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PUBLISHED BY THE AMERICAN MUSEUM OF NATURAL HISTORY CENTRAL PARK WEST AT 79TH STREET, NEW YORK, NY 10024 Number 3295, 25 pp., 61 figures, 3 tables April 25, 2000

A Cladistic Analysis of the Therini: A New Synonym of the (: Geometridae, )

SEI-WOONG CHOI1

ABSTRACT To test the monophyly of the tribe Therini and resolve the relationship between Therini and Cidariini, 49 morphological characters of 38 species of and related genera were analyzed cladistically. Five taxa from the Cidariini were chosen for outgroup comparisons. Monophyly of the Therini was not supported and the Therini newly synonymized with Cidariini. Five genera, Praethera Viidalepp, Thera Stephens, Viidalepp, Inoue, and Diathera Choi, are redefined monophyletically, and two new monotypic genera are proposed: Costicoma, n. gen. and Fascilunaria, n. gen. The combination Thera firmata (Hu¬bner) is proposed. Relationships among the ingroup taxa are largely resolved: (Fascilunaria (Heter- othera (Thera (Diathera, Pennithera)))). Synapomorphies and a diagnosis of each are given, and a key to the ingroup genera is provided.

INTRODUCTION way, 1997). Concurrently, Prout (1914) placed Thera as a subgenus within , The geometrid subfamily Larentiinae is sensu lato, and subdivided the genus into two distributed worldwide and approximately 39 subgenera separated by the structure of the genera are shared among the Palearctic and male antennae (shortly ciliated vs. bipectin- Nearctic regions. Therini is a tribe of the Lar- ate). Later, Prout (1941) upgraded Thera to entiinae and the type genus Thera Stephens the generic level. occurs in the Holarctic region. Pierce (1914) Since the works of Pierce and Prout, sev- placed Thera as a distinct group, Therinae, eral new genera and species within the Ther- which is now referred to as a tribe (Hollo- ini have been described, mainly from the

1 Kalbfleisch Fellow, Division of Invertebrate Zoology, American Museum of Natural History.

Copyright American Museum of Natural History 2000 ISSN 0003-0082 / Price $3.20 2 AMERICAN MUSEUM NOVITATES NO. 3295

Fig. 1. Viidalepp’s (1980) generic relationships of Thera and related genera. A. Viidalepp’s phylo- gram, redrawn from his fig. 1; B. the strict consensus cladogram of 55 cladograms from the reanalysis, see appendix 2. Abbreviations: A Asaphodes, H Heterothera, T Thera.

Oriental region. Inoue (1943) described the described additional members of the Cidari- monotypic genus Heterothera, and Viidalepp ini (Paradysstroma Choi, 1998 and Pseudo- (1980) subdivided Therini (sensu Prout) into Heydemann, 1961) and Therini five genera: Asaphodes Meyrick sensu Viida- (Heterothera and Diathera) (Choi, 1998a, lepp, Pennithera Viidalepp, Heterothera In- 1998b, 1999) and confused our understand- oue, Praethera Viidalepp, and Thera, al- ing of the phylogeny of these . Many though the first one (Asaphodes sensu Viida- synapomorphies for Pennithera, Heteroth- lepp) was replaced by Viidaleppia by Inoue era, and Thera were found to be homoplas- (1982). Later, Choi (1997) synonymized Vi- tic, also occurring in Paradysstroma and idaleppia with Heterothera and further pro- Pseudodysstroma. For example, the presence posed a genus from southwestern China, of cucullus hairs and saccular processes in Diathera (Choi, 1999). Presently the Therini the male genitalia were synapomorphic for includes 5 genera and about 50 species Heterothera, but these occur in Pseudodys- worldwide: Thera Stephens (1831), Praeth- stroma and Diathera as well. Similarly the era Viidalepp (1980), Pennithera Viidalepp corona-shaped processes at the distal part of (1980), Heterothera Inoue (1943), and the aedeagus, an autapomorphic character for Diathera Choi (1999). Since most species of Thera, is also observed in several species of Asaphodes occur in Australia and New Zea- Heterothera (see Choi, 1998a). Such homo- land and are closely related to Xanthorhoe plasy prompted the present analysis. Hu¬bner (Dugdale, 1988), this genus is ex- The second focal point is to understand the cluded from the Therini. generic placement of Thera exangulata Here I present a cladistic analysis of the (Warren) and T. cyphoschema Prout. These Therini based on adult morphology. The taxa were included in the Therini by Prout analysis will focus on two things. First, a test (1938, 1941). Although I have described the of the monophyly of the Therini and an ex- genitalia and other aspects of morphology of amination of relationships between Therini these species (Choi, 1998a), the placement of and Cidariini. Since my previous cladistic these taxa remains uncertain. analysis (Choi, 1997), several papers have Members of the Therini occur mainly in 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 3

Asia, except for Thera, which is Holarctic in truncatum (Hufnagel), Paradysstroma corus- distribution. Many of the Asian species are sarium (Oberthu¬r), and Pseudodysstroma al- endemic to southwestern China, Taiwan, or bovenosatum (Heydemann). The choice of northern India: four species of Pennithera these outgroups was based on (1) a close re- and two species of Heterothera are endemic lationship as shown by a previous cladistic to Taiwan; six species of Heterothera, and analysis (C. fulvata, P. rubiginata, and D. all species of Diathera are endemic to south- truncatum; Choi, 1997) and (2) the fact that western China; and eight species of Heter- they share apomorphic characters with in- othera are endemic to northern India. Thus, group taxa for which ingroup relationships understanding of interrelationships of Thera are ambiguous (Paradysstroma corussarium and other Asian genera will provide the basis and Pseudodysstroma albovenosatum). for future biogeographic studies. A parsimony-based computer program, NONA (Goloboff, 1993; Ver. 1.5; hold*hold/ MATERIALS AND METHODS 30mult*15), was used to find the most par- simonious trees. Multistate characters were TAXA AND CHARACTERS treated as unordered during analyses. The ingroup comprised 38 species from the Holarctic and northern Oriental regions: ACKNOWLEDGMENTS 7 species of Thera,1Praethera,8Pennith- My sincere thanks to Jaan Viidalepp for era,17Heterothera, 3 Diathera, as well as providing material and translating his work ‘‘Thera cyphoschema’’ and ‘‘T. exangulata’’ (1980) into English and to David Grimaldi, (see appendix 1). In total, 49 characters from Jim Miller, Fred Rindge, Toby Schuh the genitalia and other morphological struc- (AMNH), Kauri Mikkola, and Malcolm Sco- tures were used. ble for commenting on several versions of Adult specimens were obtained from the this manuscript and to Katsumi Yazaki (To- following museums and private collections: kyo), Kim Goodger (BMNH), Axel Haus- American Museum of Natural History, New mann (ZSM), and Dieter Stu¬ning (ZFMK) York; Bulgarian Natural History Museum, for loaning material. Angela Klaus (AMNH) Sofia; Hungarian Natural History Museum, provided helpful assistance with scanning Budapest; Institute of Botany and Zoology, electron microscopy. I am very grateful to Tartu; private collection of Katsumi Yazaki, David Grimaldi, Michael Engel, Michael Tokyo; The Natural History Museum, Lon- Pogue, and Jim Troubridge for reading the don; Zoologisches Forschungsinstitut und manuscript and offering appreciated criti- Museum Alexander Koenig, Bonn; Zoologi- cism. cal Museum, Helsinki; and Zoologische Staatssammlung Mu¬nchen, Munich. Male RESULTS AND DISCUSSION and female genitalia were prepared by soak- ing the abdomen in cold 10% KOH for ap- CHARACTER ANALYSIS proximately 24 hours. Scales and tissues Of the 49 characters, numbers 0Ð11 were were removed, stained with Chlorazol Black, from the head, thorax, and wing patterns; and mounted on slides in Euparal. Electron numbers 12Ð35 were from the male genitalia; micrographs were taken with a Hitachi and numbers 36Ð48 from the female genita- S4700 Field Emission Scanning Electron Mi- lia. croscope (FE-SEM). Morphological terms follow Forbes (1948) and Scoble (1992). HEAD AND BODY (figs. 2Ð23) 0. Antennae of male: (0) filiform; (1) pec- CLADISTIC ANALYSIS tinate. It is usual for species of moths in the The data matrix is provided in table 1. Larentiinae to be diagnosed on the basis of Five outgroups were used to root clado- bipectinate male antennae. Prout (1914) sug- grams, all belonging to the tribe Cidariini: gested that the groupings based on the anten- (Forster), rubigin- nal morphology of males or of both sexes do ata (Denis and Schiffermu¬ller), Dysstroma not always form natural taxa. However, even 4 AMERICAN MUSEUM NOVITATES NO. 3295

he classified Thera using this character, and A study with the scanning electron micro- his groupings were subsequently found to be scope shows that there are four modifications polyphyletic (Viidalepp, 1980; Choi, 1997). of the male antennae; filiform with either Thera and Praethera have filiform antennae short cilia (fig. 19) or long cilia (fig. 18), and in both sexes, but Pennithera and Heteroth- pectinate with either short pectinations (fig. era have bipectinate male antennae and fili- 21) or long pectinations (figs. 20, 22, 23). It form female antennae. was noted that the pectinations of some No- 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 5

Figs. 2Ð7. Adults. 2. Praethera praefecta, 3. Heterothera postalbida, 4. H. quadrifulta, 5. H. den- tifasciata, 6. H. incerta, 7. H. serraria. todontidae moths are shorter on one side figs. 20, 21, 23) or alternate (Pennithera (Miller, 1991). The difference in length is ob- comis; fig. 22). Different lengths of pectina- served in an ingroup species, Heterothera fir- tions or cilia are common among the taxa mata (fig. 20). Viidalepp (1980) and Inoue examined and are difficult to code. Thus, I (1986a) illustrated the differences in pecti- used two simple states, filiform versus pec- nation between these genera. The present re- tinate. sults show that pectinations are quadripectin- 1. Vertex: (0) black; (1) distinct with ate, but differ in orientation, opposite (Het- white scales. The vertex can be covered with erothera firmata, H. taigana, H. serraria; either black or white scales. Several species 6 AMERICAN MUSEUM NOVITATES NO. 3295

Figs. 8Ð13. Adults (continued). 8. , 9. T. vetustata, 10. T. firmata, 11. Diathera fluc- tuata, 12. Pennithera comis, 13. P. lugubris. of Heterothera and Pennithera have white Heterothera, and that this is an apomorphic scales on the vertex [e.g., Heterothera ecli- character of Cidariini. nosis Choi, H. yunannensis Choi, H. hoenei 3. Foretarsal joints: (0) indistinct in that Choi, Pennithera abolla (Inoue), P. lugubris scales covering them are same color as Inoue, P. subalpina Inoue]. scales on rest of leg; (1) distinct with white 2. Frons: (0) covered with uniformly col- scales. The tarsal joints of forelegs are usu- ored scales; (1) mixed white or ochreous and ally distinct with white scales in Thera, blackish scales. Most of the ingroup taxa Praethera, and Diathera, but are indistinct in have the frons covered with white/ ochreous several species of Pennithera [comis (Butler) or blackish scales. Choi (1997) noted also and abolla (Inoue)] and Heterothera [serrar- that a rounded frons occurs in Thera and ia (Lienig), serrataria (Prout) and kurenzovi 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 7

Figs. 14Ð17. Adults (continued). 14. P. distractata, 15. P. djakonovi, 16. Thera exangulata, 17.T. cyphoschema.

Choi, Viidalepp and Vasjurin]. Obscure tarsal medial lines: (0) unicolorous; (1) tinged joints are also observed in the outgroup taxa, with black scales. except in Cidaria fulvata (which has white 8. Basal part of forewing dorsally with scales on the joints). a black dot: (0) absent; (1) present, but dot 4. Meso- and metathorax: (0) middor- small; (2) dot large. The pattern in the area sally white; (1) black. between the basal and antemedial lines of the 5. Metathorax: (0) with white or yellow forewing varies in Therini. Pennithera is dis- tufts; (1) with blackish tufts. Strong tufts oc- tinct by having a black dot on the dorsum. cur on the dorsum of the metathorax, and this Some species of Heterothera [e.g., quadri- is often observed in some other taxa of the fulta (Prout), taigana (Djakonov), incerta In- Larentiinae (Prout, 1914). It was noted that oue, and consimilis (Warren)] also have a strong tufts occur on the metathorax in mem- small dot at the dorsum, but they differ from bers of the Hydriomenini (Forbes, 1948) and Pennithera in having a black band in the area Cidariini (Choi, 1997). between basal and antemedial lines (figs. 3, 4, 12, 13). WINGS (figs. 2Ð17) 9. Discal dot of forewing united with 6. Shape of antemedial line of forewing, costal part of antemedial line: (0) absent; from costa to dorsum: (0) transverse, (1) present. Heterothera firmata (Hu¬bner), straight; (1) rounded or medially indented; H. etes (Prout), and Thera cyphoschema have (2) outwardly oblique. a long, thick discal dot united with the costal 7. Forewing between basal and ante- part of the antemedial line (fig. 10). 8 AMERICAN MUSEUM NOVITATES NO. 3295

Figs. 18Ð23. Scanning electron micrographs of male antennae, ventral surface. 18. Thera variata (ϫ110), 19. Praethera praefecta (ϫ150), 20. H. firmata (ϫ150), 21. Heterothera taigana (ϫ130), 22. Pennithera comis (ϫ110), 23. H. serraria (ϫ150).

10. Dorsum of central fascia: (0) indis- nithera and Heterothera: (1) a large round tinct and uniform; (1) distinct with black dot between the basal and antemedial line scales. (e.g., Pennithera lugubris and P. subcomis) 11. Dorsum of forewing: (0) without a (figs. 12Ð13), (2) a long horizontal streak streak; (1) with a black horizontal streak. along the basal and central fascia (e.g., Het- There are three patterns of black dots in Pen- erothera postalbida (Wileman), H. sororcula 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 9

(Bastelberger), H. tephroptilus (Heyde- juxta that supports the ventral part of the ae- mann), H. mussooriensis Choi) (fig. 3), and deagus and are usually distinct in most (3) a blackish spot at the bottom of the cen- groups of Cidariini (Choi, 1997). In Therini, tral fascia (e.g., H. triangulata Choi) (Choi, the anellus lobes are distinctive in their 1998a). length, sclerotization, and for the long apical hairs. The different shapes of the body of the MALE GENITALIA lobes (e.g., triangular, cylindrical, or caplike) provide diagnostic features for the species of 12. Length of uncus compared to tegu- Heterothera, Diathera, and Pennithera (figs. men (figs. 24Ð28): (0) shorter; (1) longer. 32, 35, 36): Pennithera possesses anellus Three species of Diathera show a very long lobes with a strongly expanded body; and uncus compared to the length of tegumen Thera possesses digitiform lobes. But, the and this long uncus is also observed in most shapes of the lobes in Heterothera vary. In species of Pennithera and Heterothera. particular, the species from southwestern 13. Length of tegumen compared to the China have greatly modified anellus lobes total length of vinculum and saccus: (0) that are bilobed with an expanded body, of- same; (1) longer; (2) shorter. Tegumen ten with dentate surfaces: Heterothera yun- length is measured against the total length of nanensis, H. eclinosis, and Pennithera dis- the vinculum and saccus. This character di- tractata (figs. 31, 33, 34). vides Therini into three groupings: (Praeth- 22. Transtilla: (0) simple, membraneous; era), (Pennithera and Thera), and (Heteroth- (1) platelike, sclerotized; (2) thin, round, era and Diathera). Praethera has a long teg- sclerotized. The transtilla, the dorsal part of umen, whereas Thera and Pennithera have a the diaphragma, is usually simple and mem- short tegumen. Diathera and Heterothera braneous in most ingroup taxa, whereas The- have nearly the same length between the teg- ra shows a well-developed and sclerotized umen and the total length of the vinculum transtilla. Thera exangulata has a thin, round, and saccus. sclerotized transtilla. 14. Width of anterior end of tegumen 23. Shape of saccus (figs. 24Ð28): (0) flat (vinculum): or round and medially slightly projected; (1 ؍) compared to posterior end (0) same or slightly shorter; (1) greatly medially invaginated; (2) medially strongly shorter. projected. The shape of saccus shows three 15. Shape of anterior end of tegumen states: (1) broad, flat, or round saccus (figs. 24Ð28): (0) slightly invaginated; (1) (Praethera and Pennithera)(fig. 24), (2) me- deeply invaginated. dially invaginated or broad saccus (Heteroth- 16. Juxta with strongly sclerotized hairs era and Diathera)(figs. 26, 28), and (3) me- (fig. 31): (0) absent; (1) present. A unique dially strongly projected (Thera)(fig. 25). In character of the juxta is observed in all spe- Thera and Pennithera the saccus shows a cies of Diathera, which is a structure covered sclerotized internal ring. with strongly sclerotized hairs. 24. Costa of valva (figs. 37Ð44): (0) mem- 17. Length of anellus lobe: (0) strongly braneous; (1) sclerotized. extended from juxta, reaching more than half 25. Shape of costa: (0) flat, not expanded; of uncus height; (1) slightly extended from (1) basally expanded (fig. 40); (2) medially juxta, barely reaching bottom of uncus. expanded (figs. 38, 41); (3) distally expand- 18. Shape of anellus lobes (figs. 29Ð36): ed. (0) digitiform, without expanded body; (1) 26. Costa with a distal sclerotized pro- body expanded. cess: (0) absent; (1) present. The costa, 19. Anellus lobe: (0) unilobed; (1) bi- which is the dorsal part of the valva, is fre- lobed. quently modified and strongly sclerotized in 20. Surface of anellus lobe: (0) smooth; Therini and this has been most frequently (1) dentate. used by earlier authors (e.g., Viidalepp, 21. Shape of anellus lobe: (0) digitiform; 1980; Inoue, 1986a; Choi, 1997). In Thera, (1) rounded; (2) triangular; (3) flattened. Pennithera, and Praethera, the costa has im- The anellus lobes are lateral processes of the portant diagnostic characters: Thera with a 10 AMERICAN MUSEUM NOVITATES NO. 3295

Figs. 24Ð28. Complex of tegumen, vinculum, and saccus of the male genitalia. 24. Praethera prae- fecta, 25. Thera variata, 26. Diathera fluctuata, 27. Pennithera comis, 28. Heterothera postalbida. Un uncus, Tg tegumen, Vn vinculum, Sc Saccus. slender costa with a medial triangular pro- greatly enlarged; in Praethera it is long, jection; in Pennithera it is basally greatly en- slender, and distally sharply pointed. How- larged and distally sharply pointed; in Diath- ever, in Heterothera the shape of the costa era the base is minutely toothed and medially varies: with a medial, triangular process sim- 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 11

Figs. 29Ð36. Anellus lobes of male genitalia. 29. Praethera praefecta, 30. Thera variata, 31. Diath- era fluctuata, 32. Pennithera comis, 33. P. distractata, 34. Heterothera yunnanensis, 35. H. postalbida, 36. H. serraria. Jx juxta, Al anellus lobe. ilar to Thera (fig. 41), or basally or distally ally coupled with projected processes on the expanded (fig. 42). Choi (1997) noted that distal part of sacculus, which are often called the absence of the distal costal process is ple- the ‘‘harpe’’ (Inoue, 1986a). In Heterothera, siomorphic. the process varies from simple and setose 27. Ventral edge of sacculus: (0) invag- (e.g., tephroptilus) to clawlike [e.g., dentifas- inated (figs. 37Ð41, 44); (1) flat (figs. 42, ciata (Hampson)] and stellate (hoenei). Four 43). species of Heterothera have two processes 28. Shape of dorsal edge of sacculus: (0) [postalbida, sororcula, undulata (Warren) indistinct; (1) vertical (fig. 38); (2) straight and obscurata Choi; Choi, 1998a] (fig. 42). and oblique (figs. 41, 42); (3) scalloped. This character was also used in Viidalepp’s 29. Process of sacculus: (0) absent; (1) (1980) analysis (his #2). clawlike or sharply pointed (figs. 38, 41, 42, 31. Width of valva: (0) wider distally 44); (2) stellate. (figs. 39, 42); (1) even width (figs. 41, 43); 30. Number of saccular processes: (0) (2) wider basally (fig. 40). The width of the absent; (1) one; (2) two. The sacculus divides valva diagnoses some members of Therini. Therini into two groups based on whether it In Praethera, Diathera, and Thera, the distal is sclerotized or membraneous. The former part of valva is wider, whereas in Pennithera includes Thera and Heterothera, the latter the basal part is wider. In Heterothera, the Praethera, Diathera and Pennithera. The tri- valva is usually wider distally, except for angular and sclerotized sacculus diagnoses three species (serraria, serrataria and kuren- Heterothera. The sclerotized sacculus is usu- zovi) that have a basally wider valva. 12 AMERICAN MUSEUM NOVITATES NO. 3295

Figs. 37Ð44. Right valva of male genitalia. 37. Praethera praefecta, 38. Thera variata, 39. Diathera fluctuata, 40. Pennithera comis, 41. , 42. Heterothera postalbida, 43. Thera ex- angulata, 44. Thera cyphoschema. C costa, S sacculus, p saccular process.

32. Distal part of the aedeagus: (0) mem- ination of the distal part of the aedeagus is braneous, without spines; (1) scobinate (figs. morphologically part of the juxta. Because 47, 49, 50); (2) with large spines (fig. 46). these two parts are fused, they are usually Therini has several derived character states broken in genitalic preparations. on the distal part of the aedeagus. The distal 33. Shape of the vesica: (0) large, saclike spines in Thera have often been termed cor- (fig. 51); (1) tubular (fig. 45) nuti (e.g., Viidalepp, 1980; Choi, 1997), and 34. Cornutus: (0) absent; (1) large, spi- this state may be a symplesiomorphy of The- nular; (2) small setae; (3) mixed large and ra and Heterothera (Choi, 1998a). The scob- small spines. 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 13

35. Cornuti: (0) absent; (1) grouped into postalbida, sororcula, and hoenei) to large one patch; (2) grouped into several patches; hornlike processes (e.g., taigana). In Thera (3) without grouping, scattered. Viidalepp there is no distinction between the lamella (1980) divided the states of the cornuti into antevaginalis and the lamella postvaginalis, primary and secondary losses. For example, because the two structures are fused and Pennithera djakonovi (Kurentzov) is coded semicircular (fig. 53). as a primary loss because it lacks cornuti in 41. Length of ductus bursae compared the vesica. But most species of Thera are to corpus bursae: (0) shorter; (1) longer. coded as secondary because they lack cornuti 42. Ductus bursae with a colliculum: (0) in the vesica while having spinal processes present; (1) absent. at the distal part of the aedeagus. Viidalepp 43. Ductus bursae: (0) membraneous; (1) also suggested that the presence of cornuti is sclerotized; (2) with sclerites. The shape of a derived character in Therini. the ductus bursae of Heterothera is variable. The shape of the vesica is coded as tubular Unlike outgroup taxa, in which it is short and or large and saclike. For most ingroup taxa membraneous with a colliculum, Heterothera the vesica is tubular, whereas some outgroup and Pennithera have a sclerotized wall or taxa (Dysstroma truncatum, Plemyria rubi- sclerites in the ductus bursae. The shape of ginata) and Thera exangulata have a large, the ductus bursae in these taxa may be long, saclike vesica. Choi (1997) noted that several twisted, or basally greatly expanded. Thera transformations occur in the vesica of Het- and Diathera have a membraneous ductus erothera: several irregularly shaped diverti- bursae, but lack a colliculum. culae and many kinds of spinal processes and 44. Basal ductus bursae (anterior to an- arrangements of the cornuti. trum): (0) simple, without modification; (1) funnel-shaped; (2) rounded and sclerotized. FEMALE GENITALIA The basal part of the ductus bursae (anterior to the antrum) is usually simple and mem- 36. Seventh sternite: (0) simple, membra- braneous. Two species ( neous; (1) with sclerotized processes; (2) and Thera cyphoschema) have a ductus bur- scobinate. A densely scobinate seventh ster- sae basally funnel shaped (fig. 58), but in nite diagnoses Diathera, while sclerotized several species of Heterothera (serraria, ser- processes diagnose Pennithera.InThera, rataria and kurenzovi) it is strongly sclero- Praethera, and Heterothera, like outgroup tized with a rounded opening. taxa, the seventh sternite is simple and mem- 45. Distal ductus bursae: (0) membra- braneous. neous; (1) flattened and sclerotized. 37. Lamella antevaginalis (figs. 52Ð59): 46. Posterior corpus bursae: (0) simple, (0) membraneous, without structures; (1) membraneous; (1) with sclerotized striations; sclerotized with rounded structure; (2) scler- (2) simple, sclerotized. The area between the otized with two doughnutlike structures. ductus bursae and the corpus bursae is usu- 38. Lamella antevaginalis with process- ally simple and membraneous. But, several es: (0) absent; (1) minute spines; (2) large species of Heterothera (serraria, serrataria, spines. and kurenzovi) have the corpus bursae 39. Lamella postvaginalis: (0) membra- strongly sclerotized. neous; (1) sclerotized. 47. Signum: (0) one-process; (1) several, 40. Lamella postvaginalis with process- placed in a patchlike band; (2) absent; (3) es: (0) absent; (1) large, barlike processes; large, threadlike sclerite. Viidalepp (1980) (2) large, hornlike processes. Sterigmata (la- indicated that the absence of a signum is a mellae ante- and postvaginalis) occur around derived character of the Therini. In Thera, the ostium bursae and they are modified into Pennithera, Heterothera, and Diathera,a various forms. Viidalepp (1980) noted that signum is absent, whereas in Praethera,a the well-developed sterigmata are plesiomor- minute nipple-shaped signum occurs (fig. phic in this group. In Heterothera, these 52). One species (Thera exangulata) shows structures vary from simple (e.g., consimilis, a large, threadlike signum (fig. 59). etes, and dentifasciata) to spinelike (e.g., 48. Wall of the corpus bursae: (0) sim- 14 AMERICAN MUSEUM NOVITATES NO. 3295

Figs. 45Ð51. Aedeagus with everted vesica of male genitalia. 45. Praethera praefecta, 46. Thera variata, 47. Pennithera comis, 48. Thera cyphoschema, 49. Diathera fluctuata, 50. Heterothera quad- rifulta, 51. Thera exangulata. Vs vesica, Ct cornuti, D diverticulum. ple, membraneous; (1) with wavelike stria- renzovi) to minutely scobinate (e.g., incerta, tions; (2) with minute scobinations; (3) net- yunnanensis, and eclinosis)(figs. 55, 56). In like and sclerotized. In Heterothera, the Pennithera, the wall is usually membra- wall of the corpus bursae varies from simple neous and simple (fig. 57), but in Thera, (e.g., tephroptilus, sororcula, and quadri- Diathera, and Praethera wavelike striations fulta) to having wavelike striations (e.g., ku- are observed (figs. 52Ð54). Exceptionally, 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 15

Figs. 52Ð56. Sterigmata, ductus, and corpus bursae of female genitalia. 52. Praethera praefecta, 53. Thera variata, 54. Diathera fluctuata, 55. Heterothera quadrifulta, 56. H. serraria. La lamella anteva- ginalis, Sg signum, Db ductus bursae, Cb corpus bursae.

Thera exangulata has a netlike, sclerotized gram is shown in figure 60. The successive wall (fig. 59). weighting approach (Farris, 1969) for these 45 cladograms was applied by NONA and CLADISTIC ANALYSIS after two iterations one cladogram of weight- NONA produced 45 cladograms of length ed length 7819 was stabilized (table 2, fig. 256, consistency index ϭ 0.30, and retention 61). The cladogram derived from the succes- index ϭ 0.66. The strict consensus clado- sive weighting provides better resolved ge- 16 AMERICAN MUSEUM NOVITATES NO. 3295

Figs. 57Ð59. Sterigmata, ductus, and corpus bursae of female genitalia (continued). 57. Pennithera comis, 58. Thera cyphoschema, 59. Thera exangulata. neric interrelationships than the original clad- su Prout, Viidalepp (1980) divided this group ograms: (Thera cyphoschema, (Heterothera, into three genera and four subgenera (fig. 1), (Thera,(Diathera, Pennithera)))). suggesting that Prout’s classification was un- natural. A cladistic analysis of the Cidariini THERINI, A SYNONYM OF CIDARIINI resulted in some relationships different from Viidalepp’s hypothesis (Choi, 1997)—specif- Both present analyses show that the tribe ically that Heterothera was paraphyletic. Therini is not monophyletic and both (figs. Choi also indicated that the basal node of 60, 61) excluded Praethera from the ingroup three genera, Thera ϩ Pennithera ϩ Heter- taxa. Therefore, the results do not support the othera, was supported by several synapo- tribe Therini, and suggest that the Cidariini morphies. In the present analysis my pre- should be applied as the appropriate name for ferred cladogram (fig. 61) resolves the ge- all ingroup taxa. neric relationships without Praethera:(Fas- Both unweighted and successive weight- cilunaria, (Heterothera, (Thera,(Pennithera, ing approaches support most ingroup taxa, Diathera)))), and the basal clade is supported Praethera, Thera, Pennithera, and Diathera, by three synapomorphies: male antennae bi- and the positions of two unplaced taxa, T. pectinate (character #0-1), antemedial line of exangulata and T. cyphoschema, turned out forewing rounded or medially indented (#6- to be part of a very basal member. Differ- 1), and signum absent (#47-2). ences among the trees were a result of the The present analysis confirms four previ- placement of several taxa in Heterothera, ously known monophyletic taxa, which is Heterothera firmata, and the serraria group partly consistent with the results of Choi of Heterothera (serraria, serrataria, and ku- (1997). However, the result indicates that renzovi). In the original cladograms, H. fir- Praethera is not closely related to Thera, and mata was a basal taxon of Pennithera or two new monotypic genera, Fascilunaria and formed an independent clade with the ser- Costicoma, and one combination, Thera fir- raria group. mata (Hu¬bner), are suggested. Their formal In his analysis for 16 species of Thera sen- classification is presented in appendix 1. 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 17

Fig. 60. Strict consensus cladogram derived from unweighted analysis using NONA. Abbreviations: C Cidaria, D Dysstroma, P Plemyria, Pe Pennithera, Pd Paradysstroma, Pr Praethera, Ps Pseudo- dysstroma, H Heterothera, T Thera, Di Diathera. Plus signs (ϩ) indicate a taxon or node that produces the polytomy in the consensus cladogram; asterisks (*) indicate taxa that have ambiguous taxonomic positions. 18 AMERICAN MUSEUM NOVITATES NO. 3295

Fig. 61. One cladogram derived from successive weighting approach using NONA. See fig. 60 for generic abbreviations and symbols. Each node number is indicated. 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 19

neous, and presence of a colliculum and a signum. Viidalepp (1980) demonstrated that Praethera is the sister group of Heterothera. He listed five apomorphic characters for Praethera: saccus massive and round; sac- culus membraneous; last tergite of female abdomen quadrilateral; ductus bursae modi- fied; and both the collar of the bursa copu- latrix and sterigma well-developed and strongly sclerotized. However, the present analysis shows that Praethera is placed as the sister taxon of (Paradysstroma corussar- ium ϩ Costicoma exangulata), not Heteroth- era. Two species of Praethera are known, P. praefecta (Prout, 1914) and P. anomala (In- oue, 1954), but the validity of the latter spe- cies is uncertain (Viidalepp, 1996). Both spe- cies occur in east Asia.

Costicoma, new genus

SYSTEMATICS TYPE SPECIES: Perizoma exangulata War- Praethera Viidalepp, 1980 ren, 1909. DIAGNOSIS: Species of Costicoma can be TYPE SPECIES: Cidaria praefecta Prout, characterized by the forewing with blackish 1914. central fascia medially and dorsally greatly DIAGNOSIS: Species of Praethera are char- tapered, the valva with sclerotized costa and acterized by the filiform male antennae, the long hairs on the middle of costa, cornuti ab- subdorsally greatly shrunken central fascia sent from vesica, and the female genitalia with third cell half as wide as second, the having the corpus bursae with a large, male genitalia having a long tegumen, slen- threadlike signum. der and sclerotized costa of valva upturned DESCRIPTION: Antenna filiform in both sex- distally, membraneous sacculus with a me- es. Frons mixed with ochreous and brownish dially invaginated ventral edge and two bun- scales. Labial palp short. Legs dark brown- dles of cornuti on a tubular vesica, and the ish, foreleg tarsal joints distinct with ochre- female genitalia having V-shaped, sclerotized ous scales. Metathorax dorsum with blackish lamella postvaginalis. tufts. Forewing basally dark brown; basal DISCUSSION: In the present analysis, one line dentate, slanted; central fascia blackish, species of Praethera is supported by seven costally bulged; subterminal line whitish, autapomorphies: dorsum of the central fascia scalloped; apical streak blackish. Hindwing distinct with blackish scales (character #10- whitish, basally tinged with blackish scales; 1), anellus lobe with expanded and flattened postmedial line blackish. Male genitalia. body (#18-1, #21-3), valva width wider dis- Uncus long, basally tapering. Tegumen tally (#31-0), cornuti large, spinular (#34-1), dome-shaped. Saccus rounded, medially ex- and lamella postvaginalis with a V-shaped panded. Anellus lobe long, rod shaped, api- sclerotization (#39-1). cally with long hairs. Transtilla thin, round- Wing patterns of Praethera, especially the ed, sclerotized. Valva slender; costa slender, shape of the central fascia, place this group distally with long hairs; sacculus membra- in Therini, but several genitalic characters neous. Aedeagus cylindrical with large, sac- differ from Thera and are closer to Dysstro- like vesica; cornutus absent. Female geni- ma truncatum: long tegumen compared to talia. Papillae anales simple. Anterior apo- 3 vinculum and saccus, sacculus membra- physes about ⁄5 length of posterior apophy- 20 AMERICAN MUSEUM NOVITATES NO. 3295

ses. Sterigma membraneous, antrum broad, having long hairs on the costa, but differs in funnel-shaped. Ductus bursae short with a the wing pattern and female genitalia (Choi, colliculum. Corpus bursae large, subovate, 1998a). Results here place C. exangulata as with a long, threadlike signum. the sister taxon to P. corussarium, and the DISCUSSION: There are four autapomor- basal clade is supported by two apomorphic phies: uncus long (character #12-1); ventral characters: blackish metathorax and thin, edge of sacculus flat (#27-1); vesica large, round transtilla. This taxon was included in saclike (#33-0), signum large, threadlike the Therini (Prout, 1938), but is distinct from (#47-3), and wall of corpus bursae with net- other species of Therini based on several like sclerotizations (#48-3). apomorphic characters (Choi, 1998a): costa Costicoma exangulata is similar to Cidar- of valva strongly sclerotized with long hairs; ia deletaria Hampson in the male genitalia large saclike vesica without a cornutus; os- 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 21 tium bursae broad; colliculum present; sig- the costal part of antemedial line (character num large, threadlike, and wall of corpus #9-1), costa of the male valva with a distal bursae with netlike sclerotization. Differenc- expansion (#26-1), ductus bursae short (#41- es in wing pattern and genitalia separate ex- 0), and ductus bursae anterior to antrum fun- angulata from its sister genus Paradysstro- nel shaped (#44-1). ma, suggesting a new genus, Costicoma.Itis F. cyphoschema is similar to several mem- only known from northern India. bers of Heterothera in the shape of the cen- ETYMOLOGY: The name refers to long hairs tral fascia of the forewing as well as the anel- on the costa in the male genitalia, Costa lus lobe and saccular process of the male (Costa) ϩ Hairs (Coma). genitalia. However, other aspects of the male and female genitalia of F. cyphoschema are Fascilunaria, new genus different from those of Heterothera: uncus thick, short, and less sclerotized; cornutus TYPE SPECIES: Cidaria cyphoschema Prout, absent; sterigma simple; anterior to antrum 1926. funnel shaped; membraneous dustus bursae DIAGNOSIS: Synapomorphies for Fascilu- with a colliculum, and corpus bursae large. naria include bipectinate male antennae, the In the present analysis, the results demon- thin and crescent central fascia of the fore- strate that F. cyphoschema forms a basal lin- wing with a discal dot united with costal part eage of the clade comprising Heterothera ϩ of the antemedial line, the male genitalia Thera ϩ Pennithera ϩ Diathera (fig. 61). with a thick and short uncus, a saccular pro- This species is known from southwestern cess, cornuti absent from vesica and the fe- China, northern Vietnam and Burma (Choi, male genitalia with a simple sterigma, ante- 1998a). rior to the antrum funnel shaped, a collicu- ETYMOLOGY: The name refers to the shape lum, and the large corpus bursae without a of the central fascia of forewing, central fas- signum. cia (Fascia) ϩ crescent (lunaria). DESCRIPTION: Antenna. Male bipectinate with short pectination; female filiform. Frons white and ochreous scales. Labial palp long, Thera Stephens, 1831 about twice the eye diameter. Legs brownish, foreleg tarsal joints distinct with white TYPE SPECIES: Geometra variata [Denis scales. Metathorax dorsum with blackish and Schiffermu¬ller, 1775]. tufts. Forewing grayish; basal line blackish, DIAGNOSIS: Members of Thera have fili- slanted; antemedial line blackish, medially form male antennae; central fascia of the strongly indented; postmedial line medially forewing thin and medially and dorsally nar- projected; central fascia thin. Hindwing whit- rowed; male genitalia with hook-shaped un- ish, with a black discal dot; postmedial line cus, transtilla thin, platelike, saccus medially very weak. Male genitalia. Uncus short, projected, costa of valva strongly sclerotized thick. Tegumen dome shaped. Anellus lobe with medial and distal projections, sacculus digitiform, medially and apically with hairs. large, rounded, aedeagus slender with spi- Saccus rounded, medially projected. Valva nular processes distally; and female genitalia weakly sclerotized; costa distally projected; having lamella antevaginalis sclerotized, sacculus with a process. Aedeagus slender, rounded; ductus bursae thin, long without a rod shaped, with a tubular vesica; cornutus colliculum and the corpus bursae ovate with- absent. Female genitalia. Papillae anales out a signum. simple. Anterior apophyses half of posterior DISCUSSION: All species of Thera (except apophyses in length. Sterigma simple, dor- firmata) are united by four synapomorphies: sally with thin, sclerotized stripes; antrum male antenna filiform (character #0-0), sac- large, funnel shaped. Ductus bursae short, cus medially projected (#23-2), dorsal edge membraneous, with a colliculum. Corpus of sacculus vertically sclerotized (#28-1), bursae large, subspherical; signum absent. and lamella antevaginalis two doughnutlike DISCUSSION: Fascilunaria cyphoschema structures without a seta (#37-2). The basal has four apomorphies: discal dot united with clade comprising Thera and T. firmata is 22 AMERICAN MUSEUM NOVITATES NO. 3295 supported by one synapomorphy: transtilla nuti; female seventh sternite sclerotized; ste- large, platelike (#22-1). rigma simple; ductus bursae long, sclerotized Stephens (1831) proposed the genus Thera without a colliculum; and corpus bursae by separating it from Chesias Treitschke small and ovate without a signum. based on differences of the palpi, wing pat- DISCUSSION: Eight species of Pennithera tern and host plant (fir or juniper). Viidalepp are united by two synapomorphies: forewing (1980) defined the monophyly of Thera: sac- between basal and antemedial lines tinged cus tender and round; costa and sacculus of with black scales (#7-1) and ductus bursae valva distinct; ductus bursae membraneous; sclerotized (#43-1). Viidalepp (1980) pro- and sterigma sclerotized. Choi (1997) rede- posed the genus Pennithera and included fined the monophyly of the genus using 11 three species, P. comis (Butler), P. djakonovi synapomorphies and listed four uniquely de- (Kurentzov) and P. firmata (Hu¬bner), based rived characters: costa of valva with a medial on two apomorphies: saccus massive and projection; cucullus large; and cornuti on ve- round, and last tergite of the female abdomen sica fewer and arranged in a crown shape. telescopic. Inoue (1982, 1986a) added five About 12 species of Thera are known and species, mainly from Taiwan: P. abolla (In- they occur widely throughout the Holarctic oue), P. subcomis (Inoue), P. lugubris Inoue, region. P. manifesta Inoue, and P. subalpina Inoue. T. firmata has seven autapomorphies: ver- Choi (1998a) combined one additional spe- tex distinct with whitish scales (character #1- cies Thera distractata Sterneck (1928). 1), discal dot of forewing united to the costal Inoue (1986a, 1986b) noted that Pennith- part of antemedial line (#9-1), costa of valva era is distinguished from Viidaleppia, a ju- with a distal expansion (#26-1), cornuti on nior synonym of Heterothera, by quadripec- vesica large and spinular and grouped into tinate male antennae with slender branches, one-patch (#34-1, #35-1), and ductus bursae costa of valva half-expanded basally, greatly sclerotized (#48-1). The species T. firmata is invaginated medially and rounded distally; placed at the basal node of Thera in the suc- sacculus unsclerotized, cornuti few and cessive weighting approach (fig. 61), but as thornlike; simple and unsclerotized lamellae a member of Heterothera (fig. 60). The status ante- and postvaginalis, and slender ductus of this species was first discussed by Pierce bursae. Choi (1997) listed six synapomor- (1914). He distinguished it from Thera based phies of Pennithera: male antenna with long on structures of the genitalia. Later, Viida- pectinations; slender sacculus; female ninth lepp (1980) moved it to Pennithera, on the abdominal segment long, nearly the same basis of two apomorphies: indistinct sacculus length as eighth tergite; long and sclerotized and shortly bipectinate male antenna. How- antrum; and twisted ductus bursae. ever, Choi (1997) subsequently combined it with Heterothera, using three synapomor- Diathera Choi, 1999 phies: apical streak of forewing absent, anel- lus lobe expanded, and saccus flat and broad. TYPE SPECIES: Diathera fluctuata Choi, In the present analysis, firmata is again as- 1999. sociated with Thera. DIAGNOSIS: Members of Diathera have male antennae filiform; long uncus; juxta Pennithera Viidalepp, 1980 with short, strongly sclerotized hairs; costa of valva with minute dentate processes ba- TYPE SPECIES: Larentia comis Butler, 1879. sally, greatly expanded medially; vesica DIAGNOSIS: Members of Pennithera have without a cornutus; female eighth sternite male antennae bipectinate; forewing with a scobinate; ductus bursae short, sclerotized, black dot at dorsum between basal and an- and the corpus bursae slender with scalloped temedial line, ante and postmedial lines processes on the wall. strongly scalloped; tegumen short with nar- DISCUSSION: The three species of Diathera, row anterior end; costa of valva sclerotized all from southwestern China, are united by with large basal and distal projections; ae- five synapomorphies: male antenna filiform deagus bent; vesica with several spinular cor- (character #0-0), juxta with strongly sclero- 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 23 tized hairs (#16-1), medially invaginated sac- bursae relatively thick, and wall of ductus cus (#23-1), valva wider distally (#31-0), and bursae with sclerotized processes. female seventh sternite scobinate (#36-2). The species of this genus are similar to KEY TO GENERA EXAMINED BASED ON Pennithera externally, especially in the GENITALIA strongly scalloped ante and postmedial lines of the forewing. However, the structure of 1. Tegumen longer than total length of vinculum the male antennae and the male and female and saccus; corpus bursae with a signum Praethera (figs. 2, 21, 24, 29, 37, 45, 52) genitalia separate Diathera from Pennithera. — Tegumen shorter than total length of vinculum and saccus; corpus bursae without a signum Heterothera Inoue, 1943 ...... 2 2. Sacculus sclerotized with a process; sterigma TYPE SPECIES: Cidaria postalbida Wile- of female genitalia developed with sclerites man, 1911. ϭ Viidaleppia Inoue, 1982 [syn- ...... 3 onymized by Choi, 1997: 311]. Type species: — Sacculus without a process; sterigma simple, Cidaria quadrifulta Prout, 1938. without modification ...... 5 3. Vesica without a cornutus; ductus bursae DIAGNOSIS: Members of Heterothera are variable in the male antennae, the shape of membraneous ...... 4 — Vesica with cornuti; ductus bursae sclerotized central fascia of the forewing, and the geni- ... Heterothera (figs. 3Ð7, 22, 23, 28, 35, talia. However, they generally show the fol- 36, 41, 42, 50, 55, 56) lowing features: central fascia of the fore- 4. Uncus hooked; saccus strongly projected; ae- wing rather thick, outwardly oblique; tegu- deagus with distal spinular processes; ste- men same length as vinculum and saccus; rigma with sclerotized processes ...... saccus medially invaginated; costa of valva ... Thera (figs. 8, 18, 25, 30, 38, 46, 53) sclerotized with medial or distal expansion; — Uncus short and thick; saccus rounded; aedea- sacculus sclerotized with spinular process; gus distal part without spinular process; ste- cucullus with long, dense hairs; aedeagus rigma simple, membraneous ...... long, cylindrical; vesica with several scat- ...... Fascilunaria (figs. 17, 44, 48, 58) 5. Costa of male valva with long hairs; corpus tered, spinular cornuti; lamella ante- and bursae with a long signum ...... postvaginalis complicated with spinular or ...... Costicoma (figs. 16, 43, 51, 59) patchlike structures; ductus bursae thick with — Costa without long hairs; corpus bursae with- sclerites on the wall; and corpus bursae with- out a signum ...... 6 out a signum. 6. Costa of male valva with a large basal expan- DISCUSSION: In the present analysis, one sion; vesica with cornuti; ductus bursae synapomorphy, dorsal edge of sacculus scal- sclerotized . . Pennithera (figs. 11Ð14, 20, loped (character #28-3), supports the group- 27, 32, 33, 40, 47, 57) ing of 16 species of Heterothera. Inoue — Costa with a large distal expansion; vesica (1943) proposed the genus Heterothera without cornuti; ductus bursae membraneous .... Diathera (figs. 11, 26, 31, 39, 49, 54) based mainly on male genital characters: ab- sence of uncus (this character was found to be incorrect; see Choi, 1997); long anal tube; REFERENCES saccus basally flat and broad; costa weakly Choi, S.-W. sclerotized, and sacculus well-developed. Vi- 1997. A phylogenetic study on genera of Ci- idalepp (1980) noted that the broad, angulat- dariini from the Holarctic and the Indo- ed saccus is the main derived character for Australian areas (Lepidoptera: Geome- the genus. As noted above, Inoue (1982) tridae, Larentiinae). Syst. Entomol. 22: erected another genus, Viidaleppia. However, 287Ð312. Choi (1997) synonymized Viidaleppia under 1998a. Systematics of the genus Heterothera Inoue (Lepidoptera, Geometridae: Lar- Heterothera and listed seven synapomor- entiinae). Tijdschr. Entomol. 141: 19Ð phies: sacculus process (or harpe) small; cu- 47. cullus with hairs; saccus medially invaginat- 1998b. Taxonomic review of two genera Pseu- ed; cornuti scattered on vesica; lamella an- dodysstroma Heydemann and Para- tevaginalis with semicircular process; ductus dysstroma gen. n. (Lepidoptera: Geo- 24 AMERICAN MUSEUM NOVITATES NO. 3295

metridae, Larentiinae). Tinea 15: 228Ð ischen Faunengebietes. 12: 323Ð324. 239. Stuttgart: Verlag A. Kernen. 1999. Taxonomic review of a new genus Scoble, M. J. Diathera gen. n. from Southeast Asia 1992. The Lepidoptera: form, function, and (Lepidoptera, Geometridae: Larenti- diversity. New York: Oxford Univ. inae). J. Nat. Hist. 33: 1039Ð1048. Press. Dugdale, J. S. Stephens, J. F. 1988. Lepidoptera—Annotated catalogue, 1829Ð1831. Illustrations of British Entomol- and keys to family-group taxa. Fauna ogy. Haustellata 3. London: Baldwin N. Z. 14: 171Ð191. and Cradock. Farris, J. S. Viidalepp, J. 1969. A successive approximations approach 1980. Geometrid moths of the genus Thera to character weighting. Syst. Zool. 18: Steph. in the fauna of the USSR (Lep- 374Ð385. idoptera). Tartu Riikliku uˆlikooli To- Forbes, W.T.M. imetised 13(516): 54Ð84. [in Russian 1948. Lepidoptera of New York and neigh- with English summary] boring states. Part II. Cornell Univ. 1996. Checklist of the Geometridae (Lepi- Agric. Exp. Stn. Mem. 274: 10Ð175. doptera) of the former U.S.S.R. Sten- Goloboff, P. A. strup: Apollo Books. 111 pp. 1993. NONA, Ver. 1.5. San Miguel de Tucu- man. APPENDIX 1 Holloway, J. D. Formal Classification of Ingroup Taxa 1997. The moths of Borneo: family Geome- tridae, subfamilies Sterrhinae and Lar- Asterisks (*) indicate unexamined taxa. entiinae. Malay. Nat. J. 51: 1Ð242. Costicoma n. gen. Inoue, H. exangulata (Warren, 1909) n. comb. 1943. New and little known Geometridae Praethera Viidalepp, 1980 from Japan. Trans. Kansai Entomol. *anomala (Inoue, 1954) Soc. 12: 1Ð25. praefecta (Prout, 1914) 1982. Geometridae 1: 425Ð572. In H. Inoue et al., Moths of Japan. Tokyo: Kodan- Diathera Choi, 1999 sha. brunneata Choi, 1999 1986a. Further new and unrecorded species of fluctuata Choi, 1999 the Geometridae from Taiwan with metacolorata Choi, 1999 some synonymic notes (Lepidoptera). Pennithera Viidalepp, 1980 Bull. Fac. Domest. Sci., Otsuma Wom- abolla (Inoue, 1943) an’s Univ. 22: 211Ð267. comis (Butler, 1879) 1986b. Descriptions and records of some Jap- distractata (Sterneck, 1928) anese Geometridae (VI). Tinea 12: 45Ð djakonovi (Kurentzov, 1950) 71. lugubris Inoue, 1986 Miller, J. S. manifesta Inoue, 1986 1991. Cladistics and classification of the No- subalpina Inoue, 1986 todontidae (Lepidoptera: Noctuoidea) subcomis Inoue, 1978 based on larval and adult morphology. Bull. Am. Mus. Nat. Hist. 204: 1Ð230. Fascilunaria n. gen. Pierce, F. N. cyphoschema (Prout, 1926) n. comb. 1914. The genitalia of the group Geometridae Thera Stephens, 1829 of the Lepidoptera of the British Is- *britannica Turner, 1925 lands. Liverpool. 88 pp. ϩ 48 pls. cognata (Thunberg, 1792) Prout, L. B. contractata (Packard, 1873) 1914. Geometridae. In A. Seitz (ed.), Die cupressata (Geyer, 1831) Grossschmetterlinge der Erde—Die Pa- firmata (Hu¬bner, 1822) rev. comb. laearktischen Spanner. 1(IV): 216Ð219. juniperata (Linnaeus, 1758) Stuttgart: Verlag A. Kernen. *latens Barnes & McDunnough, 1917 1938. Geometridae. Ibid. (Suppl.): 110Ð170. obeliscata (Hu¬bner, 1787) Stuttgart: Verlag A. Kernen. *otsi (Dyar, 1904) 1941. Geometridae. In A. Seitz (ed.), Die variata (Denis & Schiffermu¬ller, 1775) Gross Schmetterlinge des Indoaustral- *variolata (Staudinger, 1899) 2000 CHOI: CLADISTIC ANALYSIS OF THERINI 25

vetustata (Denis & Schiffermu¬ller, 1775) Heterothera Inoue, 1943 consimilis (Warren, 1888) *comitabilis (Prout, 1923) dentifasciata (Hampson, 1895) *distinctata Choi, 1998 eclinosis Choi, 1998 etes (Prout, 1926) hoenei Choi, 1998 incerta Inoue, 1986 kurenzovi Choi, Viidalepp & Vasjurin, 1998 *mussooriensis Choi, 1998 *obscurata Choi, 1998 postalbida (Wileman, 1911) quadrifulta (Prout, 1938) serraria (Lienig, 1846) serrataria (Prout, 1914) sororcula (Bastelberger, 1909) *stamineata Choi, 1998 taigana (Djakonov, 1926) tephroptilus (Heydemann, 1961) *triangulata Choi, 1998 Characters and Character States undulata (Warren, 1888) yunnanensis Choi 1998 1. Antenna of male pectinate (0), filiform (1). 2. Sacculus processes simply pointed (0), bifid (1). APPENDIX 2 3. Uncus present (0), absent (1). Reanalysis of Viidalepp’s Data Set 4. Saccus heavily sclerotized and broad, angu- lated platelike (0), massive and rounded (1), Here I present a reanalysis of the data presented medially projected (2). by Viidalepp (1980). Fifteen characters appeared 5. The last tergite of female abdomen with long in the original diagram and most of them were binary, except for four characters (characters 4, 6, apophyses and telescopic (0), with short 7, 14) that he stated plesiomorphic, intermediate apophyses and strongly sclerotized to limit and apomorphic. I rescored these characters as the mobility of ovipositor (1). multistate and treated them as additive in the anal- 6. The ductus and the collar of corpus bursae ysis, because Viidalepp considered that the inter- strongly sclerotized (0), intermediate (1), mediate state was between the plesio- and apo- membraneous (2). morphy. Character number 15 was treated as in- 7. Sterigma well developed and sclerotized (0), applicable in the species of Thera and Heteroth- intermediate (1), simple and slightly sclero- era because the male antennae of those species tized (2). are filiform. The recoded data matrix was ana- 8. Eighth sternite of female abdomen simple (0), lyzed by Hennig86 using the implicit enumeration (ie*) using an all zero outgroup. This analysis re- fused to sterigma (1). sulted in 55 equally parsimonious trees with 33 9. Costa indistinct and fused to other part of val- steps, 57 consistency index, and 82 retention in- va (0), distinct (1). dex. The strict consensus is shown in figure 1B. 10. Sacculus indistinct and fused to other part of valva (0), distinct (1). 11. Cornuti present in vesica (0), present in the dorsal part of aedeagus, corona-shaped (1). 12. Cornutus absent (0), present (1). 13. Uncus short (0), long (1). 14. Antenna of male pectinate (0), filiform with constricted or rounded ventral surfaces (1), fi- liform with cylindrical antennomeres (2). 15. Antennal pectination short (1), long (0). AMNH NOVITATES No RH this page!! Friday Mar 31 2000 11:14 AM 2000 novi 00166 Mp 28 • Allen Press DTPro System File # 01cc

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