pale gray, with two dark brown, shortened linear patches of raised scales; a black spot near tornus; fringe brownish gray. Hindwings (Fig. 1A) narrowly round apically, gray; fringe dark yellowish gray. Male genitalia (Fig. 1B): Uncus subtrapezoidal, emarginated apically, with long, hair-like setae laterally. Valva elongate, tapered distally, curved at distal 1/5, apex acuminate; costal process 2/3 as long as valva, curved at basal 1/3, setose, digitate apically; sacculus elliptical, dense-setose dorsomarginally. Juxta nearly 1/2 as long as valva, elongate in anterior 3/4, inverted-triangular in posterior 1/4, trifid apically. Saccus short, subtriangular. Phallus (Fig. 1C) straight, about 1.4 × as long as valva; vesica finely pleated longitudinally, with a broad granular zone. Abdominal segment VIII and female genitalia (Fig. 1D): Sternite VIII with trapezoidal protrusion posteromedially. Papilla analis short, semi-globular, setose. Ovipositor telescopic with two subdivisions. Apophysis anterioris as long as apophysis posterioris. A broad emargination present near to ostium bursae. Ductus bursae constricted medially, membranous on posterior half, strongly sclerotized on anterior half, sclerotized part broadened anteriorly, interrupted by membranous invagination on a side of upper 2/3. Inception of ductus seminalis on cervical area of corpus bursae. Corpus bursae elongate, 6 × longer than ductus bursae, cucumiform, posterior 1/3 bulged on one-side, minutely pleated, anterior 1/4 globular; two dentiform signa with denticles dorsally. Distribution. Japan (Honshu) and South Korea. Etymology. The species epithet is the Latin adjective magnus, meaning “large,” and refers to the size of the new species being larger than its congeners. Remarks. The male and female specimens of W. magna were paired by similarities in their size and forewing patterns, especially the positions of the raised scales of patches on the subbasal and the antemedian lines. The female specimen, however, possesses a slightly darker head with shorter ventral cilia on the flagellomeres. These differences may be due to individual variation or sexual dimorphism.
261
Spiladarcha adamskii and five species of Wockia formed a strong monophyletic group against Anchimacheta iodes (Fig. 2: 100 in both bootstrapping and jackknifing; > 5 in Bremer support). The resulting tree placed Wockia chewbacca and Spiladarcha adamskii together, apart from other four species of Wockia. A grouping of Wockia chewbacca and Spiladarcha adamskii was, however, very weakly supported (Fig. 2: < 50% in both bootstrapping and jackknifing; 1 in Bremer support). Four species, W. koreana, W. variata, W. asperipunctella, and W. magna, were recovered as a strongly-supported monophyletic group (Fig. 2: 91% in both bootstrapping and jackknifing; Bremer support = 4). The clade including these four species was split into two species pairs, W. magna + W. asperipunctella versus W. koreana + W. variata. The monophyly of the former pair was moderately supported (Fig. 2: bootstrap = 81; jackknife = 78; Bremer support = 2) with three unambiguous character changes (Fig. 3). Character optimizations for the resulting tree proposed four synapomorphies for the other pair including W. koreana and W. variata (Fig. 3). This grouping was, however, weakly supported (Fig. 2: bootstrap = 72; jackknife = 71; Bremer support = 2). Biogeography The DIVA result required two dispersals for the optimal reconstruction with no constraint (Fig. 4). When the maximum number of possible areas was set to two, the reconstruction required two steps longer, hence four dispersals. The more differences in the reconstructed ancestral areas
Phylogeny My phylogenetic analysis resulted in only a single most parsimonious tree (Fig. 2: tree length = 38, CI = 71, RI = 70).
Fig.2.ޓThe resulting most parsimonious tree for five species of Wockia and two outgroup taxa. The nodal supports are given by bootstrap/jackknife above nodes and by Bremer supports below nodes.
Fig.3.ޓThe optimized character state changes over the tree in Fig. 2. (A) Slow optimization; (B) Fast optimization. Closed circles indicate synapomorphies; open circles indicate homoplasies and reversals. The numbers above and within circles denote the character numbers, shown in Table 1, and character states respectively. Asterisks indicate unambiguous character changes.
262
J.-C. Sohn
rate clade (Fig. 2). This proposition, however, remains inconclusive, since there exists an uncertainty in the interrelationships of W. chewbacca, Spiladarcha adamskii and the clade including other four species of Wockia. A grouping of W. chewbacca and Spiladarcha adamskii was supported by three synapomorphies (Fig. 3): the triangular uncus (7); the lack of the posterolateral bulges on the uncus (9); and the presence of winkles in the ductus bursae near the ostium bursae (19). Wockia chewbacca, however, lacks four autapomorphies of Spiladarcha proposed by Sohn (2012). Similar to these results, Adamski et al. (2009) observed the distinctiveness of W. chewbacca from W. asperipunctella, the type species of Wockia, Fig.4.ޓAncestral distributions of five species of Wockia and two outgroup taxa, optimized in DIVA. based on DNA barcoding The phylogeny was modified from Fig. 2. The letters on left and right sides of each node indicate the region. All these observations optimized ancestral distributions with maximum areas ‘five’ and ‘two’ respectively: A = Europe; B = may support a new genus for W. temperate East Asia; C = Vietnam; D = Neotropics; E = North America. Dots indicate the extant districhewbacca. This species is bution of each species. Map modified from Wikimedia Commons (http://commons.wikimedia.org/wiki/ supposedly related to at least File:WorldMap.svg). three Neotropical species (Adamski et al., 2009; Sohn, depending on the setting of the maximum area number were 2013b): i.e. W. mexicana from Mexico, and W. diabolica and observed for the nodes closer to the root: no constraint W. tetroidon, both from Jamaica. Heppner (2008) described always yielded broader ancestral distribution except for W. Wockia pelotas from Brazil, based on a single female speckoreana + W. variata and W. chewbacca + S. adamskii imen in poor condition. Its relationship with other Neotropical clades. The DIVA analysis with the maximum area number Wockia remains uncertain (Sohn, 2013b). A close relation‘2’ proposed multiple optimizations in two nodes: W. magna + ship of W. chewbacca and W. mexicana was substantiated W. asperipunctella clade and a root of all the included speusing DNA barcodes (Adamski et al., 2009). cies. Kyrki (1988) characterized Wockia based on a single The optimal reconstruction using default (no constraint) species, W. asperipunctella, and did not specify the aposuggested that Wockia and Spiladarcha once occupied all morphies for the genus. Four species of Wockia from the the areas including New World, Eurasia and Asian tropics. Holarctic and Oriental regions formed a clade (Wockia The Holarctic and Oriental Wockia were then diverged by sensu stricto in Fig. 2) substantiated by five synapomorphies: i.e. the absence of spiniform scales on the distal 1/3 vicariance events (Fig. 4). Differing from this view, the sceof the 3rd labial palpal segment (1); the presence of few long nario with two maximum areas assumed postulated that setae on the posterior margin of the uncal plate (8); the Wockia and Spiladarcha may be originated from Neotropics absence of a band-like gnathos (10); the inception of ductus plus temperate East Asia and then dispersed into Oriental seminalis near the corpus bursae (24); and the presence of Region, North America and Europe (Fig. 4). two opposable signa (26). Of these characteristics, the latter DISCUSSION two are homoplastic in Urodus. Phylogeny of Wockia Interrelationships of four species within Wockia sensu The resulting phylogeny from my analyses showed that stricto were only partly resolved from my analyses. Of the Wockia and Spiladarcha form a monophyletic clade against species, Wockia asperipunctella and W. magna were Anchimacheta iodes. These two genera are distinct from grouped together with moderate supports (Fig. 2). This relationship was substantiated by three synapomorphies (Fig. Anchimacheta in two characteristics (character numbers in parentheses; see Fig. 3 and Table 1 for the characters): the 3): i.e., the presence of the tapering, elongate valvae (12); presence of two erected scales of sub-basal dots on the the presence of a long band-like juxta (16); and the presforewing (3); the presence of erected scales of small spot in ence of an appendix bursae as the site for the inception of the tornal area of forewing (4). The genus Wockia as curthe ductus seminalis (23). The elongate valva and juxta are rently defined, however, turned out to be non-monophyletic, similar to Urodus. In addition to these, a slow character optias W. chewbacca and Spiladarcha adamskii form a sepamization scheme identified two homoplasies (character # 22
Phylogeny and biogeography of Wockia and 25) to support the grouping of the two species (Fig. 3). The male and female of W. magna were paired by superficial similarity and thus they could represent two separate species. The phylogenetic analysis reflecting this possibility recovered the identical tree as Fig. 2 (data not shown), except that the node bearing W. asperipunctella and the male and female of W. magna as three terminals was poorly supported (64 in both jackknife and bootstrap). The lower support values might be due to the increased number of missing data when the male and female of W. magna are treated as two separate terminal taxa. The other two species of Wockia sensu stricto, W. koreana and W. variata, were paired with each other in the resulting phylogeny. Their monophyly was suggested by four synapomorphies including the vertical vestiture of head covered with broad scales (0); the dorsal and lateral areas of occiput with the same shaped scales (2); the presence of the transparent hyline region in the hindwing (5); and the presence of a lobe between the cucullus and the distal end of sacculus (14). In spite of these shared features, the pair was poorly supported (Fig. 2: 72 in bootstrap; 71 in jackknife). Their relatedness is indeed undermined by several differences in the genitalia; e.g. the presence of uncal processes and additional lobes on cucullus in the male genitalia of W. variata; and the presence of nearly entire sclerotization in the female ductus bursae of W. koreana. Wockia balikpapanella Kyrki described by a single female from Borneo is undoubtedly related with W. variata in the genital features (Heppner, 2010b). Biogeography of Wockia Historical biogeography of a group of organisms requires their robust phylogenetic relationships and divergence times. Only the formal information as achieved from this study is available for Wockia, whereas dating the phylogeny is, at present, infeasible due to the lack of fossils reliably assigned to Urodidae (Sohn et al., 2012). Therefore, temporal components in the biogeography of Wockia need to be inferred from their extant distribution and the circumstantial evidences from other lepidopteran groups (cladistics biogeography approaches). Grimaldi and Engel (2005) speculated that majority of the apoditrysian superfamilies originated during the Late Cretaceous in accordance with angiosperm diversification. Therefore, the divergence of Urodidae may date back to the Late Cretaceous to the early Cenozoic. When Wockia originated, however, remains uncertain due to the lack of a robust phylogeny for Urodidae. In my DIVA analyses, the ancestral area reconstruction for Wockia was critically affected, depending on how to constrain the maximum number of areas (Fig. 4). No constraint on the parameter suggested the broad distribution of ancestral Wockia spanning the Palearctic and the Oriental regions, and the New World. This renders that their extant distribution is largely due to vicariance (Fig. 4). In the other, constraining the reconstruction with the maximum area number ‘2’ proposed the rather narrow ancestral areas for Wockia and Spiladarcha including the temperate East Asia and the Neotropical region (Fig. 4). This proposal seems to be the more likely scenario for two reasons. First, the broad ancestral distribution of Wockia assumes a similarly broad distribution of ancestral Urodidae, which is contradicted from
263
the fact that all urodid genera other than Wockia occur exclusively in the New World. Second, there are no geological references (fossils and continental associations) justifying the exclusive distribution of Wockia in the four zoogeographical ecozones. In fact, DIVA optimization tends to recover a large ancestral distribution including most of all of the areas occupied by the terminal taxa (Ronquist, 1996). Therefore, the resulting reconstruction often assumes vicariance in deep nodes, unless constrained in the maximum number of unit areas (Kodandaramaiah, 2010). DIVA needs at least two unit areas to constrain the reconstruction. The proposed ancestral area of Wockia including the temperate East Asia and the Neotropical Region is very likely due to this limitation of DIVA. Both Wockia chewbacca and Spiladarcha adamskii are exclusively Neotropical. There also are no geological events supporting the exclusive connection between East Asian and Neotropical faunas. From these reasons, it is most likely that Wockia and Spiladarcha originated in the Neotropical Region and the former genus then radiated to the temperate East Asia through a bridge of North America. Consistent with this scenario, faunal exchanges between North America and South America during the Late Cretaceous and the early Cenozoic have been observed for mammals (Pascual and Jaureguizar, 1992) and snakes (Holman, 2000). All four species of Wockia from the Holarctic and Oriental regions formed a clade (Wockia s. str.) not containing the Neotropical W. chewbacca. An ancestral distribution of this clade suggested to include all areas where the four terminal taxa occur without a constraint in optimization in ancestral area reconstruction; and to include only the temperate East Asia from the scenario with the maximum area number constrained to ‘2’. The former scenario is less likely, and may be due to the same limitation of unconstrained DIVA analysis as for Wockia and Spiladarcha. In the other, the latter scenario seems to be coincided with faunal and floral exchanges between Eurasian and North American continents during Eocene (e.g., Tiffney, 1985; Eberle and Greenwood, 2011; Missiaen et al., 2011). Once established in the temperate East Asia, the Wockia s. str. was divided following two diversification pathways. One of those represented by W. koreana was dispersed to the Oriental Region where two congeners, W. variata and W. balikpapanella, occur. The other lineage gave rise to W. magna and W. asperipunctella, which are apparently related with each other. This diversification scenario, however, is tentative, because the interrelationships of the four congeners have not been fully resolved. Therefore, the diversification of Wockia following a route from South America to North America (W. asperipunctella) and then to East Asia and the Oriental Region is also equally possible. Two Holarctic species, W. magna and W. asperipunctella, seem to be diverged relatively recently, considering the similarities in their genital structures. My DIVA analysis with a least maximum area constraint suggested two alternative scenarios in their ancestral distribution (Fig. 4). One scenario is involved in the faunal connection between East Asia and North America. It is well known that such a connection, often referred to as the ‘Bering Land Bridge,’ actually existed at various times during the Pleistocene ice ages. The other scenario assumes a trans-Eurasian distribution of the
J.-C. Sohn
264
lineage, which is undermined by the absence of Wockia in other areas than Europe, Korea and Japan. Wockia asperipunctella had long been regarded as a European endemic. Heppner (1997) found the species from the northeastern United States. The Nearctic populations of W. asperipunctella were originally thought due to anthropogenic introduction (Heppner, 1997). Landry (1998), however, suggested that the species is native to North America. It is very possible that those two regional populations of W. asperipunctella resulted from the faunal exchanges between Europe and North America during the Quaternary glaciation. It is noteworthy that all three species of Wockia whose host plants are known are associated with Salicaceae. This may suggest that the biogeography of Wockia is associated with the evolutionary history of Salicaceae. This hypothesis, however, remains elusive since the phylogeny (Leskinen and Alström-Rapaport, 1999) and the fossil record (Collinson, 1992) of Salicaceae are incomplete. ACKNOWLEDGMENTS I would like to thank Prof. Akito Kawahara (University of Florida, Gainesville and FLMNH) for critically editing my manuscript and giving me advice on cladistics analyses. I am also grateful to Drs. Donald Davis and David Adamski (Department of Entomology, USNM) for allowing my examination of the specimens from the USNM collection. I especially appreciate Prof. Charles Mitter (University of Maryland, College Park) for his encouragement and financial support for my museum visit.
REFERENCES Adamski D, Boege K, Landry J-F, Sohn J-C (2009) Two new species of Wockia Heinemann (Lepidoptera: Urodidae) from coastal dry-forests in Western Mexico. Proc Entomol Soc Wash 111: 166–182 Agnarsson I, Miller JA (2008) Is ACCTRAN better than DELTRAN? Cladistics 24: 1–7 Bruand T (1851) Catalogue systématique et synonymique des Lépidoptères du Département du Doubs. Tineides. Mém Soc emul Doubs 3(1): 23–58 Carpenter JM (1992) Random cladistics. Cladistics 8: 147–153 Cho S, Zwick A, Regier J, Mitter C, Cummings M, Yao J, et al. (2011) Can deliberately incomplete gene sample augmentation improve a phylogeny estimate for the advanced moths and butterflies (Hexapoda: Lepidoptera)? Syst Biol 60: 782–796 Clarke JFG (1941) The preparation of slides of the genitalia of Lepidoptera. Bull Brooklyn Entomol Soc 36: 149–161 Collinson ME (1992) The early fossil history of Salicaceae: a brief review. Proc R Soc Edinb 98B: 155–167 Davis JI (1995) A phylogenetic structure for the monocotyledons, as inferred from chloroplast DNA restriction site variation, and a comparison of measure of clade support. Syst Bot 20: 503–527 DeBry RW (2001) Improving interpretation of the decay index for DNA sequence data. Syst Biol 50: 742–752 Eberle JJ, Greenwood DR (2011) Life at the top of the greenhouse Eocene world, A review of the Eocene flora and vertebrate fauna from Canada’s High Arctic. Geol Soc Am Bull 124: 3–23 Goloboff P (1999) NONA. Version 2.0. Published by the author, San Miguel de Tucumán Grimaldi DA, Engel MS (2005) Evolution of the Insects. Cambridge University Press, Cambridge Heinemann H (1870) Die Schmetterlinge Deutschlands und der Schwiez, systematisch Bearbeitet. Vieweg, Braunschweig Heppner JB (1997) Wockia asperipunctella in North America (Lepidoptera: Urodidae: Galacticinae). Holarc Lepid 4: 73–74
Heppner JB (1998) Classification of Lepidoptera, Part 1. Introduction. Holarc Lepid 5(Suppl. 1): 1–148 Heppner JB (2008) The genus Wockia and a new species from southern Brazil (Lepidoptera: Urodidae: Galacticinae). Lepid Nov 1: 116–118 Heppner JB (2010a) Incawockia, a new genus and species from Peru (Lepidoptera: Urodidae: Galacticinae). Lepid Nov 3: 154– 158 Heppner JB (2010b) Notes on Vietnam moths, 10. Notes on Wockia in Vietnam (Lepidoptera: Urodidae: Galacticinae). Lepid Nov 3: 165–168 Herrich-Schäffer AW (1854) Sammlung neuer oder wenig bekannter aussereuropäischer Schmetterlinge. Vol. 1. G. J. Manz, Regenburg Hodges RW (2005) Order Lepidoptera. In “Borror and DeLong’s Introduction to the Study of Insects 7th edition” Ed by CA Triplehorn, NF Johnson, Thomson Books/Cole, Belmont, pp 571–647 Holman JA (2000) Fossil Snakes of North America: Origin, Evolution, Distribution, Paleoecology. Indiana University Press, Bloomington Klots AB (1970) Lepidoptera. In “Taxonomist’s Glossary of Genitalia in Insects” Ed by SL Tuxen, Munksgaard, Copenhagen, pp 115–130 Kodandaramaiah U (2010) Use of dispersal–vicariance analysis in biogeography – a critique. J Biogeogr 37: 3–11 Kyrki J (1984) The Yponomeutoidea: a reassessment of the superfamily and its suprageneric groups (Lepidoptera). Entomol Scand 15: 71–84 Kyrki J (1986) Wockia balikpapanella sp. n. from Borneo (Lepidoptera, Yponomeutidae auct.). Ann Entomol Fenn 52: 42–43 Kyrki J (1988) The systematic position of Wockia Heinemann, 1870, and related genera (Lepidoptera: Ditrysia: Yponomeutidae auct.). Nota Lepid 11: 45–69 Landry JF (1998) Additional Nearctic records of Wockia asperipunctella, with notes on its distribution and structural variation (Lepidoptera: Urodidae). Holarc Lepid 5: 9–13 Leskinen E, Alström-Rapaport C (1999) Molecular phylogeny of Salicaceae and closely related Flacourtiaceae: evidence from 5.8 S, ITS 1 and ITS 2 of the rDNA. Pl Syst Evol 215: 209–227 Meyrick E (1913) Hyponomeutidae. In “Exotic Microlepidoptera. Vol. 1” Ed by E Meyrick, E. W. Classey Limited, Middlesex, pp 65– 176 Missiaen P, Gunnell GF, Gingerich PD (2011) New Brontotheriidae (Mammalia, Perissodactyla) from the Early and Middle Eocene of Pakistan with implications for mammalian paleobiogeography. J Paleo 85: 665–677 Mutanen M, Wahlberg N, Kaila L (2010) Comprehensive gene and taxon coverage elucidates radiation patterns in moths and butterflies. Proc R Soc B 277: 2839–2848 Nixon KC (2002) WINCLADA. Version 1.0. Published by the author, New York Pascual R, Jaureguizar EO (1992) Evolutionary pattern of land mammal faunas during the Late Cretaceous and Paleocene in South America: a comparison with the North American pattern. Ann Zool Fenn 28: 245–252 Regier JC, Mitter C, Zwick A, Bazinet AL, Cummings MP, Kawahara AY, et al. (2013) A large-scale, higher-level, molecular phylogenetic study of the insect order Lepidoptera (moths and butterflies). PLoS ONE 8(3): e58568 Ronquist F (1996) DIVA version 1.1. Computer program and manual available by anonymous FTP from Uppsala University (ftp.systbot.uu.se) Sohn J-C (2012) Taxonomic review of Spiladarcha Meyrick (Lepidoptera: Urodoidea: Urodidae) with a new species from Venezuela. Entomol Sci 15: 303–308
Phylogeny and biogeography of Wockia Sohn J-C (2013a) A new genus and species of Urodidae (Lepidoptera: Urodoidea) from Argentina with the first report of asymmetric male genitalia in the superfamily. Fla Entomol 96: 496–476 Sohn J-C (2013b) Two new species of Wockia Heinemann (Lepidoptera: Urodidae) from Jamaica. Neotrop Entomol 42: 300–303 Sohn J-C, Adamski D (2008) A new species of Wockia Heinemann, 1890 [sic] (Lepidoptera: Urodidae) from Korea. Proc Entomol Soc Wash 111: 556–561 Sohn J-C, Park K-T (2009) A new species of Wockia Heinemann, 1870 (Lepidoptera: Urodidae) from Vietnam. Trop Lepid Res 19: 62–63 Sohn J-C, Labandeira C, Davis D, Mitter C (2012) An annotated catalog of fossil and subfossil Lepidoptera (Insecta: Holometabola) of the world. Zootaxa 3286: 1–132 Sohn J-C, Regier JC, Mitter C, Davis D, Landry J-F, Zwick A, et al. (2013) A molecular phylogeny for Yponomeutoidea (Insecta, Lepidoptera, Ditrysia) and its implications for classification, bio-
265
geography and the evolution of host plant use. PLoS ONE 8(1): e55066 Sokal RR, Rohlf FJ (1981) Taxonomic congruence in the Leptopodomorpha re-examined. Syst Zool 30: 309–325 Tiffney BH (1985) The Eocene North Atlantic Land Bridge: its importance in Tertiary and modern phytogeography of the northern hemisphere. J Arnold Arboretum 66: 243–273 Walsingham L (1914) Anchimacheta. In “Biologia Centrali Americana. Zoology: Insecta. Lepidoptera-Heterocera. Vol. 4. Tineina, Pterophorina, Orneodina, Pyralidina, and Hepialina” Ed by FD Godman, O Salvin, Taylor & Francis, London, pp 323–324 Wenzel JW (2002) Phylogenetic analysis: the basic method. In “Techniques in Molecular Systematics and Evolution” Ed by R Desalle, G Giribet, W Wheeler, Birkhäuser Verlag, Basel, pp 4– 30 Wootton RJ (1979) Function, homology and terminology in insect wings. Syst Entomol 4: 81–93 (Received September 23, 2013 / Accepted December 27, 2013)
BioOne (www.bioone.org) is a nonprofit, online aggregation of core research in the biological, ecological, and environmental sciences. BioOne provides a sustainable online platform for over 170 journals and books published by nonprofit societies, associations, museums, institutions, and presses. Your use of this PDF, the BioOne Web site, and all posted and associated content indicates your acceptance of BioOne’s Terms of Use, available at www.bioone.org/page/terms_of_use. Usage of BioOne content is strictly limited to personal, educational, and non-commercial use. Commercial inquiries or rights and permissions requests should be directed to the individual publisher as copyright holder.
BioOne sees sustainable scholarly publishing as an inherently collaborative enterprise connecting authors, nonprofit publishers, academic institutions, research libraries, and research funders in the common goal of maximizing access to critical research.
¤ 2014 Zoological Society of Japan
ZOOLOGICAL SCIENCE 31: 258–265 (2014)
Morphology-Based Phylogeny and Biogeography of Wockia (Lepidoptera: Urodidae) with Description of a New Species from Japan and South Korea Jae-Cheon Sohn1,2* 1
Department of Entomology, University of Maryland, 4112 Plant Sciences Building, College Park, MD 20742, USA 2 Department of Entomology, Smithsonian Institution, National Museum of Natural History, 10th & Constitution NW, Washington, DC 20560, USA
Wockia, one of the six genera within the lepidopteran family Urodidae, currently includes nine species distributed in Holarctic, Oriental, and Neotropical regions. A new species of the genus, W. magna sp. nov., is described from Japan and South Korea. This is the first record of Urodidae from Japan. A cladistic analysis was constructed based on 27 morphological characters from five ingroup species and two outgroup taxa. A single most parsimonious tree was found (length = 38, CI = 71, RI = 70). The resulting tree failed to recover the monophyly of Wockia chewbacca and a clade Wockia sensu stricto, including four congeners from North America, temperate East Asia and Southeast Asia. Wockia sensu stricto was resolved as monophyletic and was divided into two subgroups, one including W. asperipunctella and W. magna, and the other including W. koreana and W. variata. The sister-group relationship of W. asperipunctella and W. magna was moderately supported. Synapomorphies of Wockia sensu stricto are provided from the resulting phylogeny. Systematic definitions of Wockia and other allied genera are revised. Optimal ancestral area reconstruction implemented in DIVA resulted in different hypotheses for Wockia, depending on how to constrain the maximum number of areas. The maximum area number set as two yielded a more likely scenario suggesting that Wockia sensu stricto originated in temperate East Asia and then dispersed into the Oriental region, North America, and Europe. A working hypothesis and other equally possible alternative explanations for the biogeography of Wockia are provided. Key words: biogeography, cladistic analysis, taxonomy, Urodoidea, Wockia INTRODUCTION The False Burnet moth family, Urodidae currently include six genera: Urodus Herrich-Schäffer, 1854, Wockia Heinemann, 1870, Spildarcha Meyrick, 1913, Anchimacheta Walsingham, 1914, Incawockia Heppner, 2010a, and Anomalomeuta Sohn, 2013a. Many of the included species have long been misidentified as Yponomeutidae or Plutellidae. Kyrki (1984) first suggested that they are distinguishable from Yponomeutoidea in having the tortricoid type of thoracic-abdominal articulation and hence representing close ties with Apoditrysia. Later, Kyrki (1988) assigned them to their own family Urodidae. This proposal has been corroborated by two recent molecular studies (Sohn et al., 2013; Regier et al., 2013), whereas their systematic position within Apoditrysia remains unclear. Kyrki (1988) noticed that Urodidae share some characteristics with Schreckensteiniidae, such as the elongate prolegs of the larvae and the loosely-meshed cocoon, although those similarities are not * Corresponding author. Tel. : +1-301-717-5462; Fax : +1-301-314-9290; E-mail : [email protected]; [email protected] Supplemental material for this article is available online. doi:10.2108/zs130199
exclusively unique to those two families. Among molecular phylogenetic studies, Mutanen et al. (2010) recovered Schreckensteiniidae as sister to Urodidae. On the other hand, Cho et al. (2011) found that the Urodidae are a sister taxon to Alucitidae, Choreutidae, Pterophoridae, or Tortricidae, depending on the data set. In Regier et al. (2013), Urodidae were identified as sister to the rest of apoditrysian families. None of the abovementioned studies, however, resolved the relationships with significant support. Wockia is a relatively small genus comprising gray or fuscous micro-moths with an oblique antemedian band of raised scales (Kyrki, 1986). Members in the genus are very similar to those in Blastobasidae or Phycitinae (Pyralidae) in superficial appearance (Landry, 1998). Kyrki (1988) provided generic features, based on W. asperipunctella (Bruand, 1851), the only confirmed species of the genus in his study. Since then, the species diversity of Wockia has increased with discoveries of new species from several faunal regions. Wockia currently include nine described species: W. asperipunctella from Europe and North America (Kyrki, 1988; Heppner, 1997; Landry, 1998); W. balikpapanella Kyrki, 1986 from Borneo (Kyrki, 1986); W. koreana Sohn, 2008 from South Korea (Sohn and Adamski, 2008); W. variata Sohn and Park, 2009 from North Vietnam (Sohn and Park, 2009; Heppner, 2010b); two Mexican species, W.
Phylogeny and biogeography of Wockia chewbacca Adamski, 2009 and W. mexicana Adamski, 2009 (Adamski et al., 2009); two Jamaican species, W. diabloca Sohn, 2013 and W. tetroidon Sohn, 2013 (Sohn, 2013b); and W. pelotas Heppner, 2008 from Brazil (Heppner, 2008). This increase in species diversity has prompted a reconsideration of Kyrki’s (1988) definition of the genus. Global distribution of Wockia includes at least four zoogeographic regions, in contrast to the other five urodid genera, Anchimacheta, Anomalomeuta, Incawockia, Spiladarcha, and Urodus, which are entirely or predominantly Neotropical. Such widely scattered occurrence of Wockia may not be easy to understand without phylogenetic approaches. Thus far, however, no phylogenetic study has attempted to resolve the interspecific relationships or the biogeography of Wockia. The objectives of this study are to: 1) construct a phylogeny of Wockia, based on their morphologies, 2) understand their biogeography, and 3) describe a new species from Japan and South Korea that is the second congener from the temperate East Asia. MATERIALS AND METHODS Preparation of data set A total of seven taxa, including two outgroup taxa, Spiladarcha adamskii Sohn, 2012 and Anchimacheta iodes Walsingham, 1914, were sampled. Pinned specimens were obtained from the following collections (also see Supplementary Table S1 online): Natural History Museum, London, United Kingdom (BMNH); Chungbuk
259
National University, Cheongju, South Korea (CNU); Florida Museum of Natural History, Gainesville, Florida, USA (FLMNH); Korea National Arboretum, Pocheon, Kyonggi, South Korea (KNA); United States Museum of Natural History, Washington DC, USA (USNM). External morphologies were examined with a Leica MZ APO stereoscope and photographed using a Nikon D40 digital camera. Genitalia slides were prepared following Clarke (1941), with the modifications that Chlorazol Black was used for staining and Euparal resin for permanent slide mounting. The wings were slidemounted according to Hodges (2005). Slide-mounted specimens were examined with a Leica LETTZ-DMRX microscope. Terminology follows Klots (1970) for genitalia; Wootton (1979) for wing venation; and Heppner (1998) for other body parts. Ovipositor is defined as the segments and intersegmental membranes after the abdominal segment VIII. Verbatim label data are given only for holotype. Character coding The data matrix was constructed using WinClada ver. 1.0 (Nixon, 2002). A total of 27 adult morphological characters were coded. Parsimony-uninformative characters were excluded to avoid their influence on branch support calculation (Carpenter, 1992). The morphological characters (Table 1) were retrieved from the heads (n = 3), wings (n = 3), male genitalia (n = 12), and female genitalia (n = 9). All encoded characters were binary, non-additive, and equally weighted. Missing data were encoded as “?” and inapplicable character states as “–”. The data matrix is available at the Supplementary Table S2 online.
Table1.ޓCharacters coded for cladistic analysis. Characters Head 0: Scales on the vertex in females 1: Spiniform scales on terminal 1/3 of 3rd labial palpal segment 2: Scales on dorsal and lateral areas of occiput Wings 3: Erected scales of two subbasal dots on forewing 4: Erected scales of small spot near forewing tornus 5: Transparent hyaline area on hindwing Male genitalia 6: An articulation between tegumen and uncus 7: Shape of uncus 8: Long setae on the top of uncus 9: Posterolateral bulges or projections on uncus 10: Gnathal arms 11: Corona with strong spines 12: Stick-like terminal elongation of valva 13: Terminal saccular angulation 14: Nonsetose lobe beyond saccular angulation 15: Costal process 16: Shape of juxta 17: Spiniform cornutus or spinulate cornuti Female genitalia 18: Position of ostium bursae 19: Winkles in ductus bursae near to ostium bursae 20: Antrum 21: Sclerotization of antrum 22: Length of ductus bursae with relation to corpus bursae 23: Burge at the junction of ductus seminalis 24: Entry of ductus seminalis 25: Corpus bursae 26: Two identical signa
Character States 0) piliform, 1) broad 0) absent, 1) present 0) dorsal scales longer than lateral scales, 1) dorsal and lateral scales similar in size 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) triangular, 1) rectangular 0) absent, 1) present 0) absent, 1) present 0) not continuous or reduced, 1) continuous 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) absent, 1) present 0) U-shaped, 1) elongate 0) absent, 1) present 0) on or after anterior margin of sternite VII, 1) before anterior margin of sternite VII 0) absent, 1) present 0) absent, 1) present 0) complete, 1) partial 0) longer or almost same length, 1) shorter 0) absent, 1) present 0) on the way of ductus bursae, 1) near to corpus bursae 0) globular or ovoid, 1) elongate-elliptical 0) absent, 1) present
J.-C. Sohn
260
Phylogenetic analysis Parsimony analyses were performed using WinClada ver. 1.0 (Nixon, 2002) which implements NONA ver. 2.0 (Goloboff, 1999) to search for the most parsimonious tree. The following commands were used: hold 1000, hold/100, mult*1000, max*, and mswap+. The resulting trees were examined with WinClada under unambiguous, fast (=ACCTRAN), and slow (=DELTRAN) character optimizations. The latter two were equally considered due to no theoretical ground of favoring one of those (Agnarsson and Miller, 2008). Confidence was estimated by two resampling plans, jackknife and nonparametric bootstrap, and Bremer support. Jackknifing and bootstrapping were estimated from 500 replications or 500 pseudoreplicates respectively. Bremer supports were calculated with commands that searched for suboptimal trees at least five steps longer than the shortest tree (command: ‘bs 5;’) . The Bremer support ‘five’ represents a threshold of high support (Wenzel, 2002) and thus the values higher than five were not calculated. I regard more than 90% of jackknife or bootstrap and more than ‘3’ of Bremer supports (Davis, 1995; DeBry, 2001) as strong nodal supports. A strict consensus tree (Sokal and Rohlf, 1981) was used to summarize results. Biogeography Cladistic biogeography of Wockia was traced using the Divergence– Vicariance analysis (DIVA ver. 1.1: Ronquist, 1996). Five general distribution areas were used: Europe, eastern Palearctic (Korea and Japan), Oriental, Neotropical, and Nearctic regions. If a species is found in more than one area, all areas were coded for its extant distribution. The most parsimonious tree from my cladistic analysis was used as a tree topology for the DIVA analysis. Optimization parameters were set as (1) default (no limitation in maximum areas) and (2) two maximum areas (“maxareas = 2”).
male genitalia; having the shorter appendix bursae; and the membranous posterior half of ductus bursae in the female genitalia. Description. Head: Vertex and frontoclypeus dark brownish gray. Antenna laminate in the male, filiform in the female, 1/2 as long as forewing; flagellomeres silvery gray dorsally, naked ventrally. Labial palpus upcurved to middle of frontoclypeus, gradually narrower apically, brownish gray, darkened ventrally; 2nd segment as long as 3rd, 2 × longer than 1st, with longer scale tuft distally. Thorax: Patagia pale brownish gray; tegula and mesonotum dark brownish gray. Foreleg with coxa dark brownish gray; femur, tibia and tarsus dark gray dorsally, pale gray ventrally; tarsomeres with a pale gray ring distally. Midleg with coxa pale brownish gray; femur dark brownish gray, paler in basal 1/3; tibia brownish gray, paler in distal 1/4; tarsomeres with a pale gray ring distally. Hindleg lustrous, yellowish gray; tibia hairy dorsally. Forewing length 7.8–8.0 mm (n = 2), elongate-elliptical, dark brownish gray, intermixed with pale gray scales (Fig. 1A); sub-basal line pale gray, with two small black round patches of raised
RESULTS Description of new species Wockia magna sp. nov. (Fig. 1A–D) Type. Holotype: ۅ, “HOLOTYPE/ Wockia/ magna Sohn/ 2014” [on red label], “[Japan] Tyubu-Nagano/ Tobiraonsen/ 13 IX 1953/ [leg] T. Kodama”, “Issiki/ Collection/ 1972”, “Genitalia slide ۅ/ By J C SOHN/ USNM 115144”, deposited in USNM. Paratype: 1 ۃ, South Korea, Seoul, Mt. Inwangsan, 23 VII 1998 (leg. J.C. Sohn), genitalia slide no. SJC-470, CNU. Diagnosis. Wockia magna is the largest of all Wockia species. Wockia magna is close to Wockia asperipunctella, which occurs in Europe and North America, in overall genital structures, but differs from the latter in having the shorter posterolateral projections on the uncus; the absence of a rectangular projection at the middle of the costa in the
Fig.1.ޓWockia magna sp. nov. (A) Habitus, dorsal view, holotype; (B) Male genital capsule, missing left costal process indicated by dotted line, holotype, USNM 115144; (C) Male phallus, holotype, USNM 115144; (D) Female abdominal segment VIII and genitalia, paratype, SJC-470. The numbers marked with line indicate the morphological characters encoded in Table 1 (character number: character state).
Phylogeny and biogeography of Wockia scales; antemedian line slightly oblique, pale gray, with two dark brown, shortened linear patches of raised scales; a black spot near tornus; fringe brownish gray. Hindwings (Fig. 1A) narrowly round apically, gray; fringe dark yellowish gray. Male genitalia (Fig. 1B): Uncus subtrapezoidal, emarginated apically, with long, hair-like setae laterally. Valva elongate, tapered distally, curved at distal 1/5, apex acuminate; costal process 2/3 as long as valva, curved at basal 1/3, setose, digitate apically; sacculus elliptical, dense-setose dorsomarginally. Juxta nearly 1/2 as long as valva, elongate in anterior 3/4, inverted-triangular in posterior 1/4, trifid apically. Saccus short, subtriangular. Phallus (Fig. 1C) straight, about 1.4 × as long as valva; vesica finely pleated longitudinally, with a broad granular zone. Abdominal segment VIII and female genitalia (Fig. 1D): Sternite VIII with trapezoidal protrusion posteromedially. Papilla analis short, semi-globular, setose. Ovipositor telescopic with two subdivisions. Apophysis anterioris as long as apophysis posterioris. A broad emargination present near to ostium bursae. Ductus bursae constricted medially, membranous on posterior half, strongly sclerotized on anterior half, sclerotized part broadened anteriorly, interrupted by membranous invagination on a side of upper 2/3. Inception of ductus seminalis on cervical area of corpus bursae. Corpus bursae elongate, 6 × longer than ductus bursae, cucumiform, posterior 1/3 bulged on one-side, minutely pleated, anterior 1/4 globular; two dentiform signa with denticles dorsally. Distribution. Japan (Honshu) and South Korea. Etymology. The species epithet is the Latin adjective magnus, meaning “large,” and refers to the size of the new species being larger than its congeners. Remarks. The male and female specimens of W. magna were paired by similarities in their size and forewing patterns, especially the positions of the raised scales of patches on the subbasal and the antemedian lines. The female specimen, however, possesses a slightly darker head with shorter ventral cilia on the flagellomeres. These differences may be due to individual variation or sexual dimorphism.
261
Spiladarcha adamskii and five species of Wockia formed a strong monophyletic group against Anchimacheta iodes (Fig. 2: 100 in both bootstrapping and jackknifing; > 5 in Bremer support). The resulting tree placed Wockia chewbacca and Spiladarcha adamskii together, apart from other four species of Wockia. A grouping of Wockia chewbacca and Spiladarcha adamskii was, however, very weakly supported (Fig. 2: < 50% in both bootstrapping and jackknifing; 1 in Bremer support). Four species, W. koreana, W. variata, W. asperipunctella, and W. magna, were recovered as a strongly-supported monophyletic group (Fig. 2: 91% in both bootstrapping and jackknifing; Bremer support = 4). The clade including these four species was split into two species pairs, W. magna + W. asperipunctella versus W. koreana + W. variata. The monophyly of the former pair was moderately supported (Fig. 2: bootstrap = 81; jackknife = 78; Bremer support = 2) with three unambiguous character changes (Fig. 3). Character optimizations for the resulting tree proposed four synapomorphies for the other pair including W. koreana and W. variata (Fig. 3). This grouping was, however, weakly supported (Fig. 2: bootstrap = 72; jackknife = 71; Bremer support = 2). Biogeography The DIVA result required two dispersals for the optimal reconstruction with no constraint (Fig. 4). When the maximum number of possible areas was set to two, the reconstruction required two steps longer, hence four dispersals. The more differences in the reconstructed ancestral areas
Phylogeny My phylogenetic analysis resulted in only a single most parsimonious tree (Fig. 2: tree length = 38, CI = 71, RI = 70).
Fig.2.ޓThe resulting most parsimonious tree for five species of Wockia and two outgroup taxa. The nodal supports are given by bootstrap/jackknife above nodes and by Bremer supports below nodes.
Fig.3.ޓThe optimized character state changes over the tree in Fig. 2. (A) Slow optimization; (B) Fast optimization. Closed circles indicate synapomorphies; open circles indicate homoplasies and reversals. The numbers above and within circles denote the character numbers, shown in Table 1, and character states respectively. Asterisks indicate unambiguous character changes.
262
J.-C. Sohn
rate clade (Fig. 2). This proposition, however, remains inconclusive, since there exists an uncertainty in the interrelationships of W. chewbacca, Spiladarcha adamskii and the clade including other four species of Wockia. A grouping of W. chewbacca and Spiladarcha adamskii was supported by three synapomorphies (Fig. 3): the triangular uncus (7); the lack of the posterolateral bulges on the uncus (9); and the presence of winkles in the ductus bursae near the ostium bursae (19). Wockia chewbacca, however, lacks four autapomorphies of Spiladarcha proposed by Sohn (2012). Similar to these results, Adamski et al. (2009) observed the distinctiveness of W. chewbacca from W. asperipunctella, the type species of Wockia, Fig.4.ޓAncestral distributions of five species of Wockia and two outgroup taxa, optimized in DIVA. based on DNA barcoding The phylogeny was modified from Fig. 2. The letters on left and right sides of each node indicate the region. All these observations optimized ancestral distributions with maximum areas ‘five’ and ‘two’ respectively: A = Europe; B = may support a new genus for W. temperate East Asia; C = Vietnam; D = Neotropics; E = North America. Dots indicate the extant districhewbacca. This species is bution of each species. Map modified from Wikimedia Commons (http://commons.wikimedia.org/wiki/ supposedly related to at least File:WorldMap.svg). three Neotropical species (Adamski et al., 2009; Sohn, depending on the setting of the maximum area number were 2013b): i.e. W. mexicana from Mexico, and W. diabolica and observed for the nodes closer to the root: no constraint W. tetroidon, both from Jamaica. Heppner (2008) described always yielded broader ancestral distribution except for W. Wockia pelotas from Brazil, based on a single female speckoreana + W. variata and W. chewbacca + S. adamskii imen in poor condition. Its relationship with other Neotropical clades. The DIVA analysis with the maximum area number Wockia remains uncertain (Sohn, 2013b). A close relation‘2’ proposed multiple optimizations in two nodes: W. magna + ship of W. chewbacca and W. mexicana was substantiated W. asperipunctella clade and a root of all the included speusing DNA barcodes (Adamski et al., 2009). cies. Kyrki (1988) characterized Wockia based on a single The optimal reconstruction using default (no constraint) species, W. asperipunctella, and did not specify the aposuggested that Wockia and Spiladarcha once occupied all morphies for the genus. Four species of Wockia from the the areas including New World, Eurasia and Asian tropics. Holarctic and Oriental regions formed a clade (Wockia The Holarctic and Oriental Wockia were then diverged by sensu stricto in Fig. 2) substantiated by five synapomorphies: i.e. the absence of spiniform scales on the distal 1/3 vicariance events (Fig. 4). Differing from this view, the sceof the 3rd labial palpal segment (1); the presence of few long nario with two maximum areas assumed postulated that setae on the posterior margin of the uncal plate (8); the Wockia and Spiladarcha may be originated from Neotropics absence of a band-like gnathos (10); the inception of ductus plus temperate East Asia and then dispersed into Oriental seminalis near the corpus bursae (24); and the presence of Region, North America and Europe (Fig. 4). two opposable signa (26). Of these characteristics, the latter DISCUSSION two are homoplastic in Urodus. Phylogeny of Wockia Interrelationships of four species within Wockia sensu The resulting phylogeny from my analyses showed that stricto were only partly resolved from my analyses. Of the Wockia and Spiladarcha form a monophyletic clade against species, Wockia asperipunctella and W. magna were Anchimacheta iodes. These two genera are distinct from grouped together with moderate supports (Fig. 2). This relationship was substantiated by three synapomorphies (Fig. Anchimacheta in two characteristics (character numbers in parentheses; see Fig. 3 and Table 1 for the characters): the 3): i.e., the presence of the tapering, elongate valvae (12); presence of two erected scales of sub-basal dots on the the presence of a long band-like juxta (16); and the presforewing (3); the presence of erected scales of small spot in ence of an appendix bursae as the site for the inception of the tornal area of forewing (4). The genus Wockia as curthe ductus seminalis (23). The elongate valva and juxta are rently defined, however, turned out to be non-monophyletic, similar to Urodus. In addition to these, a slow character optias W. chewbacca and Spiladarcha adamskii form a sepamization scheme identified two homoplasies (character # 22
Phylogeny and biogeography of Wockia and 25) to support the grouping of the two species (Fig. 3). The male and female of W. magna were paired by superficial similarity and thus they could represent two separate species. The phylogenetic analysis reflecting this possibility recovered the identical tree as Fig. 2 (data not shown), except that the node bearing W. asperipunctella and the male and female of W. magna as three terminals was poorly supported (64 in both jackknife and bootstrap). The lower support values might be due to the increased number of missing data when the male and female of W. magna are treated as two separate terminal taxa. The other two species of Wockia sensu stricto, W. koreana and W. variata, were paired with each other in the resulting phylogeny. Their monophyly was suggested by four synapomorphies including the vertical vestiture of head covered with broad scales (0); the dorsal and lateral areas of occiput with the same shaped scales (2); the presence of the transparent hyline region in the hindwing (5); and the presence of a lobe between the cucullus and the distal end of sacculus (14). In spite of these shared features, the pair was poorly supported (Fig. 2: 72 in bootstrap; 71 in jackknife). Their relatedness is indeed undermined by several differences in the genitalia; e.g. the presence of uncal processes and additional lobes on cucullus in the male genitalia of W. variata; and the presence of nearly entire sclerotization in the female ductus bursae of W. koreana. Wockia balikpapanella Kyrki described by a single female from Borneo is undoubtedly related with W. variata in the genital features (Heppner, 2010b). Biogeography of Wockia Historical biogeography of a group of organisms requires their robust phylogenetic relationships and divergence times. Only the formal information as achieved from this study is available for Wockia, whereas dating the phylogeny is, at present, infeasible due to the lack of fossils reliably assigned to Urodidae (Sohn et al., 2012). Therefore, temporal components in the biogeography of Wockia need to be inferred from their extant distribution and the circumstantial evidences from other lepidopteran groups (cladistics biogeography approaches). Grimaldi and Engel (2005) speculated that majority of the apoditrysian superfamilies originated during the Late Cretaceous in accordance with angiosperm diversification. Therefore, the divergence of Urodidae may date back to the Late Cretaceous to the early Cenozoic. When Wockia originated, however, remains uncertain due to the lack of a robust phylogeny for Urodidae. In my DIVA analyses, the ancestral area reconstruction for Wockia was critically affected, depending on how to constrain the maximum number of areas (Fig. 4). No constraint on the parameter suggested the broad distribution of ancestral Wockia spanning the Palearctic and the Oriental regions, and the New World. This renders that their extant distribution is largely due to vicariance (Fig. 4). In the other, constraining the reconstruction with the maximum area number ‘2’ proposed the rather narrow ancestral areas for Wockia and Spiladarcha including the temperate East Asia and the Neotropical region (Fig. 4). This proposal seems to be the more likely scenario for two reasons. First, the broad ancestral distribution of Wockia assumes a similarly broad distribution of ancestral Urodidae, which is contradicted from
263
the fact that all urodid genera other than Wockia occur exclusively in the New World. Second, there are no geological references (fossils and continental associations) justifying the exclusive distribution of Wockia in the four zoogeographical ecozones. In fact, DIVA optimization tends to recover a large ancestral distribution including most of all of the areas occupied by the terminal taxa (Ronquist, 1996). Therefore, the resulting reconstruction often assumes vicariance in deep nodes, unless constrained in the maximum number of unit areas (Kodandaramaiah, 2010). DIVA needs at least two unit areas to constrain the reconstruction. The proposed ancestral area of Wockia including the temperate East Asia and the Neotropical Region is very likely due to this limitation of DIVA. Both Wockia chewbacca and Spiladarcha adamskii are exclusively Neotropical. There also are no geological events supporting the exclusive connection between East Asian and Neotropical faunas. From these reasons, it is most likely that Wockia and Spiladarcha originated in the Neotropical Region and the former genus then radiated to the temperate East Asia through a bridge of North America. Consistent with this scenario, faunal exchanges between North America and South America during the Late Cretaceous and the early Cenozoic have been observed for mammals (Pascual and Jaureguizar, 1992) and snakes (Holman, 2000). All four species of Wockia from the Holarctic and Oriental regions formed a clade (Wockia s. str.) not containing the Neotropical W. chewbacca. An ancestral distribution of this clade suggested to include all areas where the four terminal taxa occur without a constraint in optimization in ancestral area reconstruction; and to include only the temperate East Asia from the scenario with the maximum area number constrained to ‘2’. The former scenario is less likely, and may be due to the same limitation of unconstrained DIVA analysis as for Wockia and Spiladarcha. In the other, the latter scenario seems to be coincided with faunal and floral exchanges between Eurasian and North American continents during Eocene (e.g., Tiffney, 1985; Eberle and Greenwood, 2011; Missiaen et al., 2011). Once established in the temperate East Asia, the Wockia s. str. was divided following two diversification pathways. One of those represented by W. koreana was dispersed to the Oriental Region where two congeners, W. variata and W. balikpapanella, occur. The other lineage gave rise to W. magna and W. asperipunctella, which are apparently related with each other. This diversification scenario, however, is tentative, because the interrelationships of the four congeners have not been fully resolved. Therefore, the diversification of Wockia following a route from South America to North America (W. asperipunctella) and then to East Asia and the Oriental Region is also equally possible. Two Holarctic species, W. magna and W. asperipunctella, seem to be diverged relatively recently, considering the similarities in their genital structures. My DIVA analysis with a least maximum area constraint suggested two alternative scenarios in their ancestral distribution (Fig. 4). One scenario is involved in the faunal connection between East Asia and North America. It is well known that such a connection, often referred to as the ‘Bering Land Bridge,’ actually existed at various times during the Pleistocene ice ages. The other scenario assumes a trans-Eurasian distribution of the
J.-C. Sohn
264
lineage, which is undermined by the absence of Wockia in other areas than Europe, Korea and Japan. Wockia asperipunctella had long been regarded as a European endemic. Heppner (1997) found the species from the northeastern United States. The Nearctic populations of W. asperipunctella were originally thought due to anthropogenic introduction (Heppner, 1997). Landry (1998), however, suggested that the species is native to North America. It is very possible that those two regional populations of W. asperipunctella resulted from the faunal exchanges between Europe and North America during the Quaternary glaciation. It is noteworthy that all three species of Wockia whose host plants are known are associated with Salicaceae. This may suggest that the biogeography of Wockia is associated with the evolutionary history of Salicaceae. This hypothesis, however, remains elusive since the phylogeny (Leskinen and Alström-Rapaport, 1999) and the fossil record (Collinson, 1992) of Salicaceae are incomplete. ACKNOWLEDGMENTS I would like to thank Prof. Akito Kawahara (University of Florida, Gainesville and FLMNH) for critically editing my manuscript and giving me advice on cladistics analyses. I am also grateful to Drs. Donald Davis and David Adamski (Department of Entomology, USNM) for allowing my examination of the specimens from the USNM collection. I especially appreciate Prof. Charles Mitter (University of Maryland, College Park) for his encouragement and financial support for my museum visit.
REFERENCES Adamski D, Boege K, Landry J-F, Sohn J-C (2009) Two new species of Wockia Heinemann (Lepidoptera: Urodidae) from coastal dry-forests in Western Mexico. Proc Entomol Soc Wash 111: 166–182 Agnarsson I, Miller JA (2008) Is ACCTRAN better than DELTRAN? Cladistics 24: 1–7 Bruand T (1851) Catalogue systématique et synonymique des Lépidoptères du Département du Doubs. Tineides. Mém Soc emul Doubs 3(1): 23–58 Carpenter JM (1992) Random cladistics. Cladistics 8: 147–153 Cho S, Zwick A, Regier J, Mitter C, Cummings M, Yao J, et al. (2011) Can deliberately incomplete gene sample augmentation improve a phylogeny estimate for the advanced moths and butterflies (Hexapoda: Lepidoptera)? Syst Biol 60: 782–796 Clarke JFG (1941) The preparation of slides of the genitalia of Lepidoptera. Bull Brooklyn Entomol Soc 36: 149–161 Collinson ME (1992) The early fossil history of Salicaceae: a brief review. Proc R Soc Edinb 98B: 155–167 Davis JI (1995) A phylogenetic structure for the monocotyledons, as inferred from chloroplast DNA restriction site variation, and a comparison of measure of clade support. Syst Bot 20: 503–527 DeBry RW (2001) Improving interpretation of the decay index for DNA sequence data. Syst Biol 50: 742–752 Eberle JJ, Greenwood DR (2011) Life at the top of the greenhouse Eocene world, A review of the Eocene flora and vertebrate fauna from Canada’s High Arctic. Geol Soc Am Bull 124: 3–23 Goloboff P (1999) NONA. Version 2.0. Published by the author, San Miguel de Tucumán Grimaldi DA, Engel MS (2005) Evolution of the Insects. Cambridge University Press, Cambridge Heinemann H (1870) Die Schmetterlinge Deutschlands und der Schwiez, systematisch Bearbeitet. Vieweg, Braunschweig Heppner JB (1997) Wockia asperipunctella in North America (Lepidoptera: Urodidae: Galacticinae). Holarc Lepid 4: 73–74
Heppner JB (1998) Classification of Lepidoptera, Part 1. Introduction. Holarc Lepid 5(Suppl. 1): 1–148 Heppner JB (2008) The genus Wockia and a new species from southern Brazil (Lepidoptera: Urodidae: Galacticinae). Lepid Nov 1: 116–118 Heppner JB (2010a) Incawockia, a new genus and species from Peru (Lepidoptera: Urodidae: Galacticinae). Lepid Nov 3: 154– 158 Heppner JB (2010b) Notes on Vietnam moths, 10. Notes on Wockia in Vietnam (Lepidoptera: Urodidae: Galacticinae). Lepid Nov 3: 165–168 Herrich-Schäffer AW (1854) Sammlung neuer oder wenig bekannter aussereuropäischer Schmetterlinge. Vol. 1. G. J. Manz, Regenburg Hodges RW (2005) Order Lepidoptera. In “Borror and DeLong’s Introduction to the Study of Insects 7th edition” Ed by CA Triplehorn, NF Johnson, Thomson Books/Cole, Belmont, pp 571–647 Holman JA (2000) Fossil Snakes of North America: Origin, Evolution, Distribution, Paleoecology. Indiana University Press, Bloomington Klots AB (1970) Lepidoptera. In “Taxonomist’s Glossary of Genitalia in Insects” Ed by SL Tuxen, Munksgaard, Copenhagen, pp 115–130 Kodandaramaiah U (2010) Use of dispersal–vicariance analysis in biogeography – a critique. J Biogeogr 37: 3–11 Kyrki J (1984) The Yponomeutoidea: a reassessment of the superfamily and its suprageneric groups (Lepidoptera). Entomol Scand 15: 71–84 Kyrki J (1986) Wockia balikpapanella sp. n. from Borneo (Lepidoptera, Yponomeutidae auct.). Ann Entomol Fenn 52: 42–43 Kyrki J (1988) The systematic position of Wockia Heinemann, 1870, and related genera (Lepidoptera: Ditrysia: Yponomeutidae auct.). Nota Lepid 11: 45–69 Landry JF (1998) Additional Nearctic records of Wockia asperipunctella, with notes on its distribution and structural variation (Lepidoptera: Urodidae). Holarc Lepid 5: 9–13 Leskinen E, Alström-Rapaport C (1999) Molecular phylogeny of Salicaceae and closely related Flacourtiaceae: evidence from 5.8 S, ITS 1 and ITS 2 of the rDNA. Pl Syst Evol 215: 209–227 Meyrick E (1913) Hyponomeutidae. In “Exotic Microlepidoptera. Vol. 1” Ed by E Meyrick, E. W. Classey Limited, Middlesex, pp 65– 176 Missiaen P, Gunnell GF, Gingerich PD (2011) New Brontotheriidae (Mammalia, Perissodactyla) from the Early and Middle Eocene of Pakistan with implications for mammalian paleobiogeography. J Paleo 85: 665–677 Mutanen M, Wahlberg N, Kaila L (2010) Comprehensive gene and taxon coverage elucidates radiation patterns in moths and butterflies. Proc R Soc B 277: 2839–2848 Nixon KC (2002) WINCLADA. Version 1.0. Published by the author, New York Pascual R, Jaureguizar EO (1992) Evolutionary pattern of land mammal faunas during the Late Cretaceous and Paleocene in South America: a comparison with the North American pattern. Ann Zool Fenn 28: 245–252 Regier JC, Mitter C, Zwick A, Bazinet AL, Cummings MP, Kawahara AY, et al. (2013) A large-scale, higher-level, molecular phylogenetic study of the insect order Lepidoptera (moths and butterflies). PLoS ONE 8(3): e58568 Ronquist F (1996) DIVA version 1.1. Computer program and manual available by anonymous FTP from Uppsala University (ftp.systbot.uu.se) Sohn J-C (2012) Taxonomic review of Spiladarcha Meyrick (Lepidoptera: Urodoidea: Urodidae) with a new species from Venezuela. Entomol Sci 15: 303–308
Phylogeny and biogeography of Wockia Sohn J-C (2013a) A new genus and species of Urodidae (Lepidoptera: Urodoidea) from Argentina with the first report of asymmetric male genitalia in the superfamily. Fla Entomol 96: 496–476 Sohn J-C (2013b) Two new species of Wockia Heinemann (Lepidoptera: Urodidae) from Jamaica. Neotrop Entomol 42: 300–303 Sohn J-C, Adamski D (2008) A new species of Wockia Heinemann, 1890 [sic] (Lepidoptera: Urodidae) from Korea. Proc Entomol Soc Wash 111: 556–561 Sohn J-C, Park K-T (2009) A new species of Wockia Heinemann, 1870 (Lepidoptera: Urodidae) from Vietnam. Trop Lepid Res 19: 62–63 Sohn J-C, Labandeira C, Davis D, Mitter C (2012) An annotated catalog of fossil and subfossil Lepidoptera (Insecta: Holometabola) of the world. Zootaxa 3286: 1–132 Sohn J-C, Regier JC, Mitter C, Davis D, Landry J-F, Zwick A, et al. (2013) A molecular phylogeny for Yponomeutoidea (Insecta, Lepidoptera, Ditrysia) and its implications for classification, bio-
265
geography and the evolution of host plant use. PLoS ONE 8(1): e55066 Sokal RR, Rohlf FJ (1981) Taxonomic congruence in the Leptopodomorpha re-examined. Syst Zool 30: 309–325 Tiffney BH (1985) The Eocene North Atlantic Land Bridge: its importance in Tertiary and modern phytogeography of the northern hemisphere. J Arnold Arboretum 66: 243–273 Walsingham L (1914) Anchimacheta. In “Biologia Centrali Americana. Zoology: Insecta. Lepidoptera-Heterocera. Vol. 4. Tineina, Pterophorina, Orneodina, Pyralidina, and Hepialina” Ed by FD Godman, O Salvin, Taylor & Francis, London, pp 323–324 Wenzel JW (2002) Phylogenetic analysis: the basic method. In “Techniques in Molecular Systematics and Evolution” Ed by R Desalle, G Giribet, W Wheeler, Birkhäuser Verlag, Basel, pp 4– 30 Wootton RJ (1979) Function, homology and terminology in insect wings. Syst Entomol 4: 81–93 (Received September 23, 2013 / Accepted December 27, 2013)
