Lepidoptera: Geometridae: Ennominae) Linda M

This article was downloaded by: [Michigan State University] On: 24 December 2014, At: 11:29 Publisher: Taylor & Francis Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer House, 37-41 Mortimer Street, London W1T 3JH, UK

Systematics and Biodiversity Publication details, including instructions for authors and subscription information: http://www.tandfonline.com/loi/tsab20 Moths of the Neotropical genera Ischnopteris, Stegotheca and Rucana (Lepidoptera: Geometridae: Ennominae) Linda M. Pitkin a a Department of Entomology , Natural History Museum , Cromwell Road, London, SW7 5BD, UK E-mail: Published online: 11 Mar 2010.

To cite this article: Linda M. Pitkin (2005) Moths of the Neotropical genera Ischnopteris, Stegotheca and Rucana (Lepidoptera: Geometridae: Ennominae), Systematics and Biodiversity, 3:1, 13-96, DOI: 10.1017/S1477200004001616 To link to this article: http://dx.doi.org/10.1017/S1477200004001616

PLEASE SCROLL DOWN FOR ARTICLE

Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”) contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors make no representations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose of the Content. Any opinions and views expressed in this publication are the opinions and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of the Content should not be relied upon and should be independently verified with primary sources of information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to or arising out of the use of the Content. This article may be used for research, teaching, and private study purposes. Any substantial or systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http:// www.tandfonline.com/page/terms-and-conditions Systematics and Biodiversity 3 (1): 13-96 Issued 29 April 2005 doi: 10.1017/S1477200004001616 Printed in the United Kingdom © The Natural History Museum

Moths of the Neotropical genera Ischnopteris, Stegotheca and Rucana (Lepidoptera: Geometridae: Ennominae)

Linda M. Pitkin Department of Entomology, Natural History Museum, Cromwell Road, London SW7 5BD, UK Email: [email protected]

submitted June 2004 accepted September 2004

Contents Abstract 13 Introduction 14 Taxonomic history of Ischnopteris, Stegotheca and Rucana 14 Structures of taxonomic note 15 Materials and methods 15 Depositories of material 16 Ischnopteris Hubner 16 Description 17 Checklist of species 46 Key to species 46 Ischnopteris fabiana species group 48 Ischnopteris chryses species group 57 Species not placed to group 63 Species excluded from Ischnopteris: 'Ischnopteris' discolor species group 77 Stegotheca Warren 80 Description 80 Checklist of species 81 Key to species 81 Species 81 Rucana Rindge 86 Description 86 Checklist of species 86 Key to species 87

Downloaded by [Michigan State University] at 11:29 24 December 2014 Species 87 Acknowledgements 94 References 94 Index 95

Abstract Three closely related Neotropical genera of ennomine moths of the tribe Nacophorini are revised: Ischnopteris, Stegotheca and Rucana. These genera are all redefined, and the concepts of Stegotheca and Rucana are extended; the previously monobasic Stegotheca is markedly expanded by the inclusion of two other genera as junior synonyms: Salasaca and Canelo. In Ischnopteris 37 species are recognized, 17 of which are newly described; six species, including one newly described, are recognized in Stegotheca, and nine species, including two newly described, are recognized in Rucana. All species of these genera are described fully, and several new synonyms and other taxonomic changes are published. A vital aspect of the work is to serve as an identification guide. To this end keys to the species are provided as are illustrations of all species: the moths are illustrated in colour; genitalia and other structures are in half-tone. Information on the larvae and hostplants is given for several species of Ischnopteris, and some larvae are figured, including the larva of a new species. This study also describes highly unusual morphological modifications of some moths in these genera, which have not been previously recorded in the Geometridae.

13 14 Linda M. Pitkin

Keywords Ennominae, Geometridae, hostplants, larvae, Lepidoptera, moths, Nacophorini, Neotropical region, retinaculum, taxonomy

Introduction Although the relationships of Ischnopteris have yet to be resolved fully, some advance is made here in establishing af- The Geometridae are one of the three largest families of moths finities with certain other genera within the Nacophorini, and in number of species, and one of the chief areas of focus of so my initial study was expanded to include them in this pa- the Macrolepidoptera Research Group's programme of work per. These are Stegotheca and Rucana, smaller genera with at the Natural History Museum, London, UK (BMNH). Geo- six and nine species respectively recognised here. There are metrid moths occur worldwide but they are most diverse in some apomorphic features shared by some members of the the tropics, particularly the Neotropical region, which is home three genera, suggesting a close relationship between them. to about one-third of the species of the subfamily Ennominae. The most notable character is an enlarged retinaculum of the My recent review of the genera of Neotropical ennomine moths male fore wing, usually with a central semi-transparent 'win- (Pitkin, 2002) provided a framework for that subfamily at the dow' (see 'Structures of taxonomic note'). This is present in genus level, preparing the ground for better assessment of nearly all Ischnopteris species and in one or more species of species names and the wealth of data associated with them. each of the other two genera, but not in any other member of the The broad coverage of that study (267 genera with about Nacophorini. However, without further analysis of characters 3400 species) inevitably left tremendous scope for more de- across the entire tribe, there is insufficient evidence to determ- tailed studies, and the present work selects a few genera of ine whether or not the three genera under study constitute a Neotropical Ennominae for in-depth revision. monophyletic group. For future investigation, comparison with The genera chosen for this study, Ischnopteris Hubner, the genus Quillaca Rindge could be useful. Certain characters Stegotheca Warren, and Rucana Rindge, are members of the of Quillaca and Rucana are similar (see under 'diagnosis and Nacophorini. This tribe of mainly large and often sturdy monophyly' of Rucana), although moths of Quillaca are much ennomine moths is strongly represented in the Neotrop- larger and have distinctively modified genitalia. ics, with more than 40 genera (Pitkin, 2002). The mono- Nacophorine moths often have fairly elongate fore wings, phyly of the Nacophorini is not clearly established. Members and this is particularly marked in Ischnopteris, Stegotheca and of the tribe typically have a pair of processes in the anel- Rucana (e.g. Figs 11, 73, 82), so much so that they do not have lus of the male genitalia, which is situated usually postero- a typically geometrid moth appearance. They could even be laterally at the end of the juxta. Similar processes are mistaken for Notodontidae, except for the long slender abdo- present in the tribe Lithinini, but they are usually more men in the male sex, particularly in Ischnopteris and Rucana. slender and setose. Nacophorine processes are usually curved Like many other nacophorine moths, Ischnopteris species are and tapered from a wide base to a pointed apex, without generally fairly large geometrids (with a fore wing length of up setae, and are typically large and heavily sclerotized. How- to 29 mm), whereas those of Stegotheca and Rucana look like ever, there are exceptions to this in both tribes. For fur- smaller versions of Ischnopteris. Ischnopteris and Rucana are ther discussion of characters of both tribes see Pitkin (2002: typical nacophorines in that they have a pair of well-sclerotized pp. 133-134). The New World genera of the Nacophorini processes, which are situated postero-laterally at the end of the were revised by Rindge (1983), with the description of 19 new juxta, in the anellus of the male genitalia. These processes are genera, and have since been the subject of further research and curved, usually pointed, and not setose, and they are tapered or Downloaded by [Michigan State University] at 11:29 24 December 2014 revision (e.g. by Pitkin (2002) and studies on individual genera have an extended base. Stegotheca, like several other nacophor- by Parra & Beeche (1986) and Dias (1998)). The Nacophorini ine genera, usually has the processes of the anellus secondarily are also well represented in Australia, and research on the tribe reduced or lost. Stegotheca is unusual in having a fovea in the there has been carried out by McQuillan et al. (e.g. 2001). male fore wing, a feature not known in any other genera con- Ischnopteris, Stegotheca and Rucana occur widely across firmed as belonging in the Nacophorini, although present in the Neotropics but not outside that region. Ischnopteris is the various ennomines in some other tribes. second largest genus of Neotropical nacophorines, exceeded in numbers of species only by Mallomus Blanchard. Prior to the present study, Ischnopteris comprised 34 named species Taxonomic history of Ischnopteris, (Pitkin, 2002), and I now revise that to 37 species. Ischnop- teris is a particularly significant genus because its concept Stegotheca and Rucana has been confused in the past. Many species have at times The concept of Ischnopteris has undergone enormous change been incorrectly placed in this genus and later moved to since the genus was described by Hubner in 1823. Early stud- other nacophorine genera, with further transfers in this paper, ies were based solely on external features, with the result that which has had an extensive impact on the understanding of the many unrelated species were misplaced in the genus. Genitalic Neotropical component of the tribe (see 'Taxonomic history characters were not examined until much later, and it was not of Ischnopteris, Stegotheca and Rucana' for details). until Rindge's generic revision of the New World Nacophorini Neotropical geometrid moths 15

(1983) and my review of the genera of Neotropical ennom- Structures of taxonomic note ine moths (Pitkin, 2002), that many of the misconceptions involving Ischnopteris were resolved. In all, more than 20 spe- Some highly unusual morphological modifications occur in cies that were either originally described in Ischnopteris or its Ischnopteris and related genera. Most species of Ischnop- junior synonyms, or subsequently transferred to Ischnopteris, teris have the retinaculum of the male fore wing enlarged and have since been moved to various other genera: Ceratonyx with a central semi-transparent 'window' (e.g. Fig. 92). The Guenee,´ Cidariophanes Warren, Erilophodes Warren, Canelo 'window' forms a slight depression in the retinaculum and is Rindge (synonymized here with Stegotheca), Charca Rindge, a membrane free of scales, although sometimes overlapped by Cundinamarca Rindge, Nazca Rindge, Quillaca, and Rucana surrounding scales. This character occurs also in Stegotheca (all Nacophorini), and Dasciopteryx Warren (possibly Neph- speculifera and in Rucana mediosecta, and the retinaculum is odiini). Rindge (1983) synonymized Ischnopterix Hubner and enlarged but lacks a transparent central 'window' in Stegotheca Amblurodes Warren with Ischnopteris, but he recognised two amissa. An enlarged retinaculum occurs in two Asian genera other genera, Trichostichia Warren andAnischnopteris Rindge, in the tribe Baptini of the Ennominae: Parasynegia Warren and which were synonymized with Ischnopteris by Parsons et al. to a lesser degree Yashmakia Warren (see Holloway, [1994], (1999) and by Pitkin (2002), respectively. pp. 72), but the 'window' described here, in Ischnopteris and The present study brings further refinement: nine more relatives, has not previously been noted in the Geometridae, or species are moved out of Ischnopteris, some of which remain in any other Lepidoptera as far as I am aware. unplaced, but others go to Quillaca, Rucana and Stegotheca, A fovea is present on the male fore wing of Stegotheca, and one other species that had previously been transferred from and although this is not an uncommon structure in the Ischnopteris to Chrysomima Warren is now moved to Rucana. Ennominae, it is rare in the tribe Nacophorini. In the Neotrop- Seventeen new species of Ischnopteris are described here, and ical component it is known only in two other genera, Mimo- five species plus one subspecies are synonymised. phyle Warren and Plesiophyle Dognin, neither of which are The genera Stegotheca and Rucana have also undergone confirmed as belonging in this tribe. The fovea of Stegotheca substantial revision. Stegotheca had been largely ignored since is sometimes made more conspicuous by the presence of raised Warren's description of it as a monobasic genus in 1900, until scales at its edge, particularly in cythereata, which has these in Pitkin (2002: 205) I noted the possibility that Rindge's scales large and blade-shaped (Fig. 91). The fovea is usually genera Canelo and Salasaca might be synonymous with it. situated near the base of the wing, between the cubital and That is confirmed here, and with species newly transferred to anal veins, but S. speculifera has instead a fovea-like structure Stegotheca, and the description of a new species, Stegotheca between the subcosta and radius. This structure in speculifera now has six species. Rucana was described by Rindge (1983), lies over the retinaculum, and it is flanked on its costal edge by who suggested that it should include three species, but men- a double comb-like row of modified scales on the underside of tioned only one by name, abnormipalpis Warren. I transferred the wing (Fig. 92). Stegotheca speculifera also has a modified three species from Ischnopteris to Rucana in Pitkin (2002), male frenulum, which is elongate and slightly swollen subap- and add five more species in the present study, including two ically, and extends to directly below the fovea-like structure. newly described here, bringing the total to nine species. I suspect that there are further species to be discovered in Materials and methods Ischnopteris, Stegotheca and Rucana. Although I have madeMaterialsandme efforts to study as much material as possible, with help- Study of as much available material as possible was invaluable ful support from colleagues in other institutions, some spe- in helping to establish species identities and variation. This cies are apparently very rare. Others, with undistinguished work is based on more than 900 specimens, examined from external appearance, have been discovered only by dissec- the collections of the BMNH, and from several other important Downloaded by [Michigan State University] at 11:29 24 December 2014 tion. One intriguing mystery cannot be cleared up at present. sources (listed under 'Depositories of material'). I examined It has been suggested to me (by Vitor Becker and John about 400 genitalia slides, the majority of which were made Rawlins, pers. comm.) that the type specimen of Pagrasana specifically for this study by Shayleen James (BMNH) and bermejona Schaus might belong in Ischnopteris. This type, a myself. My study benefited enormously from specimens I re- female specimen, is the sole representative not only of berme- ceived on loan, and also some digital images, from helpful jona but also of the genus Pagrasana Schaus, and is currently colleagues in other institutions. The current worldwide move placed in the Notodontidae where it was originally described. towards digitisation of collections is in its early stages, and The specimen resides in the Carnegie Museum, Pittsburgh, only a tiny percentage of specimens held in collections is USA, but, unfortunately, it has been mislaid since Vitor Becker available as images on the internet. However, already, the in- and John Rawlins saw it, and so I have been unable to include it creasing ease of producing and sending digital images around in my study. If their suspicions should later prove to be correct, the world is proving a useful alternative to conventional exam- then Pasagrana would become a junior synonym of Ischnop- ination of material, as has been noted by Valdecasas, Becerra & teris. However, the figure of the type specimen (Schaus, 1939: Marshall (1997). I have been able to enhance the material ex- 329, pl. 31, fig. 14) seems to me to be unlikely to be an Ischnop- amined in my study, in species that have moths of sufficiently teris species. Instead the general appearance of the moth re- distinctive appearance, by reference to some digital images of sembles that of other genera of Nacophorini — Oratha Walker moths in lieu of actual specimens. This has involved only a and Marcodava Walker. small percentage of the total material studied, but it has been 16 Linda M. Pitkin

highly valuable in several instances, notably in extending the Additional digital images published here were kindly provided paratype series of the new species Ischnopteris chavesi and as follows: a moth (Fig. 77) by the photographic depart- Rucana chaconi. It has also enabled the association of biolo- ment of the Zoological Museum, University of Copenhagen, gical information, including images of larvae, with I. chavesi Denmark, during my study visit supported by a grant from and several other species; such information is hard to come Copenhagen Biosystematics Centre (COBICE), and all the by for Neotropical geometrid moths. Digital images were also digital images of larvae (Plate 1) by Daniel H. Janzen, Uni- useful in checking identities of some type specimens and, in versity of Pennsylvania, Philadelphia, USA. The latter im- the case of Ischnopteryx velledata Moschler, obviated the need ages are taken from high resolution versions of images on to borrow from abroad a unique and fragile specimen dating the Janzen & Hallwachs Caterpillar Rearing Database at back to the 1880s. I have cited specimens examined in this way http://janzen.sas.upenn.edu/. Moths in the BMNH Collection as 'digital photographs' under 'material examined'. It should that were imaged have had their data added to the Entomology also be noted that depositories of specimens cited under 'ma- specimen database of the BMNH. terial examined' state the current holders of the material, but In addition to The Times Atlas of the World (various edi- a small number of duplicate specimens (not primary types) tions), Appendix I in Brown (1979) proved useful for check- may subsequently be requested for the BMNH. All specimens ing locality names, and I located many obscure place names examined in this study bear my det. labels, except for a small on various websites of the internet using the search engine number of specimens where only photographs were examined. GoogleTM. I have cited data of altitude in metres, converting I have made lectotype designations where necessary to them from feet where necessary. stabilise the nomenclature, particularly as there are instances of mixed syntype-series. In selecting lectotypes, I chose spe- Depositories of material cimens that had already been labelled as the 'Type' in the collections of the BMNH and the USNM, so long as these proved AMNH American Museum of Natural History, New York, entirely suitable in other respects. Lectotypes I selected from USA Warren's syntypes are specimens bearing Warren's salmon- BMNH Natural History Museum, London, UK (formerly coloured determination labels. I have noted that there are sev- British Museum (Natural History)) eral minor discrepancies between the data as stated by Warren CMNH Carnegie Museum, Pittsburgh, USA and those on the specimen labels. Type specimens of species DHJ DHJ (Daniel Janzen) and WH (Winnie Hallwachs) in this study that were described by Felder & Rogenhofer ACG inventory collection, University of Pennsyl- (1875) are regarded as holotypes, as in each case the descrip- vania, Philadelphia, USA tion takes the form solely of a single figured specimen, with INBio Instituto Nacional de Biodiversidad, Santo Domingo, citation of its name, sex and locality. Heredia, Costa Rica Within each of the three genera under study, species are MNHU Museum fur Naturkunde der Humboldt-Universit¨at, treated in alphabetical order, except that some are placed in Berlin, Germany species groups where applicable. I have followed the Res- SMF Forschungsinstitut Senckenberg, Frankfurt am Main, olution adopted by the Societas Europaea Lepidopterologica Germany (Sommerer, 2002: 202) in using the original gender given for SMNS Staatliches Museum fur Naturkunde, Stuttgart, species-group names. In describing new species, I have either Germany chosen names based on descriptive terms referring to the mor- USNM National Museum of Natural History, Washington, phology, or I have named the species after the collector of DC, USA (formerly United States National Museum) the holotype (or in the case of brehmi sp. n. after the primary VOB Vitor O. Becker private collection, Brasilia, Brazil collector of material of the species). Descriptions of wing pat- ZMUC Zoological Museum, University of Copenhagen, Downloaded by [Michigan State University] at 11:29 24 December 2014 tern of moths refer to the upper side of the wings, unless stated Denmark otherwise. The terminology of the genitalia largely follows ZMUJ Zoological Museum, Jagiellonian University, that of Klots (1956), and some structures are labelled in Figs Krakow, Poland 95 and 209A. Various spines and other sclerotised structures ZSM Zoologische Staatssammlung, Munich, Germany are referred to on the aedeagus; where these structures occur on the vesica, they are cornuti, whereas those on other parts of the aedeagus are not cornuti. In dissecting females, Ischnopteris Hubner abdominal tergum 7 was sometimes separated from its cor- Ischnopteris Hubner, [1823] 1806: pl. [222]. Type species: responding sternum using the technique explained in Pitkin Ischnopteris chlorata Hubner, [1823] 1806, by monotypy. (1993: 41). The illustrations of whole moths and of their gen- Group 3 of Rindge (1983: 224, 228, 230). italia were prepared from digital scans made with a Synoptics Ischnopterix Hubner, [1825] 1816: 332. Type species: Imaging System (http://www.synoptics.co.uk) mainly by Tara Phalaena fabiana Stoll, 1782, by monotypy. [Synony- Lewis (BMNH volunteer) and Shayleen James, and were sub- mized by Rindge, 1983: 224.] sequently edited in Adobe® Photoshop® by Ms James and Ischnopteryx Agassiz, 1847: 196. [Unjustified emendation of myself. The combined use of Synoptics and Photoshop tools Ischnopterix Hubner.] gave results of enhanced clarity, particularly in achieving high Syrtodes Guenee, [1858]: 451. Type species: Syr- contrast and sharply focused detail in images of moth genitalia. todes chloroclystata Guenee, [1858], by subsequent Neotropical geometrid moths 17

designation by Fletcher, 1979: 199. [Synonymy cited by stilla not often forming distinct pair of bulbous lobes, but these Druce, 1893: 157.] sometimes well developed (e.g. in bryifera). Pair of processes Trichostichia Warren, 1895: 131. Type species: Syrtodes of anellus extending postero-laterally from juxta, processes bifinita Walker, 1862, by original designation. [Synonymy large, curved and usually pointed, tapered or with base ex- cited by Parsons et al., in Scoble (1999).] tended; juxta usually with, but occasionally without, long, Amblurodes Warren, 1900: 200. Type species: Amblur- medio-posterior, spinulose, furca-like process; cristae often odes commixta Warren, 1900, by original designation. weak but sometimes moderately prominent. Aedeagus with [Synonymized by Rindge, 1983: 224.] tapered posterior end or extension, with longitudinal wrinkles Anischnopteris Rindge, 1983: 230. Type species: Ischnop- and denticulate ridges; vesica with spines or setae usually in teryx chryses Druce, 1893, by original designation [given elongate group, sometimes arising from sclerotized strip. as chryses Godman & Salvin]. [Synonymized by Pitkin, FEMALE GENITALIA (Figs 209-234). Bursa copulatrix 2002: 213.] sometimes long. Signum usually present: dentate, elliptical to triangular or rounded, hollow, sometimes forming pocket; Description base of signum (neck-like region that connects signum to wall ADULTS (Figs 1-65, 92, 250). Fore wing length: , 15- of corpus bursae) situated off-centre, adjacent to rim of signum 25 mm; , 18—29 mm. Antenna usually simple filiform in both (except in pronubata and klagesi, where base is almost central sexes, although thicker in male, but occasionally serrate or in signum, and in chlorophaearia); signum absent in ochro- bipectinate in male (xylinata and bipectinata). Front of head prosthia. swollen. Wing pattern variable and often sexually dimorphic: fore wing mottled brown, dark to pale, with or without white Diagnosis blotches or grey-green markings, usually without red flecks Moths of Ischnopteris are moderately large, typical nacophor- or these inconspicuous if present; hind wing often plainer and ines, but with fairly elongate fore wings and an unusually ranging from much paler to slightly darker than fore wing, slender male abdomen. Some Ischnopteris species have an or- but, in some species, with large orange or yellowish region, ange blotch on the hind wings, but that is seen also in several either apical or over most of wing except for broad brown other nacophorine genera: Charca, Chrysomima, sometimes margin. Fore wing of female often with roughly diamond- in Rucana and to a lesser extent Stegotheca, and Quillaca ver- shaped central area. Moths that have orange or yellow on the sipennis (Warren). The last species is very easily confused upper surface of the hind wing have similar colour also on with Ischnopteris chryses (except for the very different gen- parts of the under-side of fore and hind wing. Brown colour italic structures). Male members of Ischnopteris can usually and markings on under-side of wings subject to individual be recognized by the enlarged male retinaculum, with a 'win- variation. Wings with outer margin often weakly wavy; fore dow', but these modifications are occasionally absent, and they wing generally elongate, without fovea; male retinaculum are not exclusive to Ischnopteris; they occur also occasionally usually enlarged and with transparent or semi-transparent cent- in Stegotheca and Rucana. The expanded anal margin of the ral 'window' varying in size, but occasionally not modified; hind wing is often conspicuous in Ischnopteris but it occurs male with or without hairy lobe at midpoint of dorsum of fore also in some other nacophorine genera, including Charca and wing; hind wing of male with anal margin slightly expanded or Quillaca. strongly expanded to form distinct flap, usually with weak to pronounced hair-scale tuft. Metathorax with dorsal tuft; male Ischnopteris can be distinguished and defined best by abdomen elongate and slender, with longitudinal row of dorsal several putative autapomorphies present in the male genitalia. tufts, second tuft usually large, tufts not usually evenly rounded These are: the presence of one or more dense patches of ventral except in chlorata. Tympanal cavity usually with slight pouch setae on the uncus (except in bipectinata), the small pointed Downloaded by [Michigan State University] at 11:29 24 December 2014 or sac extending from wall. Sternite 3 of male abdomen with process of the sacculus (weakest in xylinata and bipectinata), patch of setae subtriangular or semicircular, occasionally very situated near to the small fold towards the base of the valva weak or absent. bearing a dense patch of dorsal setae. The last feature occurs to a lesser degree in a few other nacophorines, but it is more MALE GENITALIA (Figs 95-132, 154-188). Uncus fre- strongly defined in Ischnopteris. In the female, a signum is quently a hood-shaped structure, large and complex, but vary- usually present in Ischnopteris species (except ochroprosthia), ing from narrow (as in chlorata) to broad (as in bifinita), and whereas it is absent in Rucana. strongly developed in the fabiana-group; uncus with ventral setae forming one or more dense patches, except in bipectinata. Variation Socii better developed and longer than usual for Ennom- Many Ischnopteris species are variable in having male moths inae, digitate, usually membranous but semi-rigid in latijuxta. with or without a conspicuous white blotch on the fore wing, Gnathos with one to two median processes: sclerotized, near the midpoint of the costa. Sexual dimorphism is common pointed, usually spine-like and curved. Valva with apical costal in the wing pattern of Ischnopteris, females but not males lobe or process and with costal band; sacculus with small, typically having a roughly diamond-shaped central area of the sometimes minute, pointed ventral process; valva with small fore wing, and sexual dimorphism sometimes also occurs in dense patch of dorsal setae (usually deciduous, very long and the wing shape (notably in bryifera). As many species differ fine), situated towards base of valva, in sacculus region. Tran- only subtly in their wing patterns, the presence of sexual 18 Linda M. Pitkin Downloaded by [Michigan State University] at 11:29 24 December 2014

Figs 1-15 Moths. Ischnopteris. 1, beckeri sp. n. ; 2, bifinita ; 3, bifinita ; 4, bryifera ; 5, bryifera (variation); 6, bryifera ; 7, chlorophaearia ; 8, chlorophaearia ; 9, fabiana ; 10, fabiana ; 11, illineata ; 12, illineata (variation); 13, illineata ; 14, klagesi sp. n. ; 15, klagesi . Scale line = 10 mm. Neotropical geometrid moths 19

25 Downloaded by [Michigan State University] at 11:29 24 December 2014

Figs 16-30 Moths. Ischnopteris. 16, rostellaria form A ; 17, rostellaria form B ; 18, chavesi sp. n. ; 19, chavesi sp. n. ; 20, chryses ; 21, chryses ; 22, fasslisp. n. ; 23, FASSLIsp. n. ; 24, hoffmani sp. n. ; 25, inornata sp. n. ; 26, janzeni sp. n. ; 27, lata sp. n. ; 28, watsoni sp. n. ; 29, watsoni sp. n. ; 30, albiguttata . Scale line = 10 mm. 20 Linda M. Pitkin Downloaded by [Michigan State University] at 11:29 24 December 2014

Figs 31-45 Moths. Ischnopteris. 31, albiguttata (variation); 32, albiguttata ; 33, aurudaria ; 34, aurudaria ; 35, bifurcata sp. n. form A cf ; 36, bifurcata sp. n. ; 37, bipectinata sp. n. ; 38, brehmi sp. n. ; 39, brehmi sp. n. ; 40, catocalata ; 41, chlorata ; 42, chlorata ; 43, chloroclystata ; 44, chloroclystata ; 45, fasciata sp. n. . Scale line = 10 mm. Neotropical geometrid moths 21 Downloaded by [Michigan State University] at 11:29 24 December 2014

58 Figs 46-60 Moths. Ischnopteris. 46, hirsuta sp. n. ; 47, latijuxta sp. n. ; 48, lemoulti sp. n. ; 49, miseliata ; 50, miseliata ; 51, multistrigata ; 52, multistrigata ; 53, obtortionis ; 54, obtortionis ; 55, ochroprosthia ; 56,palmerisp. n. ; 57,pronubata ; 58, pronubata ; 59, seriei ; 60, seriei . Scale line = 10 mm. 22 Linda M. Pitkin Downloaded by [Michigan State University] at 11:29 24 December 2014

Figs 61-75 Moths. Ischnopteris and Stegotheca. 61, Ischnopteris stenoptila ; 62, Ischnopteris xylinata ; 63, Ischnopteris xylinata ; 64, Ischnopteris xylinata small form A ; 65, Ischnopteris xylinata small form B ; 66, 'Ischnopteris' discolor , 67, 'Ischnopteris' discolor ; 68, 'Ischnopteris'festa ; 69, 'Ischnopteris' festiva ; 70, Stegotheca albipennis ; 71, Stegotheca amissa ; 72, Stegotheca costiplaga ; 73, Stegotheca costiplaga (variation); 74, Stegotheca cythereata typical form ; 75, Stegotheca cythereata brown form . Scale line = 10 mm. Neotropical geometrid moths 23 Downloaded by [Michigan State University] at 11:29 24 December 2014

Figs 76-90 Moths. Stegotheca and Rucana. 76, Stegotheca phalangifera sp. n. ; 77, Stegotheca speculifera form A ; 78, Rucana abnormipalpis ; 79, Rucana bisecta form A ; 80, Rucana bisecta form B ; 81, Rucana brunneoviridis ; 82, Rucana chaconi sp. n. ; 83, Rucana degener ; 84, Rucana fidelis ; 85, Rucana fidelis ; 86, Rucana marmorata ; 87, Rucana mathanisp. n. ; 88, Rucana mediosecta ; 89, Rucana sclerovesica sp. n. ;90, head of Rucana abnormipalpis (front view). Scale line (for whole moths) = 10 mm. 24 Linda M. Pitkin Downloaded by [Michigan State University] at 11:29 24 December 2014

Figs 91-99 Stegotheca and Ischnopteris. 91 detail of fore wing (upper side) showing fovea of Stegotheca cythereata ; 92, detail of fore wing (under-side) showing retinaculum of Ischnopteris bifurcata ; 93, detail of fore wing (under-side) showing fovea-like structure of Stegotheca speculifera ; 94, abdomen of Stegotheca amissa ; 95-99, male genitalia of Ischnopteris: 95, beckeri sp. n.; 96, bifinita; 97, bryifera; 98, chlorophaearia; 99, fabiana. Neotropical geometrid moths 25

103 Downloaded by [Michigan State University] at 11:29 24 December 2014

1Û4B Figs 100-105 Male genitalia. Ischnopteris. 100, illineata; 101, klagesi sp. n. ; 102, rostellaria form A; 103, rostellaria form B; 104A, chavesi sp. n.; 104B, detail of chavesi sp. n.; 105, chryses. 26 Linda M. Pitkin

108A Downloaded by [Michigan State University] at 11:29 24 December 2014

108B Figs 106-109 Male genitalia. Ischnopteris. 106A, fassli sp. n.; 106B, detail of fassli sp. n.; 107, hoffmani sp. n.; 108A, inornata sp. n.; 108B, detail of inornata sp. n.; 109, janzeni sp. n. Neotropical geometrid moths 27

HOB Downloaded by [Michigan State University] at 11:29 24 December 2014

112A 112B

Figs 110-112 Male genitalia. Ischnopteris. 110A, lata sp. n.; 110B, detail of lata sp. n.; 111A, watsoni sp. n.; 111B, detail of watsoni sp. n.; 112A, albiguttata; 112B, detail of albiguttata. 28 Linda M. Pitkin

116 Downloaded by [Michigan State University] at 11:29 24 December 2014

117B Figs 113-117 Male genitalia. Ischnopteris. 113, aurudaria; 114, bifurcata sp. n. form A; 115, bifurcata sp. n. form B; 116, bipectinata sp. n.; 117A, brehmi sp. n.; 117B, detail of brehmi sp. n. Neotropical geometrid moths 29

121 Downloaded by [Michigan State University] at 11:29 24 December 2014

122

Figs 118-123 Male genitalia. Ischnopteris. 118, chlorata; 119, chloroclystata; 120, fasciata sp. n.; 121, hirsuta sp. n.; 122, latijuxta sp. n.; 123, lemoulti sp. n. 30 Linda M. Pitkin

127 Downloaded by [Michigan State University] at 11:29 24 December 2014

128

Figs 124-129 Male genitalia. Ischnopteris. 124, miseliata; 125, multistrigata; 126, obtortionis; 127, ochroprosthia; 128, palmeri sp. n.; 129, pronubata. Neotropical geometrid moths 31

133 Downloaded by [Michigan State University] at 11:29 24 December 2014

134

Figs 130-131 Male genitalia. Ischnopteris. 130, seriei; 131, xylinata small form A; 132, xylinata small form B; 133, 'I.' discolor; 134, 'I.' festa; 135, 'I.' festiva. 32 Linda M. Pitkin Downloaded by [Michigan State University] at 11:29 24 December 2014

140

Figs 136-141 Male genitalia. Stegotheca. 136, albipennis; 137, amissa; 138, costiplaga; 139, cythereata; 140, cythereata brown form; 141, phalangifera sp. n. Neotropical geometrid moths 33 Downloaded by [Michigan State University] at 11:29 24 December 2014 144

146B Figs 142-146 Male genitalia. Stegotheca and Rucana. 142A, Stegotheca speculifera form A; 142B, detail of Stegotheca speculifera form A; 143, Stegotheca speculifera form B; 144, Rucana abnormipalpis; 145, Rucana bisecta; 146A, Rucana brunneoviridis; 146B, detail of brunneoviridis. in Linda M. Pitkin Downloaded by [Michigan State University] at 11:29 24 December 2014 150 A

152

Figs 147-152 Male genitalia. Rucana. 147, chaconi sp. n.; 148, degener; 149, fidelis; 150A, marmorata form A; 150B, marmorata form B; 152, mediosecta. Neotropical geometrid moths 35 Downloaded by [Michigan State University] at 11:29 24 December 2014

Figs 153-166 Male genitalia. 153, Rucana sclerovesica sp. n. 154-166, aedeagus of Ischnopteris. 154, beckeri sp. n.; 155, bifinita; 156, bryifera; 157, chlorophaearia; 158, fabiana; 159, illineata; 160, klagesi sp. n.; 161, rostellaria form A; 162, chavesi sp. n.; 163, s; 164, fassli sp. n.; 165, hoffmani sp. n.; 166, inornata sp. n. 36 Linda M. Pitkin

167 Downloaded by [Michigan State University] at 11:29 24 December 2014

177

Figs 167-181 Male genitalia (aedeagus). Ischnopteris. 167, janzeni sp. n.; 168, lata sp. n.; 169, watsoni sp. n.; 170, albiguttata; 171, aurudaria; 172, bifurcata sp. n. typ. form; 173, bipectinata sp. n.; 174, brehmi sp. n.; 175, chlorata; 176, chloroclystata; 177, fasciata sp. n.; 178, hirsuta sp. n.; 179, latijuxta sp. n.; 180, lemoulti sp. n.; 181, miseliata. Neotropical geometrid moths 37

187 Downloaded by [Michigan State University] at 11:29 24 December 2014

Figs 182-196 Male genitalia (aedeagus). 182-191, Ischnopteris: 182, I. multistrigata; 183, I. obtortionis; 184, I. ochroprosthia; 185, I. palmeri sp. n.; 186, I. pronubata; 187, I. seriei; 188, I. xylinata typical form; 189, 'I.' discolor; 190, 'I.' festa; 191, 'I.' festiva; 192-196, Stegotheca: 192, S. albipennis; 193, S. amissa; 194, S. costiplaga; 195, S. cythereata; 196, S. cythereata brown form. 38 Linda M. Pitkin Downloaded by [Michigan State University] at 11:29 24 December 2014

Figs 197-209A Genitalia. 197-198, male (aedeagus) of Stegotheca: 197, S. phalangifera sp. n.; 198, S. speculifera; 199-209A, male (aedeagus) of Rucana: 199, R. abnormipalpis; 200, R. bisecta; 201, R. brunneoviridis; 202, R. chaconi sp. n.; 203, R. degener; 204, R. fidelis; 205, R. marmorata form A; 206, R. mathani sp. n.; 207, R. mediosecta; 208, R. sclerovesica sp. n.; female: 209A, detail of Ischnopteris bifinita. Neotropical geometrid moths 39

210B

209B Downloaded by [Michigan State University] at 11:29 24 December 2014

212 213A

Figs 209B-214 Female genitalia. Ischnopteris. 209B, bifinita; 210A, bryifera; 210B, detail of bryifera; 211, chlorophaearia; 212, fabiana; 213A, illineata; 213B, detail of illineata; 214, klagesi sp. n. 40 Linda M. Pitkin

>•>

217B à ! ! : 215 216 217A Downloaded by [Michigan State University] at 11:29 24 December 2014

218A

\ '

Figs 215-219 Female genitalia. Ischnopteris. 215, rostellaria form A; 216, chavesi sp. n.; 217A, chryses; 217B, detail of chryses; 218A, fassli sp. n.; 218B, detail of fassli sp. n.; 219A, watsoni sp. n.; 219B, detail of watsoni sp. n. Neotropical geometrid moths 41

220A Downloaded by [Michigan State University] at 11:29 24 December 2014

222

Figs 220-224 Female genitalia. Ischnopteris. 220A, albiguttata; 220B, detail of albiguttata; 220C, signum of albiguttata; 221A, aurudaria; 221B, signum of aurudaria; 222, bifurcata sp. n.; 223, signum of brehmi sp. n.; 224A, catocalata; 224B, detail of catocalata. 42 Linda M. Pitkin

225 " Downloaded by [Michigan State University] at 11:29 24 December 2014

228 229 / 230A

Figs 225-230 Female genitalia. Ischnopteris. 225, chlorata; 226A, chloroclystata; 226B, signum of chloroclystata; 227A, miseliata; 227B, signum of miseliata; 228, multistrigata; 229, ochroprosthia; 230A, pronubata; 230B, detail of pronubata. Neotropical geometrid moths 43

232 B

231 233A

Downloaded by [Michigan State University] at 11:29 24 December 2014 •

**''*,

233B 234 235' 236

Figs 231-236 Female genitalia. Ischnopteris. 231, seriei; 232A, stenoptila; 232B, detail of stenoptila; 233A, xylinata typical form; 233B, detail of xylinata typical form; 234, detail of xylinata small form A; 235, 'I.' discolor; 236, 'I.' festiva. 44 Linda M. Pitkin

237 240

\r Downloaded by [Michigan State University] at 11:29 24 December 2014

241 244

Figs 237-244 Female genitalia. Stegotheca and Rucana. 237, Stegotheca albipennis; 238, Stegotheca amissa; 239, Stegotheca costiplaga; 240, Stegotheca cythereata; 241, Stegotheca phalangifera sp. n.; 242, Rucana abnormipalpis form A; 243, Rucana abnormipalpis form B; 244, Rucana brunneoviridis. Neotropical geometrid moths 45

- \

246 247 . 248 Downloaded by [Michigan State University] at 11:29 24 December 2014

249 250

Figs 245-250.245-249 Female genitalia of Rucana: 245, R. chaconi sp. n.; 246, R. degener; 247, R. fidelis; 248, R. marmorata; 249, R. mathani sp. n. 250, male abdomen (detail) of Ischnopteris bifinita. 46 Linda M. Pitkin

dimorphism adds to the difficulty of associating the sexes janzeni sp. nov. within a species. lata sp. nov. watsoni sp. nov. Biological notes Moths of Ischnopteris are known from elevations ranging from Species not placed to group 5 m to 2450 m. Information about their biology is limited, but albiguttata Warren, 1904a (Ischnopteris) larvae have been recorded on a wide range of hostplants in the praeluteata Warren, 1904b (Ischnopteris) following families: Acanthaceae, Annonaceae, Araliaceae, aurudaria (Schaus, 1901) (Ischnopteryx) Bignoniaceae, Celastraceae, Clusiaceae, Euphorbiaceae, bifurcata sp. nov. Fabaceae, Grossulariaceae, Lauraceae, Malpighiaceae, Mal- bipectinata sp. nov. vaceae, Melastomataceae, Menispermaceae, Monimiaceae, brehmi sp. nov. Myrtaceae, Ochnaceae, Piperaceae, Rosaceae, Rubiaceae, catocalata (Guenee, [1858]) (´Syrtodes) Sabiaceae, Sapindaceae, Solanaceae and Urticaceae (Brehm, prognata Dognin, 1923 (Ischnopteris) syn. nov. 2003; Janzen & Hallwachs' Caterpillars, pupae, butterflies & chlorata Hubner, [1823] (¨Ischnopteris) moths of the ACG, northwestern Costa Rica: http://janzen. chloroclystata chloroclystata (Guenee, [1858]) (´Syrtodes) sas.upenn.edu/; Roy A. Mora Arias records: http://www. chloroclystata juvencata (Guenee, [1858]) (´Syrtodes) nomen inbio.ac.cr/papers/Geometridae/Plantas hospedantes.html). dubium Host plants include sapling trees, woody shrubs, woody vines, fasciata sp. nov. and large perennial herbs (Daniel H. Janzen, pers. comm.). hirsuta sp. nov. For further mention see under the species bifinita, bryifera, latijuxta sp. nov. chavesi, chryses and brehmi. The larvae are known for those lemoulti sp. nov. species and their last instars are cryptic, twig or stick mimics miseliata (Guenee, [1858]) (´Syrtodes) with bark-like coloration, and with a swelling on each side of inconspicua Warren, 1907 (Ischnopteris) the body, towards the anterior end (particularly noticeable in multistrigata Warren, 1909 (Ischnopteris) Plate 1G). The top of the head is usually notched. Photographs obtortionis (Prout, 1928) (Ischnopterix) of a number of larvae are given by Janzen & Hallwachs (see ochroprosthia (Prout, 1929) (Ischnopterix) the URL above) and Brehm (2003); some are featured in this palmeri sp. nov. paper in Plate 1. pronubata (Felder & Rogenhofer, 1875) (Syrtodes) seriei (Giacomelli, 1911) (Ischnopteryx) Distribution stenoptila Warren, 1907 (Ischnopteris) Ischnopteris occurs widely thoughout the Neotropical Region, xylinata (Guenee, [1858]) (´Syrtodes) except in the temperate zones of the region: Chile and southern xylinata ockendeni (Prout, 1928) (Ischnopterix) syn. Argentina. nov.

Checklist of species Species excluded from Ischnopteris [placement unresolved, Ischnopteris fabiana- group see treatment at end of Ischnopteris] beckeri sp. nov. discolor-group bifinita (Walker, 1862) (Syrtodes) discolor Warren, 1904b (Ischnopteris) pexatata (Moschler, 1881) (Ïschnopteryx) conjugens Warren, 1905a (Ischnopteris) bryifera (Felder & Rogenhofer, 1875) (Syrtodes) fes ta Dognin, 1912 (Ischnopteris) velledata (Moschler, 1881) (Ïschnopteryx) festiva Warren, 1904b (Ischnopteris) Downloaded by [Michigan State University] at 11:29 24 December 2014 commixta (Warren, 1900) (Amblurodes) syn. nov. Other species excluded from Ischnopteris chlorophaearia (Walker, 1866) (Cidaria) callistrepta (Prout, 1928) (Ischnopterix) obfuscata Warren, 1909 (Ischnopteris) syn. nov. viriosa (Dognin, 1904) (Ischnopteryx) [transferred here to fabiana (Stoll, 1782) (Phal[aena] Geomet[ra]) Quillaca, as a synonym of viridifascia Warren] chlorosata (Hubner, [1825]) (Ïschnopterix) [Unnecess- ary replacement name for fabiana Stoll] apicemaculata Dognin, 1923 (Ischnopteris) syn. nov. Key to species choritis (Prout, 1929) (Ischnopterix) syn. nov. 1. Hind wing with orange region 2 illineata Warren, 1909 (Ischnopteris) - Hind wing without orange region 15 klagesi sp. nov. 2. Orange marking on hind wing more or less confined to a rostellaria (Felder & Rogenhofer, 1875) (Syrtodes?) large apical blotch (e.g. Figs 18-24) 3 - Orange marking extending across middle and base of wing Ischnopteris chryses-group (e.g. Figs 40, 57, 58, 61) 12 chavesi sp. nov. 3. Dorsum of male fore wing with only weak median tuft chryses (Druce, 1893) (Ischnopteryx) or without tuft; female genitalia with signum a short oval fassli sp. nov. or disc-shaped (Figs 221B, 223). [N.B. Female of palmeri hoffmani sp. nov. unknown.] 4 inornata sp. nov. - Dorsum of male fore wing with distinct median tuft; Neotropical geometrid moths 47

female genitalia with signum a moderately long oval projection of antrum pointed (Fig. 219B). Venezuela.... (Figs 216, 217A, 218A, 219A) 6 watsoni sp. nov. 4. Male retinaculum with small transparent central 12. Hind wing with dark region extended from dorsum up 'window'; female hind wing with dark brown streaks outer margin (Figs 40, 57, 58, 61) 13 extending from margin of orange region to near tornus, — Hind wing with dark region not extended from dorsum up clearly visible against greyish ground (Fig. 39), and fe- outer margin (Figs 33, 34) aurudaria male genitalia with base of signum much smaller than 13. Hind wing with distinct dark blotch at apex, or band from signum (Fig. 223) brehmi sp. nov. apex along outer margin (Figs 57, 58) pronubata - Male retinaculum with large transparent central ' window ' — Hind wing without distinct dark blotch or band at apex.. (similar to Fig. 92 of bifurcata); female hind wing with 14 or without distinct dark streaks extending from margin of 14. Hind wing with discal spot, but without a dark line along- orange region to near tornus, female genitalia with base of side dark blotch at tornus (Fig. 61) stenoptila signum almost as large as signum (Fig. 221B) or female — Hind wing without discal spot, but with a dark line along- unknown 5 side dark bar at tornus (Fig. 40) catocalata 5. Male genitalia with median process of juxta much shorter 15. Fore wing dark brown with contrasting pale costal band than main part of juxta (Fig. 113), female genitalia with (Fig. 55) ochroprosthia base of signum almost as large as signum (Fig. 221B)... — Fore wing not dark brown with contrasting pale costal aurudaria band 16 - Male genitalia with median process of juxta similar in 16. Fore wing with transverse slanting white band (Fig. 45) length to main part of juxta (Fig. 128). [Female unknown.] fasciata sp. nov. palmeri sp. nov. — Fore wing without transverse slanting white band .... 17 6. Hind wing with distinct dark spot at or very near apex 17. Hind wing shape: male with anal area between vein CuA1 (Figs 18,19) chavesi sp. nov. and dorsum expanded, forming lobe, usually prominent - Hind wing without distinct apical spot 7 (Fig. 4) but occasionally slight (Fig. 5); female with hind 7. Apical region of hind wing usually appearing dirty orange- wing fairly broad and outer margin projecting less at CuA1 yellow due to presence of fine brown speckles; with dark than normal for Ischnopteris (Fig. 6) bryifera brown lines and blotch leading from edge of this region — Hind wing shape not as in Figs 4—6, male without anal to near tornus. Male genitalia with very broad squar- lobe, female with outer margin projecting at CuA1 e.g. as ish processes of the anellus (Fig. 106B), female gen- in Figs 8, 10, 32 18 italia with digitate projection of antrum extending well 18. Hind wing of male with outer margin projecting, often beyond narrow lamella antevaginalis (Fig. 218B) strongly, around veins M1 to M3 (Fig. 2); female genitalia fassli sp. nov. with slanting ridge at postero-lateral edge of each of the - Apical region of hind wing pure orange-yellow without two depressions in lamella postvaginalis (Fig. 209A) ... speckles, and usually without lines and blotch between bifinita this region and tornus. Male genitalia with processes of — Hind wing of male with outer margin angled or projecting, the anellus not broad and squarish, female genitalia, where not strongly, at CuA1 (e.g. as in Figs 9,11,31); female gen- known, with projection of antrum not digitate if lamella italia either without depressions in lamella postvaginalis, antevaginalis is narrow 8 or with depressions that do not have slanting postero- 8. Fore wing long (male 22-23 mm). [Female unknown.].. lateral ridges 19 lata sp. nov. 19. Male with dark blotches as in Figs 46, 53: on median lobe - Fore wing length moderate (male 18-21 mm; female, of dorsum of fore wing and on anal lobe of hind wing, Downloaded by [Michigan State University] at 11:29 24 December 2014 where known, 21—23 mm) 9 pronounced against pale hind wing 20 9. Male retinaculum slightly enlarged and with small trans- — Male without pronounced dark blotches on median lobe of parent central 'window'; female, where known, with gen- dorsum of fore wing and on anal lobe of hind wing... 21 italia as in Figs 217, 219 10 20. Male genitalia with processes of anellus bifurcate - Male retinaculum strongly enlarged and with large trans- (Fig. 121) hirsuta sp. nov. parent central 'window'. [Female unknown.] — Male genitalia with processes of anellus not bifurcate (Fig. janzeni sp. nov. 126) obtortionis 10. Male genitalia with median process of juxta broad (Fig. 21. Fore wing of male with well-marked dark brown lines 107). [Female unknown.] Brazil hoffmani sp. nov. forming a series of longitudinal dashes, mainly in outer - Male genitalia with median process of juxta narrower half of wing (Fig. 51). Metathorax of both sexes dorsally (Figs 105, 11 1B); female genitalia, where known, as in with small dark longitudinal dash multistrigata Figs 217, 219. Not known from Brazil 11 — Fore wing of male without well-marked longitudinal 11. Male genitalia with digitate posterior process on apical dashes. Metathorax without dark dorsal longitudinal dash costal lobe of valva (Fig. 105); female genitalia with 22 posterior projection of antrum digitate (Fig. 217B). Not 22. Fore wing pattern with combination of: fine transverse known from Venezuela chryses streaks, small pale dark-edged wedge at base of wing, and - Male genitalia (Fig. 111A) without digitate process on well-marked dash at apex of wing (pale zigzag contrasting apical costal lobe of valva; female genitalia with posterior with dark brown marking, and with pale slanting subapical ifi Linda M. Pitkin

line, more pronounced than in most other species) (Figs long, or if disc-shaped then with base of signum situated 9, 10) fabiana at rim of signum 32 — Fore wing without the above combination of mark- 32. Male genitalia with gnathos process longer than gnathos ings 23 arms (Figs 100, 102, 103); female genitalia with base of 23. Male antenna strongly bipectinate. [Female unknown.].. signum situated at rim of signum; signum broad or disc- bipectinata sp. nov. shaped (Figs 213A, 215) 33 — Antenna of both sexes either not bipectinate, or male - Male genitalia with gnathos process not longer than bipectinate but with very short rami 24 gnathos arms (Fig. 98); female: signum broad, with base 24. Male antenna bipectinate but with very short rami, or situated in from rim of signum (Fig. 211) serrate; head of both sexes with dark brown dorsal line.. chlorophaearia xylinata 33. Male genitalia with gnathos process narrow and rod-like — Antenna of both sexes simple filiform; head without dark (Fig. 100); female genitalia with signum broad and dis- brown dorsal line 25 tinctly triangular (Fig. 213A) illineata 25. Hind wing pale except for margin, contrasting with fore - Male genitalia with gnathos process tongue-shaped wing which is uniformly dark or has dark areas (Figs 59, (Figs 102, 103); female: signum disc-shaped or only 60) seriei weakly subtriangular (Fig. 215) rostellaria — Hind wing not much paler than fore wing 26 34. Male genitalia with processes of the anellus bifurcate 26. Hind wing greyish brown, with diffuse patch paler brown (Figs 114,115,118); female genitalia with lamella anteva- and warmer in hue, from apex of wing to dark postmedial ginalis as a broad band (Figs 222, 225) 35 line, and as narrower band along outer margin of wing - Male genitalia with processes of the anellus not bifurc- (Figs 30-32) albiguttata ate; female genitalia (where known) with lamella anteva- — Hind wing without the above combination of markings . ginalis as a narrow or bilobed band (Figs 226A, 227A).. 27 36 27. Fore wing with dark brown and pale greenish speckles, 35. Male with processes of anellus bifurcate with prongs well and very fine transverse streaks, not forming any well- apart (Fig. 118); female genitalia with signum a fairly long defined blotches, and with three wavy whitish transverse oval (Fig. 225) chlorata lines (Fig. 1); male genitalia with uncus strongly sinuous - Male with processes of anellus bifurcate with prongs close (Fig. 95) beckeri sp. nov. together (Figs 114, 115); female genitalia with signum a — Fore wing without the above combination of markings; short oval (Fig. 222) bifurcata sp. nov. male genitalia with uncus not strongly sinuous (Fig. 95). 36. Processes of anellus each with large basal region extend- 28 ing over anellus and with smaller spine at posterior end 28. Dorsum of male fore wing with distinct median tuft.... of that region (Fig. 119); female genitalia with base of inornata sp. nov. signum almost as large as signum (Fig. 226B) — Dorsum of male fore wing without distinct median chloroclystata tuft 29 - Processes of anellus without accompanying spine; female 29. Male genitalia with gnathos as a simple loop without pro- genitalia (where known) with base of signum much smal- cess latijuxta sp. nov. ler than signum proper 37 — Male genitalia: gnathos with one or two median processes. 37. Male genitalia with uncus tapered at apex (Fig. 124); fe- 30 male genitalia with elongate pointed signum (Fig. 227B) 30. Male genitalia with uncus strongly hood-shaped and usu- miseliata ally moderately broad, with large tuft of ventral setae (Figs - Male genitalia with uncus narrowing abruptly to small Downloaded by [Michigan State University] at 11:29 24 December 2014 98,100-103). Female genitalia with signum situated well rod-like apex (Fig. 123). [Female unknown.] away from anterior end of corpus bursae (Figs 211, 213, lemoulti sp. nov. 215), or signum with base situated well away from rim of signum (Fig. 214); signum not a long oval 31 Ischnopteris fabiana species group — Male genitalia with uncus narrow, at least apically, and not The wing pattern of the fabiana-group is fairly typical of that of strongly hood-shaped (Figs 114,115,118,119,123,124). Ischnopteris in general, but the hind wing never has an orange Female genitalia with signum situated less well away from region. The fore wing is predominantly dark or mid brown, anterior end of corpus bursae (Figs 222, 226A, 227A), or often mottled with greenish or paler brown flecks, and often in signum a fairly long oval (Fig. 225), or unknown; base of the female with a roughly diamond-shaped central area of con- signum at or near rim of signum 34 trasting colour, usually paler but sometimes darker. The fore 31. Male genitalia with pair of postero-lateral lobes on uncus, wing of the male does not have a white blotch near the mid- resembling ears (Fig. 101); female genitalia with signum point of the costa, as is sometimes seen in other Ischnopteris disc-shaped and base of signum almost central (Fig. 214) species, but some specimens of illineata have other whitish klagesi sp. nov. blotches. The dorsum of the male fore wing does not have — Male genitalia: uncus without pair of postero-lateral lobes; the distinct median tuft of the chryses-group and a few other female: signum subtriangular to triangular, or broader than Ischnopteris species. Neotropical geometrid moths 49

The male genitalia have a definitive character of the un- apex. Aedeagus with or without a few extremely minute spin- cus: this is a more strongly hood-shaped structure than in other ules near posterior end; vesica with row of spines extending Ischnopteris species, and it is often large and broad, with vent- along short caecum. ral setae more or less merged as a single large tuft; also, the uncus often has a pair of minute digitate apical projections, FEMALE GENITALIA. Unknown. one lying over the other. The gnathos usually has a single median process, and only bifinita has two processes (which Diagnosis are not hook-tipped as in many other Ischnopteris species). In The fore wing pattern of beckeri, as described above for males, the valva the longitudinal ridge is sinuous (except in beckeri). is fairly distinctive, but the species is better defined by charac- The median process of the juxta is spinulose over much of ters of the male genitalia. Like other members of the fabiana- its length or just apically. The aedeagus usually has spinules, group, beckeri has a distinctly hood-shaped uncus, but it is the which are situated near the posterior tongue, less commonly only member of this group to have the apical costal process on the tongue itself. The vesica has a spinose sclerite (but of the valva narrow and sharply pointed, and the longitudinal not long as in the chryses-group and some other Ischnopteris ridge of the valva not strongly sinuous. The strongly sinuous species); the sclerite is usually on a short or very short caecum, shape of the uncus is diagnostic and putatively autapomorphic but occasionally not on a distinct caecum. for beckeri. In the female genitalia, the lamella postvaginalis often has a pair of depressions, varying from pronounced to weak and Biological notes indistinct, or absent; the antrum does not have a posterior pro- Moths have been found at an altitude of 1300 m, in April. jection (except in klagesi); the corpus bursae sometimes has a differentiated narrower anterior part. The signum is often broad Distribution and sometimes subtriangular; except in klagesi, the signum is Known only from the type locality in eastern Brazil. situated well away from the anterior end of the corpus bursae (outside of the fabiana-group, xylinata and multistrigata are the only species comparable in this respect). Ischnopteris bifinita (Walker) Plate 1A-C; Figs 2, 3, 96,155, 209, 250 Ischnopteris beckeri sp. nov. Syrtodes? bifinita Walker, 1862:1374. Holotype , [BRAZIL] Figs 1, 95,154 (BMNH) [examined]. Material examined Ischnopteryx pexatata Moschler, 1881: 412, pl. 18, fig. 24. 2 (genitalia preparations of both) LECTOTYPE , SURINAM (MNHU) here designated Holotype , Brazil: M[inas] G[erais], Carac¸a, 1300 m, 14- [examined]. [Synonymy cited by Parsons et al., in Scoble, 15.iv.1998 (Becker) (genitalia slide LMP 454; VOB). 1999: 519.] Paratype, 1 , data as holotype but 1-2.iv.1992 (VOB). Ischnopteris bifinita (Walker); Parsons etal., in Scoble, 1999: 519. Description ADULTS (Fig. 1). Fore wing length: , 19 mm. Wing pat- Material examined tern (only male known): fore wing with dark brown and pale 36 , 15 (including 5 , 7 genitalia preparations) (AMNH, greenish markings as speckles and very fine transverse streaks, BMNH, CMNH, DHJ, INBio, USNM, ZMUC, ZSM). not forming any well-defined blotches (and not with a small pale basal wedge as is present in fabiana); with three wavy Holotype (bifinita), Brazil: Para (genitalia slide Geom: whitish transverse lines; hind wing brown with darker but 21095, BMNH). Downloaded by [Michigan State University] at 11:29 24 December 2014 sometimes indistinct postmedial line, and with diffuse, dark Lectotype (pexatata), Surinam: no further data (MNHU). submarginal band. Under-side of hind wing with dark discal spot well marked. Male retinaculum moderately enlarged and Localities of additional material. Guatemala: Cayuga; with transparent central 'window'. Hind wing of male with Quirigua. Belize: no further data; Rio Temash. Nicaragua: anal margin not expanded and without pronounced hair-scale Chontales; 10 miles NW of Eden, Pis Pis River, Great tuft. Falls. Costa Rica: Alajuela Province, Area de Conserva- cion Guanacaste, Rincon Rainforest, Estaci´on Caribe; data MALE GENITALIA (Figs 95, 154). Uncus distinctly hood- as before, Rincon Rainforest, Camino Rio Francia; province shaped, strongly sinuous-sided, apex without processes. and area as before, Sector San Cristobal, Rio Blanco Abajo; Gnathos with one median process, curved, narrow and not province, area and sector as before, Quebrada Cementerio [all tongue-shaped, not longer than gnathos arms. Valva with long- Alajuela records based on digital photographs, DHJ]; [Cartago itudinal ridge not strongly sinuous, and with apical costal pro- Province,] Juan Vinas; Punta[renas] Prov[ince], [Carara Bio- cess narrow and sharply pointed. Processes of anellus claw- logical Reserve,] Est[acion] Q[uebrada] Bonita; Puntarenas like, smoothly curved, with narrow and pointed apical part. Province, Osa Peninsula, Corcovado National Park, Sirena; Median process of juxta long and slender (about twice length Punt[arenas] Prov[ince], 35 km S. of Palmar Norte, Fila Es- of main part of juxta), almost straight, with spinules towards quinas; Santa Clara Vall[ey]. Panama: Chiriqui; La Chorrera. 50 Linda M. Pitkin Downloaded by [Michigan State University] at 11:29 24 December 2014

Plate 1 Larvae (last instar) of Ischnopteris. A-C, bifinita 00-SRNP-14209: A, lateral entire (DHJ55615); B, lateral rear (DHJ55617); C, front (DHJ55618); D, bryifera 00-SRNP-4413 lateral entire (DHJ51043); E, bryifera 01-SRNP-5456 lateral entire (DHJ63418); F-G, chavesi sp. n. 02-SRNP-9835: F, dorsal entire (DHJ67792); G, dorsal front (DHJ67795); H-I, chavesi sp. n. 99-SRNP-1126: H, lateral entire (DHJ50490); I, dorsal front (DHJ50488). All photographs by Daniel Janzen (© D. Janzen & W. Hallwachs).

Colombia: near Cali, 'Juntas'; Don Amo. Venezuela: no fur- St Laurent du Maroni. Ecuador: Esmeraldas, San Mateo; Para- ther data; Palma Sola; San Esteban. Guyana: no further data; mba. Brazil: Minas Gerais, Teofilo Otoni, San Jacintho Valley; Bartica District, Kartabo; confluence of Oroneque and New Para; Rio Purus, Hyutanahan; Santa Catarina, Nova Bremen. Rivers; Potaro; Potaro River. French Guiana: no further data; Bolivia: S[anta] Cruz de la Sierra. Neotropical geometrid moths 51

Description Remarks ADULTS (Figs 2, 3). Fore wing length: , 21-25 mm; , 22- The type series of pexatata consists of two male specimens 25 mm. Wing pattern: fore wing mottled dark to mid brown, in the MNHU, both labelled 'Type' but not conspecific. I female with roughly diamond-shaped central area paler, darker, have chosen to designate as lectotype the specimen figured in or more-or-less concolorous and marked out only by dark lines Moschler's original description, which has a reference to the around it; hind wing of male plain dark brown, female mid original description and figure on its type label. The identity to dark brown with darker but indistinct postmedial line and of the lectotype is bifinita, thus the status of pexatata remains diffuse submarginal band. Male retinaculum enlarged and with unchanged, but the identity of the other specimen (now the transparent central 'window' usually small. Hind wing of male paralectotype) is chlorophaearia. with outer margin projecting, often strongly, around veins M1 to M3 (instead of angled or projecting, not strongly, at CuA 1, as Biological notes is usual for the genus); anal margin strongly expanded forming Ischnopteris bifinita is primarily a lowland, to low intermedi- distinct flap, with pronounced hair-scale tuft; hind wing with ate elevation, species. Moths have been examined from alti- extensive patches of hairs in rows between veins; female with tudes of 50-610 m; Janzen & Hallwachs (http://janzen.sas. fairly broad hind wing. Male abdomen (Fig. 250): segment 4 upenn.edu/caterpillars/database.lasso) record also 700 m; re- with strong constriction in lightly sclerotized areas. cords of elevations higher than 700 m are unconfirmed). Moths examined were collected in all months of the year except March MALE GENITALIA (Figs 96, 155). Uncus strongly hood- and October. Larvae (Plate1A-C) have been reared by Janzen shaped, rounded, very large and broad, with pair of dorsal tufts & Hallwachs (see the URL above for further details; vouchers: prominent and well defined; with pair of tiny digitate ap- 00-SRNP-11398, 11879, 01-SRNP-4237), from hostplants: ical projections, one lying over the other, upper one often spatulate, both projections bent over and directed anteriorly. Clusiaceae: Vismia baccifera (L.) Triana & Planch. Gnathos with two median processes: short, tapered and diver- Monimiaceae: Siparuna andina (Tul.) A. Dc. ging, without hooked tip. Valva with strongly sinuous longitud- Urticaceae: Myriocarpa longipes Liebm. inal ridge and with low, rounded, apical costal lobe. Processes of anellus claw-like, evenly curved and tapered. Median pro- The following records from Janzen & Hallwachs' rearings of cess of juxta moderately short (similar in length to main part Ischnopteris may also be of bifinita, but this is unconfirmed: of juxta), slightly curved to right (in ventral view) or almost Euphorbiaceae: Acalypha diversifolia Jacq. straight, minutely spinulose over much of its length, with lar- Fabaceae: Lonchocarpus guatemalensis Benth. ger spines at apex. Aedeagus with spinulose region situated Malpighiaceae: Heteropterys obovata (Small) Cuatrec. & adjacent to posterior tongue, which has no spinules; vesica Croat with spinose sclerite on very short caecum. Piperaceae: Piper auritum Kunth Sapindaceae: Cupania glabra Sw. FEMALE GENITALIA (Figs 209A, B). Lamella postvaginalis with pair of depressions, each with slanting postero-lateral Distribution ridge. Lamella antevaginalis a narrow band, curved or sinuous Central and South America from Guatemala to Brazil and but without distinct lobes. Antrum without posterior projec- Bolivia. tion. Corpus bursae with narrower anterior part; signum broad, subtriangular, situated in bulbous part of corpus bursae, well away from anterior end. Ischnopteris bryifera (Felder & Rogen hofer) Plate 1D-E; Figs 4-6, 97,156, 210 Downloaded by [Michigan State University] at 11:29 24 December 2014 Syrtodes bryifera Felder & Rogenhofer, 1875: pl. 131, fig. 28. Diagnosis Holotype , FRENCH GUIANA (BMNH) [examined]. The outer margin of the hind wing, projecting as described Ischnopteryx velledata Moschler, 1882: 413, pl. 18, fig. above, usually gives the male moths of bifinita a distinctive 25. Holotype , SURINAM (MNHU) [digital photograph shape, and is autapomorphic for the species. Females, having examined]. [Synonymy cited by Parsons et al., in Scoble broad hind wings, might sometimes be confused with those of (1999).] bryifera except that the latter have a slightly less projecting Amblurodes commixta Warren, 1900: 201. Syn. nov. outer margin (compare Fig. 3 with Fig. 6). Another similarity LECTOTYPE , [ECUADOR] (BMNH), here desig- with bryifera is in the male abdomen, the constriction in the nated [examined]. [Treated as a subspecies of Ischnopteris sclerotized areas of segment 4. The male genitalia resemble bryifera by Parsons et al., in Scoble (1999).] those of bryifera in the very large, broad, uncus, but bifinita Ischnopteris bryifera (Felder & Rogenhofer); Pitkin et al., is the only member of the fabiana-group to have two gnathos 1996: 135. processes. The female genitalia are characteristic in having a narrow lamella antevaginalis, exposing to clear view the Material examined slanting postero-lateral ridge at each of the depressions in the 85 , 43 (including 3 , 10 genitalia preparations) (BMNH, lamella postvaginalis. CMNH, DHJ, INBio, SMNS, USNM). 52 Linda M. Pitkin

Holotype (bryifera), French Guiana: no further data (gen- projections, one lying over the other, upper one usually spat- italia slide Geom: 21097, BMNH). ulate, both projections sometimes bent over at right angles to Holotype (velledata), Surinam: no further data (MNHU) uncus or directed anteriorly, situated between pair of short in- [digital photograph]. conspicuous lobes. Gnathos with one median process. Valva Lectotype (commixta), Ecuador: Paramba, 1070 m ('3500 with strongly sinuous longitudinal ridge and with apical costal ft'), vi.[18]97 (Rosenberg) (genitalia slide Geom: 21008, lobe. Processes of anellus broad in basal half, with angular BMNH). edge. Median process of juxta similar in length to main part of juxta or slightly longer, straight, minutely spinulose over Localities of additional material. Costa Rica: no further much of its length, with larger spines at apex. Aedeagus with data; Alajuela Province, Area de Conservacion Guanacaste, spinulose region situated adjacent to posterior tongue, which Sector San Cristobal, Quebrada Cementerio (voucher: 99- has no spinules; vesica with sclerite, spinose and not rounded SRNP-4413) [digital photograph, DHJ]; Alajuela Province, at tip, on very short caecum. 8 km S. of Santa Cecilia, Estacion Pitilla; Alajuela Province, 5 km N. of Col. Palmarena, San Ramon For[est] Res[erve], Rio FEMALE GENITALIA (Figs 210A, B). Lamella postvaginalis San Lorencito; [Cartago Province,] Juan Vinas; Guanacaste with a pair of shallow rounded depressions, mainly covered by Province, Rincon National Park, 8 km E. of Casetilla; Heredia the lamella antevaginalis. Lamella antevaginalis moderately Province, Puerto Viejo de Sarapiqui, La Selva Biol. Sta.; broad, forming a pair of rounded lobes separated medially by Tuis. Panama: Volcan de Chiriqui. ´Trinidad: no further a V-shaped indentation. Antrum without conspicuous posterior data; Caparo. Colombia: [Magdalena Province,] Cacagualito; projection; antrum longer than in bifinita (compare Figs 210 Choco Province, Condoto. Venezuela: no further data. and 209). Corpus bursae with narrower anterior part; signum Guyana: no further data; Amazon-Courantyne divide, head broad, weakly subtriangular shape but usually bilobed, situated of Oronoque River; Kartabo; New River; Rio Demerara. in bulbous part of corpus bursae, well away from anterior end. Surinam: Surinam River. French Guiana: no further data; St Jean du Maroni; St Laurent du Maroni. Ecuador: Bol´ıvar Diagnosis Province; [Bol´ıvar Province,] Balzapamba; Corondalet; The shape of the hind wing in males of bryifera, with expan- Esmeraldas, 5 km E. [of] Alto Tambo; Paramba; Sarayacu; ded anal area and lobe, is unique in the genus and is highly Zamora; Zamora-Chinchipe, Parque Nacional Podocarpus, distinctive, even in specimens with only slight development [Rio] Bombuscara. Peru: [Loreto,] 'Amazones, Cavallo- of the lobe (see Figs 4 and 5). The conspicuous pale greenish Cocho'; Carabaya, R. Huacamayo, La Union; E. Peru, curved line across the fore wing is also distinctive. Females Chanchamayo; Huanuco, Pozuzo. ´Brazil: Santa Catarina. are less easily recognised, but the shape of the hind wing (described above) is subtly different from that of other species. The constriction in the sclerotized areas of segment 4 of the Description male abdomen is similar to that in bifinita (Fig. 250). In the ADULTS (Figs 4-6). Fore wing length: , 21-24 mm; , 22- male genitalia the uncus is very large and broad, as in bifin- 26 mm. Wing pattern: fore wing mottled dark brown, male ita, but that species differs in having two gnathos processes. with pale grey-green markings, most prominent of which is a The female genitalia of bryifera are characterized by a com- curved transverse line, situated medially, female with roughly bination of features, the V-shaped indentation of the lamella diamond-shaped central area sometimes paler (straw-coloured, antevaginalis, and the broad, weakly subtriangular but usually whitish or grey), or darker, but often more-or-less concolorous bilobed, signum. and central area marked out only by dark lines around it; hind wing of male plain dark brown, female dark to mid brown with Variation darker but indistinct postmedial line and diffuse submarginal Some of the male specimens examined from Costa Rica, band. Male retinaculum slightly or not enlarged and with trans- Downloaded by [Michigan State University] at 11:29 24 December 2014 Colombia, and Ecuador, have only a slight lobe at the anal parent central 'window' very weak and inconspicuous. Hind angle of the hind wing. Specimens of this form are also an wing of male modified with anal area between vein CuA1 and exceptionally dark chocolate-brown colour. dorsum expanded, usually forming a prominent lobe at anal angle although lobe slight in some specimens (see 'Variation'); Biological notes presence of lobe producing indentation to outer margin of wing Moths of bryifera have been found at altitudes of 40 m to at CuA1 (instead of outer margin projecting at that point, as is 1120 m; in Costa Rica it appears to be primarily a low- usual for the genus); anal margin strongly expanded forming land, to low intermediate elevation, species, with only one distinct flap, with pronounced hair-scale tuft; hind wing with record above 730 m. Moths examined were collected in all extensive patches of hairs in rows between veins. Female also months of the year except May, July, and September. Lar- with outer margin of hind wing projecting less at CuA1 than vae (Plate 1 D-E) have been reared by Janzen & Hallwachs normal for the genus, and with hind wing fairly broad. Male (see http://janzen.sas.upenn.edu/caterpillars/database.lasso for abdomen: segment 4 with strong constriction in areas of light further details; vouchers: 99-SRNP-4413, 00-SRNP-9961, sclerotization. 11465, 12929, 22201, 01-SRNP-612, 5456, 02-SRNP-7295, 7652, 7691), from hostplants: MALE GENITALIA (Figs 97, 156). Uncus strongly hood- shaped, oval, very large and broad, with pair of dorsal tufts Annonaceae: Annona sp. prominent and well defined; with pair of tiny digitate apical Araliaceae Dendropanax arboreus (L.) Decne. & Planch. Neotropical geometrid moths 53

Malvaceae: Sida sp. Description Melastomataceae: Conostegia xalapensis (Bonpland) D. ADULTS (Figs 7, 8). Fore wing length: , 18-22 mm; , 20- Don 23 mm. Wing pattern: fore wing of male mottled mid to dark Monimiaceae: Siparuna andina (Tul.) A. Dc. brown with small paler flecks; sometimes with pale blotches: a Ochnaceae: Cespedesia macrophylla Seem. basal wedge, and one to two small subapical markings; darker Piperaceae: Piper sp. antemedial line and zigzag postmedial line commonly present Sapindaceae: Cupania glabra Sw. but rather indistinct, with black dash at dorsum between those Solanaceae: Cestrum sp. two lines often present in male and strongly marked in female; Urticaceae: Myriocarpa longipes Liebm. female predominantly pale brown, speckled and marked with dark brown, with dark antemedial and postmedial lines en- The following records from Janzen & Hallwachs' rearings of closing roughly diamond-shaped central area. Hind wing of Ischnopteris may also be of bifinita, but this is unconfirmed: both sexes more or less concolorous with fore wing, in male ususally plain and only occasionally with some diffuse band- Acanthaceae: Iresine sp. ing, and in female with darker, dull brown, postmedial line Rubiaceae: Psychotria sp. and submarginal band. Under-side of wings sometimes with Solanaceae: Solanum sp. more distinct postmedial line. Male retinaculum enlarged and with moderate-sized transparent central 'window'. Hind wing Distribution of male with anal margin expanded forming distinct flap, with Central and South America from Costa Rica to Brazil. I can- slight hair-scale tuft. not confirm a record from Mexico by Delf´ın-Gonzalez & MALE GENITALIA (Figs 98, 157). Uncus strongly hood- Beutelspacher (1986:453), and I consider it more likely to shaped, broad; with pair of tiny digitate apical projections, be a misidentification of rostellaria. one lying over the other, both projections either at right angles to uncus or bent over and directed anteriorly. Gnathos with Ischnopteris chlorophaearia (Walker) one median process, narrow and not tongue-shaped, similar in Figs 7, 8, 98,157, 211 length to gnathos arms or shorter. Valva with sinuous longitudinal ridge and with apical costal lobe. Processes of anel- Cidaria chlorophaearia Walker, 1866: 1690. LECTO- lus claw-like, evenly curved and tapered. Median process of TYPE , [COLOMBIA]: (BMNH), here designated juxta varying in length from nearly twice to nearly three times [examined]. length of main part of juxta; straight or almost straight, pointed, Ischnopteris obfuscata Warren, 1909: 103. LECTOTYPE minutely spinulose over much of its length, with larger spines , [BRAZIL] (BMNH), here designated [examined]. medially, towards apex. Aedeagus with one or a few triangu- Syn. nov. lar spinules, occasionally weak or absent, on posterior tongue, Ischnopteris chlorophaearia (Walker); Parsons et al., in and with group of spinules situated opposite and subapically Scoble, 1999: 519. to tongue; vesica with spinose sclerite on short caecum.

Material examined FEMALE GENITALIA (Fig. 211). Lamella postvaginalis with 30 , 3 (including 17 , 3 genitalia preparations) (BMNH, a pair of shallow rounded depressions. Lamella antevaginalis INBio, MNHU, USNM, ZSM). a band neither particularly narrow or broad, forming a pair of rounded lobes, pronounced or slight, separated medially Lectotype (chlorophaearia), Colombia: Bogota (´Birchall) by a broad indentation. Antrum without conspicuous posterior (genitalia slide Geom: 21072, BMNH). projection. Corpus bursae with or without slightly narrower Downloaded by [Michigan State University] at 11:29 24 December 2014 Lectotype (obfuscata), Brazil: [Amazonas] Upper anterior part; signum much broader than long, varying in shape Amazons, Fonte Boa, vii. 1906 (Klages) (genitalia slide but sometimes subtriangular, base of signum situated in from Geom: 21067, BMNH). Paralectotypes, 4 (obfuscata), rim of signum, signum situated in posterior half of corpus data as lectotype but vii, viii, xi. 1906 (BMNH). [Dates of bursae. collection given here are from label data, but Walker (1866) cited viii for all the syntypes.] Diagnosis Localities of additional material. Costa Rica: Moths of chlorophaearia are not readily distinguishable by [Puntarenas Province,] Osa Penin[sula], Corcovado Nat[ional] external features but the genitalia have some definitive features. P[ar]k, Sirena. Panama: Bugaba; La Chorrera; Volcan In the male, the median process of the juxta is longer than de Chiriqui; Rio Trinidad. Colombia: Bogota.´ Venezuela: in other members of the fabiana-group except fabiana itself, Cucuta; W Caracas, Carabobo, Bejuma, Casa Maria. Suri- which has a narrower uncus with putatively autapomorphic nam: no further data (paralectotype of Ischnopteryx pexatata features of a dorsal, basal, single patch of hair-scales and a Moschler, see 'remarks' under bifinita); Paramaribo. Ecuador: collar-like plate. In the female, the signum is broad like that of R[io] Anzu; Corondalet; S[an] Javier. Peru: 'Amazones, bifinita and several other members of the fabiana-group, but Cavallo-Cocho'. Brazil: [Amazonas,] Fonte Boa. Bolivia: E. chlorophaearia differs from those species in having the base Bolivia, Buenavista; [Cochabamba,] Chapare. of the signum situated in from the rim of the signum. 54 Linda M. Pitkin

Specimens of obfuscata were found to have a longer juxta with dark brown subapical band and postmedial line (some- process, and the anellar processes tend to be more slender, but times faint). Male retinaculum enlarged and with moderately as variation in both those characters ranges continuously across large transparent central 'window'. Hind wing of male with obfuscata and chlorophaearia, the species are treated here as anal margin little expanded, without pronounced hair-scale synonymous. tuft.

Biological notes MALE GENITALIA (Figs 99, 158). Uncus distinctly hood- Moths have been found at altitudes of 400 m to 920 m, in March shaped but not broad; with flattened or slanting top forming to August, November and December. apical projection, and with very weak division at extreme tip of that into pair of tiny projections at extreme tip, one above the Distribution other; base of uncus with dorsal single patch of hair-scales (de- Central and South America from southern Costa Rica to Brazil ciduous but scale bases always remain), and with a collar-like and Bolivia. plate situated between that patch and the socii. Gnathos with one median process, narrow or tapered and not tongue-shaped, similar in length to gnathos arms. Valva with strongly sinuous Ischnopteris fabiana (Stoll) longitudinal ridge and with angular apical costal lobe. Pro- Figs 9,10, 99,158, 212 cesses of anellus claw-like, evenly curved, tapered and pointed. Phal[aena] Geomet[ra] fabiana Stoll, 1782: 227, 249 Median process of juxta a long, straight, narrow, pointed blade (index), pl. 397, fig. H. Syntype(s), SURINAM [not (approximately twice length of main part of juxta), spinulose examined]. in apical half or one-third. Aedeagus with spinulose region Ischnopterix chlorosata Hubner, [1825]: 332. Syntype(s), situated adjacent to posterior tongue, which has no spinules; SURINAM. [Unnecessary replacement name for fabiana vesica with sclerite, spinose at tip and not rounded, on short Stoll.] caecum. Ischnopteris apicemaculata Dognin, 1923: 18. Holotype , [BRAZIL] (USNM) [examined]. Syn. nov. FEMALE GENITALIA (Fig. 212). Lamella postvaginalis with Ischnopterix choritis Prout, 1929: 68. Holotype , PERU pair of rounded depressions small and weak or absent. Lamella antevaginalis forming pair of rounded lobes separated medi- (BMNH), [examined]. Syn. nov. ally by strong V-shaped indentation. Antrum without posterior Ischnopteris fabiana (Stoll); Rindge, 1983: 226. projection. Corpus bursae without differentiated narrower anterior part; signum very small, weakly developed, situated well Material examined away from anterior end of corpus bursae. 10 , 3 (including 5 , 3 genitalia preparations) (AMNH, BMNH, CMNH, USNM). Diagnosis Holotype (apicemaculata), Brazil: Amazon[as], Sao Paulo The wing pattern of fabiana described above is fairly distinc- de Olivenc¸a, xi-xii (Fassl) (Type No. 32553, genitalia slide tive. Dark males may resemble bryifera (both species have a 60136, USNM). pale or olive transverse line medially), but the pale wedge at Holotype (choritis), Peru: SE. Peru, Carabaya, Rio Huaca- the base of the wing in fabiana is not present in bryifera, and mayo, 950 m ('3100 ft'), vi. 1904 (Ockenden) (genitalia the latter species has a modified hind wing shape. The male slide Geom: 21016, BMNH). genitalia of fabiana have unique characters, notably the single basal patch of hair-scales and the collar-like plate of the uncus, Localities of additional material. Colombia: upper ('Ob. ') and the angular apical costal lobe of the valva, and in the Rio Negro. Guyana: Upper Kutari R. French Guiana: St female the signum is the smallest known in Ischnopteris (with Downloaded by [Michigan State University] at 11:29 24 December 2014 Jean du Maroni. Ecuador: R[io] Anzu; Sarayacu. Peru: the exception of a single female unidentified to species). 'Amazones, Cavallo-Cocho'; Carabaya, R[io] Huacamayo; [Huanuco,] Pozuzo. ´Brazil: [Amazonas] ('Upp. Amazon'), Biological notes Fonte Boa; Amazonas, Rio Manes; Rio Purus, Hyuatanahan; Moths have been found at altitudes of 300-950 m, in February, [Amazonas,] Rio Purus, Nova Olinda; Rondonia, 62 km S. April to June, July, and November to December. Ariquemes, Fazenda Rancho Grande.

Distribution Description South America from Colombia eastwards to French Guiana ADULTS (Figs 9, 10). Fore wing length: , 21-24 mm; and southwards to Peru. , 24-25 mm. Wing pattern: fore wing of both sexes dark and mid brown mottled in very fine transverse streaks; with some Ischnopteris illineata Warren olive and whitish markings including: curved or almost straight Figs 11-13,100,159, 213 transverse line situated medially (broken up in female), small pale dark-edged wedge at base of wing, and well-marked dash Ischnopteris illineata Warren, 1909: 101. Holotype , at apex of wing (pale zigzag contrasting with dark brown mark- ECUADOR (BMNH) [examined]. ing, and with pale slanting subapical line, more pronounced Ischnopteris illineata ab. trimaculata Warren, 1909: 101. than in most other species); hind wing mid to dark brown, [Infrasubspecific name.] Neotropical geometrid moths 55

Material examined than the females) can sometimes be a useful pointer, but it 11 , 8 (including 5 , 7 genitalia preparations) (AMNH, is not conclusive. The definitive features of illineata are in BMNH, ZSM). the genitalia, chiefly those of the male, with rod-like, almost straight, gnathos process. The female genitalia are similar to Holotype Ecuador: West Ecuador, [Los Rios,] Quevedo (v. those of bifinita in having a triangular signum, but the paired Buchwald) (genitalia slide Geom: 20996, BMNH). depressions of the lamella postvaginalis are rounded in illineata whereas those in bifinita have a slanting ridge at the Localities of additional material. Colombia: Magdalena postero-lateral edge. River; Magdalena, Sierra Nevada de Santa Marta. French Guiana: Nouveau Chantier. Ecuador: Esmeraldas, San Variation Mateo; Guayas: Bucay; 4km SW. [of] Bucay, Hacienda Two males from Peru, probably belonging to illineata, have San Joaquin; Imbabura: Perucho; N. Perucho, near Otavalo; some slight differences in characters from those given above. Los Rios Prov[ince], [near?] Bahahoyo, Rio de las Juntas, They have a fore wing length of 19—20 mm, and the me- La Chima; N. Ecuador, Montecristi; W. Ecuador, [Los dian process of the gnathos is slightly broader and less obvi- Rios Province,] Quevedo; [El Oro Province,] Zaruma. Peru: ously rod-like than in typical illineata, but not broad enough Hacienda Taulis. to appear tongue-like as in rostellaria. A further male from Localities of further atypical material (see 'Variation'). 2 Peru is not identified to species but mentioned here because it d (genitalia preparations of both) (BMNH). Ecuador: Loja. has some illineata-like characteristics, particularly in gnathos Peru: [Cajamarca,] R. Tabaconas. shape. However, this specimen is larger (fore wing length 20 mm), with a longer median process of the juxta (similar Description to that of chlorophaearia), and does not have a spinulose re- ADULTS (Figs 11-13). Fore wing length: , 16-18 mm; , gion opposite the posterior tongue of the aedeagus. 21—25 mm. Wingpattern: fore wing mottled dark to mid brown; male without conspicuous antemedial and postmedial lines, but sometimes with small, paler but inconspicuous blotch towards Biological notes anal angle, or infrequently with that blotch and some other Moths have been found at altitudes of 250 m to 2000 m, in markings conspicuously whitish; female with patchy markings March, May to July, and December. over extensive area of fore wing, this area often paler than base and apex of wing but occasionally darker. Hind wing Distribution mid to dark brown with only faint markings. Male retinaculum South America from Colombia, French Guiana, Ecuador and enlarged and with small to moderately large transparent central Peru. 'window'. Hind wing of male with anal margin not strongly expanded and with hair-scale tuft usually slight. Ischnopteris klagesi sp. nov. Figs 14,15,101,160, 214 MALE GENITALIA (Figs 100, 159). Uncus strongly hood- shaped and moderately elongate; with pair of tiny digitate Material examined apical projections, one lying over the other, upper one pointed, 1 , 1 (genitalia preparations of both) both projections at right angles to uncus, or slightly bent over Holotype , Brazil: Rio Purus, Hyutanahan, iii. 1922 (Klages) and directed anteriorly. Gnathos with one median process, nar- (genitalia slide LMP 423; CMNH). row rod-like and almost straight, but with pointed, hooked tip; Paratype, 1 , data as holotype but iv.1922 (CMNH). process longer than gnathos arms. Valva with strongly sinuous longitudinal ridge and with apical costal lobe. Processes of Additional specimen excluded from the type series: 1 , Brazil:

Downloaded by [Michigan State University] at 11:29 24 December 2014 anellus claw-like, evenly curved and tapered. Median process Ce[ara], Guaramiranga (VOB). This specimen possibly be- of juxta tapered, not curved to one side, similar in length to longs to this species; it differs mainly in fore wing pattern, in main part of juxta or slightly shorter, with a few spines at apex. which the roughly diamond-shaped central area is white. Aedeagus with spinulose region situated opposite to posterior tongue, which has no spinules; vesica with spinulose sclerite, Description not rounded at tip, on short caecum. ADULTS (Figs 14,15). Fore wing length: ,22 mm; , 23 mm. Wing pattern: fore wing of both sexes mottled pale and dark FEMALE GENITALIA (Figs 213A, B). Lamella postvagina- brown with some very fine transverse streaks; female with lis with a pair of shallow rounded depressions. Lamella ante- dark postmedial and antemedial lines broken and indistinctly vaginalis a crescent-shaped band, varying in width, with or outlining roughly diamond-shaped central area; hind wing mid without very weak pair of lobes. Antrum without posterior brown with darker but indistinct subapical band and (female) projection. Corpus bursae with narrower anterior part; signum trace of postmedial line. Under-side of both wings with dark broad and triangular, situated in bulbous part of corpus bursae, subapical band. Male retinaculum strongly enlarged and with well away from anterior end. large transparent central 'window'. Hind wing of male with anal margin slightly expanded, and with slight hair-scale tuft. Diagnosis Moths of illineata are not easily distinguished by external MALE GENITALIA (Figs 101, 160). Uncus distinctly hood- features. The small size of the male moths (much smaller shaped and moderately elongate, with small digitate apical 56 Linda M. Pitkin

projection at right angles to uncus, and with pair of postero- Localities. Guatemala: no further data; Alta Verapaz, lateral lobes resembling ears. Gnathos with one median pro- Municipio San Cristobal Verapaz, Baleu; Geronimo. Belize: cess, fairly narrow, tapered, and not a distinct tongue-shape, no further data; Orange Walk. Nicaragua: Sebaco, 15 km S. similar in length to gnathos arms, or slightly longer. Valva [of] Sebaco, 'Cehite Grande'. Costa Rica: no further data; with sinuous longitudinal ridge and with apical costal lobe. [Cartago Province,] Irazu; Guanacaste Province, SW. side of Processes of anellus claw-like but with strongly broadened Volcan Cacao, Estaci´on Cacao ('Mengo'); Heredia [Province], base and narrow apex. Median process of juxta narrow and Santo Domingo; Puntarenas [Province], Monteverde; San tapered, almost straight and only marginally curved to one Jose. side, longer than (but less than 1.5 times length of) main part of juxta, with a short row of spines at apex. Aedeagus without Description spinulose region; with short, broad, rounded, posterior tongue; ADULTS (Figs 16, 17). Fore wing length: , 18-22 mm; , vesica with sclerite not on distinct caecum, rounded at tip and 19—24 mm. Wing pattern: fore wing of both sexes mottled bearing a row of squat spines. mid to dark brown with paler, sometimes whitish or olive- greenish flecks or diffuse patches, with darker antemedial and FEMALE GENITALIA (Fig. 214). Lamella postvaginalis postmedial lines sometimes moderately conspicuous, and also without pair of rounded depressions. Lamella antevaginalis (in some males) dark spot towards anal angle. Fore wing of broad, with no more than extremely weak lobes and median female often with extensive area of mainly cream or white indentation. Antrum with approximately digitate posterior pro- (consisting of roughly diamond-shaped central area extended jection. Bursa copulatrix elongate, with slightly bulbous an- to outer margin of wing), contrasting with darker base and terior region; signum disc-shaped and with small base situated apex of wing, but sometimes wing more or less concolorous, not far off centre; signum situated towards anterior end of bursa or with barely darker central area. Hind wing varying from copulatrix. moderately pale brown in both sexes to dark brown (male) or to mid brown (female), either with some darker markings Diagnosis or almost plain. Male retinaculum enlarged and with small to Moths of klagesi do not have very distinctive external features, moderate-sized transparent central 'window'. Hind wing of but the species is clearly defined by putative autapomorphies male with anal margin not strongly expanded and with hair- of the genitalia. The male is unique in having a pair of postero- scale tuft usually slight. lateral lobes on the uncus. In the female genitalia, klagesi is unusual in having the base of the signum nearly central MALE GENITALIA (Figs 102, 103, 161). Uncus strongly in the signum, not at the rim as in most other Ischnopteris hood-shaped, broad; form A without, but form B with, dorsal species, and the signum is round, not broader than long as in patch of hair-scales (situated well away from base of uncus); chlorophaearia. Ischnopteris pronubata has a signum similar with pair of tiny digitate apical projections, one lying over the to that of klagesi but with a large base, and the bursa copulatrix other, upper one pointed, both projections either at right angles is not elongate as in klagesi. to uncus or bent over and directed anteriorly. Gnathos with one median process, tongue-shaped with apex a tiny hook or occa- Distribution sionally merely a point, much longer than gnathos arms. Valva Brazil. with sinuous longitudinal ridge and with apical costal lobe. Processes of anellus varying strongly: very broad, squarish blades, except for pointed tip (form A); slender to moderately broad, evenly curved and tapered, sometimes with region of Ischnopteris rostellaria (Felder & Rogen hofer) very fine, inconspicuous spinules on inner curve (form B). Figs 16, 17, 102,103, 161, 215 Median process of juxta tapered, straight or curved to right (in Downloaded by [Michigan State University] at 11:29 24 December 2014 Syrtodes? rostellaria Felder & Rogenhofer, 1875: pl. 125, fig. ventral view), with spinules mainly in apical half; length of 1. Holotype , MEXICO (BMNH) [examined]. median process varying from same as length of main part of Ischnopteris rostellaria (Felder & Rogenhofer); Pitkin juxta to more than 1.5 times length. Aedeagus with or without etal., 1996: 135. spinules on posterior tongue (1-2 spinules sometimes present in form A, absent in form B), both forms with group of spinules Material examined situated opposite tongue, subapically to it; vesica with spinose Form A: 9 , 5 (including 5 , 2 genitalia preparations) sclerite on very short caecum. (BMNH, CMNH, VOB, ZMUC). FEMALE GENITALIA (Fig. 215). Lamella postvaginalis with Holotype , Mexico: no further data (genitalia slide Geom: pair of depressions occasionally pronounced but more usually 21092, BMNH). weak and indistinct. Lamella antevaginalis a narrow curved band, occasionally forming a very weak pair of lobes. Antrum Localities of additional material. Mexico: no further without posterior projection. Corpus bursae with anterior part data; Col[ima], Comala; Jalapa; Tam[aulipas], El Encino; often narrower but not clearly differentiated. Signum situated Tam[aulipas], Gomez Farias. well away from anterior end of corpus bursae, varying from Form B: 16 , 10 (including 9 , 5 genitalia preparations) disc-shaped to weakly subtriangular and slightly broader than (AMNH, BMNH, CMNH, INBio). long, small or moderate-sized. Neotropical geometrid moths 57

Diagnosis subapical spinules are sometimes also present; the vesica has a Moths of rostellaria are not readily distinguishable by external long, narrow, finely spinulose sclerite, not rounded at its apex features, but the male genitalia have a definitive character: the and not on a distinct caecum. tongue-shaped median process of the gnathos. The female does The female genitalia differ from those of the fabiana- not have such a broad signum as that of other members of the group in that the lamella postvaginalis never has a pair of fabiana-group (except fabiana, which has a smaller signum, depressions, and the antrum always has a posterior projection. and klagesi, which is unusual in having the base of the signum That projection extends to varying degree beyond the lamella situated nearly in the centre of the signum). I. rostellaria is antevaginalis, and is often digitate but occasionally oval or tri- treated here as a single variable species, but two forms are angular. The corpus bursae never has a differentiated narrower noted, differing in several respects as described above. Further anterior part, and the signum is a moderately long oval with a study is needed to show if the two forms are truly conspecific base no more than one half the length of the signum. or not. Form B is notable in having a dorsal patch of hair-scales The chryses-group is a tight-knit assemblage of species on the uncus of the male genitalia, but not situated at the base exhibiting less variation than in the genus as a whole. For of the uncus as in fabiana and multistrigata. instance, the signum is uniform in shape, whereas a wide range of shapes is seen elsewhere in the genus. Biological notes Moths of form A have been found at altitudes of 250-1600 m, from June to August; moths of form B have been found at Ischnopteris chavesi sp. nov. altitudes of 1100 m to 1830-2140 m, in January, June to Au- Plate 1F-I; Figs 18,19,104,162, 216 gust, October and November. Material examined Distribution 16 , 13 (including 5 , 3 genitalia preparations) Form A of rostellaria is known only from Mexico; form B occurs from Guatemala to Costa Rica. Holotype , Costa Rica: Guanacaste Prov[ince], W. side of V[olcan] Cacao, F[in]ca La Luz, 1000 m, 3-8. viii. 1986 Ischnopteris chryses species group (Janzen & Hallwachs) (INBio). The wing pattern of the chryses-group is fairly typical of that of Paratypes. Costa Rica: 1 , Guanacaste Province, Area Ischnopteris in general, but the hind wing nearly always has a de Conservacion Guanacaste, Sector Cacao, Estacion Ca- large orange or yellowish region, apical and sometimes also at cao, latitude: 10.92691, longitude: -85.46822, LN 323104- costa (inornata is the only member without orange), whereas 375725, 1150 m, larva (penultimate instar) on Conostegia an orange apical marking occurs in only a few other species of xalapensis, 27. vi. 2002, moth emerged 12. viii. 2002 Ischnopteris). The fore wing in the chryses-group is predom- (Quesada) (voucher: 02-SRNP-9835, DHJ); 1 , data as inantly mottled dark or mid brown, often with greenish tinge before but larva (last instar) on Inga longispica, 25. ii. or with other markings; a white blotch near the midpoint of the 1997, moth emerged 4. iv. 1997 (Janzen's 'gusaneros') costa may be present or absent in the male; the fore wing of (voucher: 97-SRNP-732, DHJ); 1 , data as before, Sec- the female often has a central area, roughly diamond-shaped, tor Cacao, Sendero Circular, latitude: 10.92714, longitude: of contrasting colour, paler or darker. As in other species that -85.46683, LN 323129-375878, 1185 m, larva (2nd instar) have an orange blotch on the hind wing, the under-side of on Conostegia xalapensis, 28. vi. 2002, moth emerged the fore wing usually has a slanting transverse yellowish or or- 9. ix. 2002 (Quesada) (voucher: 02-SRNP-9864, DHJ); ange band, bordered by a dark brown band or patch towards the 1 , data as before, Sector Cacao, Sendero Derrumbe, outer margin of the wing, and a line towards the middle of the latitude: 10.92918, longitude: -85.46426, LN 323354-

Downloaded by [Michigan State University] at 11:29 24 December 2014 wing, and often also a yellowish region at base of wing; these 376160, 1220 m, larva (penultimate instar) on Prunus an- markings are often indistinct in fassli and absent in inornata. nularis, 9. ii. 2001, moth emerged 31. iii. 2001 (Ramirez) The dorsum of the male fore wing always has a distinct median (voucher: 01-SRNP-6415, DHJ); 1 , data as before, Sec- tuft; this occurs in only a few other Ischnopteris species. tor Cacao, Sendero Maritza, latitude: 10.93644, longitude: The male genitalia have a definitive character of the un- -85.47764, LN 324162-374701,760 m, larva (last instar) on cus: the ventral setae form a distinct pair of lateral patches on Cinnamomum aff. triplinerve, 13. vii. 1999, moth emerged low cushion-like lobes, which are well demarcated along their 20. viii. 1999 (Pereira) (voucher: 99-SRNP-1126, DHJ); 1 inner edge (although sometimes weakly developed in janzeni), , data as before, Sector Cacao, Sendero Nayo, latitude: with a narrow median patch situated between them subapic- 10.92446, longitude: -85.46953, LN 322834-375582,1090 ally. The uncus is no more than slightly hood-shaped and it is m, larva (last instar) on Perrotettia longistylis, 18. vi. 1997, digitate to moderately broad; shaped like a fountain pen nib, moth emerged 25. vii. 1997 (Moraga) (voucher: 97-SRNP- with a short, tapered, digitate apex. The gnathos always has 1356, DHJ); 1 , data as before but larva (penultimate instar) two median processes, each with a strongly hooked tip; paired on Psychotria sp., 18. vi. 2002, moth emerged 21. vii. 2002 processes that have a strongly hooked tip occur only occasion- (Quesada) (voucher: 02-SRNP-9671, DHJ); 1 , data as be- ally elsewhere in the genus. The median process of juxta has fore but larva (penultimate instar) on Senna papillosa, 21. spinules in its apical half or at its apex. The aedeagus always v. 2002, moth emerged 11. vii. 2002 (Quesada) (voucher: has several short spinules on the posterior tongue, and other 02-SRNP-9493, DHJ); 1 , larva on Cissampelos pareira, 58 Linda M. Pitkin

moth emerged 11. vii. 2003 (voucher: 03-SRNP-4239, forming a narrow band with no more than very weak develop- DHJ); 1 , larva on Cissampelos pareira, moth emerged ment of lobes and median indentation. Antrum with posterior 9. viii. 2003 (voucher: 03-SRNP-4670, DHJ); 1 , larva oval or digitate projection on right side of antrum (in ventral on Hampea appendiculata, moth emerged 1. viii. 2003 view), projection extending to varying degree beyond lamella (voucher: 03-SRNP-4664, DHJ) [all 11 records above antevaginalis. Bursa copulatrix long, fairly slender posteriorly are based on digital photographs]; 1 , Guan[acaste] but with large bulbous anterior part; posterior end of bursa Prov[ince], SW. side of Vol[can] Cacao, Est[acion] Ca- copulatrix with wrinkled sclerotized patch. cao, LN 323300-375700, 1000-1400 m, iv. 1991 (Chaves) (INBio); 1 , 2 , Guan[acaste] Prov[ince], SW. side of Diagnosis Vol[can] Cacao, Estac[ion] Cacao ('Mengo'); 10°55'43"N Moths of chave si resemble those of chryses, and do not have the 85°28'10"W [ ], 1100 m, 3. i. 1987 [ ], 7. v. 1988 [ ], orange-yellow region of the hind wing speckled as in fassli. ii.1989 [ ], (GNP Biodiversity Survey [ ], Janzen & They can usually be recognised by the distinct dark apical Hallwachs [ ]) (INBio); 1 , 2 , Guanacaste Pr[ovince], spot on the hind wing (more pronounced than in chryses and W. side of Volcan Cacao, Estac[ion] Cacao ('Mengo'), other similar members of the chryses-group). A species out- Derrumbe, 1400 m, 5.vi, 11. vii. 1988 (Janzen & Hallwachs) side the chryses-group, aurudaria, sometimes has an apical (INBio); 4 , 4 , Puntarenas [Province]: Monteverde, spot, but aurudaria has a much larger male retinaculum than 1400 m, 1500 m, 26. iii. 1960, 1-4. ix. 1999 (Becker, Palmer) that of chavesi. The male genitalia of chavesi are typical of (AMNH, VOB); 1 , R. B. Monteverde, Est[acion] La the chryses-group and are similar to those of fassli. The two Casona, LN 253250-449700, 1520 m, vii 1993 (Obando) species share a putatively synapomorphic character, the very (INBio). Panama: 1 , Chiriqui, Rio Hornito, 800 m, v. 1994 broad squarish processes of the anellus, but there are several (Snyder) (AMNH). distinctions defining chavesi. The valva has a short digitate apical costal lobe (whereas that of fassli is irregularly roun- Description ded), the process of the juxta is longer, and the aedeagus has ADULTS (Figs 18, 19). Fore wing length: , 21-23 mm; , a subapical triangular spine instead of a group of spinules. In 22—24 mm. Wing pattern: fore wing of male usually without the female genitalia, chavesi and fassli both have the lamella whitish blotches, but occasionally with several of these (at antevaginalis as a narrow band, but chavesi has a more bulbous costa, towards apex, and forming broken postmedial band); bursa copulatrix than fassli. fore wing of female with roughly diamond-shaped central area ranging from much paler, occasionally almost white, to almost Biological notes concolorous. Hind wing of both sexes dark to mid brown, Moths have been found at altitudes of 800 m to 1520 m, in with apical orange-yellow region, and with distinct dark spot February to September. Larvae (Plate 1F—I) have been reared at or very near apex. Under-side of fore wing with irregular, by Janzen & Hallwachs (see http://janzen.sas.upenn.edu/ slanting, dark brown band (towards outer margin) and with caterpillars/database.lasso for further details), from hostplants: slanting line (towards middle), and hind wing with dark brown band or line sometimes forming an almost separate blotch near Celastraceae: Perrottetia longistylis Rose tornus. Male retinaculum slightly enlarged, and with small Fabaceae: Inga longispica Standl. transparent central 'window'. Hind wing of male with tufts Fabaceae: Senna papillosa (Britton & Rose) Irwin & Barneby of hairs more dense than in chryses (comparison made using Lauraceae: Cinnamomum aff. triplinerve (Ruiz & Pav.) specimens in very good condition as hairs are deciduous); hind Kosterm. wing of male with anal margin slightly to moderately expanded Malvaceae: Hampea appendiculata (Donn. Sm.) Standl. and with slight to moderate hair-scale tuft. Melastomataceae: Conostegia xalapensis (Bonpland) Downloaded by [Michigan State University] at 11:29 24 December 2014 D. Don MALE GENITALIA (Figs 104A-B, 162). Uncus moderately broad, slightly hood-shaped; with ventral setae grouped in Menispermaceae: Cissampelos pareira L. distinct well-separated pair of lateral patches on low cushion- Rosaceae: Prunus annularis Koehne like lobes, and small, narrow, median subapical patch. Gnathos Rubiaceae: Hamelia patens Jacq. [record unconfirmed] with two median processes, which are as long as gnathos arms Rubiaceae: Psychotria sp. (measured from base of arm to point where process arises). Sabiaceae: Meliosma glabrata (Liebm.) Urb. [record uncon- Valva with apical costal lobe short and digitate, with a minute firmed, larva died] projection on the lobe. Processes of anellus very broad blades, squarish except for pointed tip. Median process of juxta a Distribution blade, usually tapered irregularly, more than twice to almost Costa Rica and Panama. three times length of main part of juxta, occasionally deflected to right (in ventral view), apical half or less with spinules along Ischnopteris chryses (Druce) one edge. Aedeagus with triangular spine (situated opposite Figs 20, 21, 105, 163, 217 and subapically to posterior tongue bearing the usual spinules). Ischnopteryx chryses Druce, 1893: 158; ibidem 3: pl. 40, figs FEMALE GENITALIA (Fig. 216). Lamella postvaginalis 23-25. LECTOTYPE , PANAMA (BMNH), here desig- without pair of rounded depressions. Lamella antevaginalis nated [examined]. Neotropical geometrid moths 59

Ischnopteris chryses ab. dispar Warren, 1900: 203. FEMALE GENITALIA (Figs 217A, B). Lamella postvaginalis [Infrasubspecific name.] without pair of rounded depressions. Lamella antevaginalis Ischnopteris chryses (Druce); Pitkin, 2002: 213–4. varying in shape but never as narrow as in chavesi or fassli; with or without pair of weak lobes separated by slight median Material examined indentation. Antrum with posterior digitate projection extend- 26 , 14 (including 11 , 5 genitalia preparations) ing a short way beyond lamella antevaginalis. (AMNH, BMNH, DHJ, INBio, USNM, VOB, ZSM). Lectotype , Panama: Volcan de Chiriqui, 610-920 m Diagnosis ('2000-3000 ft') (Champion) (genitalia slide Geom: 20789, The hind wing pattern of chryses is like that of palmeri, and BMNH). Paralectotypes: Panama: 1 , data as lectotype sometimes also aurudaria, but chryses differs from those spe- (BMNH); 1 , Bugaba (Champion) (BMNH). cies in having a distinct median tuft on the dorsum of the male fore wing. Also, the male retinaculum of chryses is smal- Localities of additional material. Mexico: [Veracruz,] ler than that of aurudaria. The wing pattern of chryses is not Misantla; Tam[aulipas], Gomez Farias. ´Guatemala: Hue- clearly distinguishable from that ofhoffmani, lata, andwatsoni huetenango, Municipio Santa Cruz, Barillas, Chiblac. Costa (chryses-group members allopatric from chryses), and janzeni Rica: Alajuela Province, Area de Conservacion Guanacaste, (sympatric with chryses). Ischnopteris chryses is better distin- Sector San Cristobal, Sendero Corredor [digital photograph, guished and defined by genitalic characters, most notably the DHJ]; [Alajuela Province], 9 km S. of S[an]ta Cecilia, shape of the apical costal lobe of the valva, unique in combin- Est[acion] Pitilla; [Cartago Province:] I.I.C.A. grounds; Orosi; ing a rounded lobe with a digitate posterior process. Tuis; Turrialba, Chirripo, Grano de Oro; Guanacaste Province: Rincon National Park, Mirador; Puntarenas [Province], 800 m NW. of Tigra, Embalce, Finca Cafrosa; San Jose Province: Biological notes Braulio Carrillo National Park, Estacion Carrillo. ´Panama: no Moths have been found at altitudes of 600-1615 m, in March further data; Bugaba; Volcan de Chiriqui. ´Ecuador: Chimbo; to September, and November. A larva was reared by Jan- Cotopaxi, San Francisco de Las Pampas; Esmeraldas, San zen & Hallwachs (see http://janzen.sas.upenn.edu/caterpillars/ Mateo; Lita; Paramba; Pichincha, 12 km SE. [of] Santo database.lasso for further details; voucher: 00-SRNP-11624), Domingo, Tinalandia. Peru: Chanchamayo. from the hostplant Monimiaceae: Siparuna andina (Tul.) A. Dc. Description ADULTS (Figs 20,21). Fore wing length: ,18-20 mm; , 21- Distribution 23 mm. Wing pattern: fore wing of male with or without white Central America from Mexico to Panama; also in Ecuador and blotch near midpoint of costa, sometimes plus several other Peru. smaller diffuse whitish markings. Hind wing of both sexes dark to mid brown with apical orange or yellow region; margin between brown and yellow regions not as sharply angled as Ischnopteris fassli sp. nov. in brehmi. Under-side of fore wing with slanting dark brown Figs 22, 23,106,164, 218 band (towards outer margin) and line (towards middle), and hind wing with dark brown band or line sometimes forming Material examined an almost separate round blotch near tornus. Male retinaculum 10 , 11 (including 3 , 3 genitalia preparations) slightly enlarged, and with small transparent central 'window'. Holotype , Costa Rica: [Cartago Province,] Orosi, Hind wing of male with anal margin slightly to moderately 1200 m (Fassl) (genitalia slide Geom: 21007; BMNH). expanded, with slight to moderate hair-scale tuft. Downloaded by [Michigan State University] at 11:29 24 December 2014 Paratypes: Costa Rica: 3 , 9 , data as holotype (BMNH); MALE GENITALIA (Figs 105,163). Uncus not broad, slightly 3 , Carta[go] Province, R[io] Grande de Orosi, from hood-shaped; with ventral setae grouped in distinct well- Puente R. Dos Amigos to la Represa, LN 186600-562000, separated pair of lateral patches on low cushion-like lobes, 1800 m, ii.1995 (Delgado) (INBio); 2 , 2 , San Jose and small, narrow, median subapical patch. Gnathos with two Province, Par[que] Nac[ional] Braulio Carrillo, Estacion median processes that are shorter than gnathos arms (meas- Zurqui (el Tunel), 10°04'N 84°01'W, 1500 m, xi, xii.1985, ured from base of arm to point where process arises). Valva i.1986 (I. & A. Chacon) (INBio); 1 , San Jose Province, with weakly rounded apical costal lobe tapering to a digitate Estacion Zurqui (500 m before Tunel), LN 226800-535200, posterior process. Processes of anellus narrow blades, tapered, 1600 m, 26. ix-x. 1990 (Maass) (INBio). with pointed apex, sometimes almost spine-like, sometimes claw-shaped. Median process of juxta a long straight blade Description (approximately twice length of main part of juxta), usually ADULTS (Figs 22,23). Fore wing length: ,22-23 mm; , 22- fairly narrow and constricted at base, but occasionally not 26 mm. Wing pattern: fore wing of male with or without whit- constricted; densely spinose apically. Aedeagus with one tri- ish blotch near midpoint of costa, plus several smaller whitish angular spine plus several minute spinules on sclerite (situated markings; hind wing of both sexes mid to pale brown, with opposite and subapically to posterior tongue bearing the usual apical orange to dirty-yellow region (often with fine brown spinules). speckles present), and with dark brown lines and blotch leading 60 Linda M. Pitkin

from edge of this region to near tornus. Under-side of each but fassli has a narrower lamella antevaginalis than chryses, wing with a dark brown line situated medially or postmedially, and a less bulbous bursa copulatrix than chavesi. and with a round dark brown blotch, situated towards outer margin of each wing, usually fairly distinct in male but not Biological notes always in female; brown line on hind wing usually continued Moths have been found at altitudes of 1200 m to 1800 m, in alongside blotch, but sometimes only faintly; hind wing usu- February, and September to December. ally mainly straw colour, tinged with yellow or dull orange. Male retinaculum varying in size, often only very slightly en- Distribution larged, and with small transparent central 'window'. Hind wing Costa Rica. of male with anal margin little expanded, without pronounced hair-scale tuft. Ischnopteris hoffmani sp. nov. MALE GENITALIA (Figs 106A-B, 164). Uncus moderately Figs 24,107,165 broad, slightly hood-shaped; with ventral setae grouped in Material examined distinct well-separated pair of lateral patches on low cushion- 4 (including 3 genitalia preparations) like lobes, and small, narrow, median subapical patch. Gnathos with two median processes. Valva with apical costal lobe weak Holotype , Brazil: Santa Catarina, Blumenau, x. 1937 (Hoff- and irregularly rounded (not digitate), forming a minute point man) (genitalia slide Geom: 21188; BMNH). posteriorly which is sometimes indistinct. Processes of anellus Paratypes: Brazil: 1 , Ba[hia], Camaca, 400-700 m, 21-30. very broad blades, squarish except for pointed tip. Median ix. 1991 (Becker) (USNM); 2 , Sao Paulo, Alto da Serra, process of juxta a pointed blade, more than 1.5 times to almost xii. 1922, vi. 1923 (Spitz) (BMNH). twice length of main part of juxta, slightly or distinctly curved to right (in ventral view), apical two-fifths to one half of length Description with spinules along one edge. Aedeagus with group of spinules ADULTS (Fig. 24). Fore wing length: , 19-20 mm. Wing (situated opposite and subapically to posterior tongue bearing pattern (only male known): fore wing with whitish blotch near the usual spinules, which include one larger spinule and form midpoint of costa, plus several smaller whitish markings; hind a longitudinal band). wing dark brown with apical orange region. Under-side of fore wing with dark brown broad band (towards outer margin) FEMALE GENITALIA (Figs 218 A, B). Lamella postvaginalis and line (towards middle); hind wing with dark brown line, without pair of rounded depressions. Lamella antevaginalis and with round blotch near tornus. Male retinaculum barely forming a narrow band without lobes or median indentation. enlarged, and with small transparent central 'window'. Hind Antrum with posterior digitate projection on left side of antrum wing of male with anal margin slightly to moderately expan- (in ventral view), projection extending well beyond lamella ded, with hair-scale tuft not pronounced. antevaginalis. Bursa copulatrix long, very slender posteriorly and with small bulbous anterior. MALE GENITALIA (Figs 107,165). Uncus not broad, slightly hood-shaped; with ventral setae grouped in distinct well- Diagnosis separated pair of lateral patches on low cushion-like lobes, Moths of fassli can generally be distinguished by the hind wing and small, narrow, median subapical patch. Gnathos with two pattern, with the brown region never dark, with speckles often median processes. Valva with minute pointed apical costal lobe present in the orange/yellow region giving a dirty appearance, (not digitate). Processes of anellus blade-like, not broad. Me- and with brown lines and other markings leading from the or- dian process of juxta broad with spatulate apex, not curved ange/yellow region to the tornus. There is some resemblance to one side, as long as main part of juxta or slightly shorter, Downloaded by [Michigan State University] at 11:29 24 December 2014 to brehmi, but the two species differ in the predominant colour and with many spinules at apex. Aedeagus without subapical of the under-side of the hind wing (mainly yellowish straw spines or spinules other than those on the posterior tongue. in fassli and greyish brown, often dark, in brehmi). Moths of chavesi and watsoni (females) and aurudaria (both sexes) FEMALE GENITALIA. Unknown. are occasionally also similar to fassli, but with a brighter orange/yellow region, and chavesi has a distinctive dark apical Diagnosis spot, (compare Figs 22 and 23 of fassli with Figs 19, 29 and Moths of hoffmani resemble those of chryses, but hoffmani is 34 of the other species). clearly defined by an autapomorphy of the male genitalia, the The male genitalia of fassli are typical of the chryses- unique shape of the median process of the juxta, being broad group and are similar to those of chavesi. The two species with a spatulate apex. share a putatively synapomorphic character, the very broad squarish processes of the anellus, but there are several dis- Biological notes tinctions defining fassli. The valva has an irregularly rounded One moth was found at an altitude of 400-700 m; moths have apical costal lobe (whereas that of chavesi is short and digit- been found in June, September, October and December. ate), the process of the juxta is shorter, and the aedeagus has a subapical group of spinules and not a triangular spine. The Distribution female genitalia are similar to those of chryses and chavesi, Brazil. Neotropical geometrid moths 61

Ischnopteris inornata sp. nov. dark tufts on the wings as in hirsuta and obtortionis, nor do Figs 25,108,166 they have the white band on the fore wing that is characteristic offasciata. The male genitalia of inornata are similar to those Material examined of watsoni in some respects, but differ in having the apex of 12 (including 6 genitalia preparations) the valva somewhat extended and tapered on the non-costal side, and the aedeagus without a subapical spine. Also, the Holotype , Brazil: Espirito Santo, S[an]t[a] Leopoldina, Dorf median process of the juxta, in its combination of features Tirol, 650 m, 8-20 xii. 1996 (leg. Thony) (ZSM). described above, is unlike any other in the chryses-group and Paratypes: Brazil: 1 , Ba[hia], Bonito, 1000 m, 25. iv. is putatively autapomorphic for inornata. 1998 (Becker) (VOB); 1 , Ba[hia], Camacan, 600m, 6. ii. 1998 (Becker) (VOB); 1 , [Minas Gerais,] Nova Lima, 850 m, 1-3. iv. 1983 (Becker) (VOB); 1 , Biological notes Parana, Castro, 950 m (´Jones) (BMNH); 1 , Parana ('PR'), Moths have been found at altitudes of 600-1100 m, in Febru- [near Antonina,] Guaratuba, 600 m, 5. viii. 1975 (Becker) ary, April, August, November and December. (VOB); 2 , [Rio Grande do Sul,] Sao Leopoldo, 29. ii. 2000 (leg. Moser) (ZSM); 2 , Rio de Janeiro, Federal Distribution Dist[rict], Corcovado Forest, 1958 (Kettlewell) (BMNH); Brazil. 1 , R[io de]J[aneiro]: Nova Friburgo, 1100 m, 9. xi. Ischnopteris janzeni sp. nov. 1998 (Becker) (VOB); 1 , Santa Catarina, [Nova] Neu Figs 26,109,167 Bremen, RioLaeiss, viii. 1931 (Hoffmann) (BMNH). Material examined Description 2 (genitalia preparations of both) ADULTS (Fig. 25). Fore wing length: , 19-22 mm. Wing Holotype , Costa Rica: San Jose Province: Braulio Car- pattern (only male known): fore wing usually without white rillo National Park, La Montura, 1100 m, 17.xii.1981 blotch near midpoint of costa, but occasionally with diffuse (Janzen & Hallwachs) (genitalia slide LMP 472; INBio). blotch and other whitish markings; hind wing grey-brown, Paratype: Costa Rica: 1 , Cart[ago] Prov[ince], P[arque] sometimes with faint traces of tan, but without orange region. N[acional] Tapanti, Quebrada Segunda, LN 194000- Under-side of wings brown with some darker markings, not 560000,1250 m, i.1993 (Mora) (INBio). contrasting strongly. Male retinaculum slightly to moderately enlarged, and with small to moderate transparent central 'win- Description dow'. Hind wing of male with tuft of dense hairs, and anal ADULTS (Fig. 26). Fore wing length: , 19-21 mm. Wing margin expanded forming distinct flap with moderate hair- pattern (only male known): fore wing with diffuse, slanting, scale tuft. medial region, which is paler than the mottled buff and brown, grey-green tinged, ground colour, and with small pale spot MALE GENITALIA (Figs 108A-B, 166). Uncus moderately near apex of wing, but no distinct white blotches (in the sample broad, slightly hood-shaped; ventral setae grouped in dis- studied); hind wing mid greyish brown with apical yellow re- tinct well-separated pair of lateral patches on low cushion-like gion. Under-side of fore wing with broad slanting dark brown lobes, and small, narrow, median subapical patch. Gnathos band (towards outer margin) and line (towards middle), and with two median processes, which are as long as gnathos arms hind wing with dark brown band or line forming a round (measured from base of arm to point where process arises). blotch, sometimes almost separate, near tornus. Male retin- Valva with apical costal lobe rounded (not digitate) and form- aculum strongly enlarged and with large transparent central ing a small point posteriorly; apex of valva somewhat extended 'window'. Hind wing of male with anal margin expanded,

Downloaded by [Michigan State University] at 11:29 24 December 2014 and tapered on non-costal side. Processes of anellus curved and with slight to moderate hair-scale tuft. tapered, narrow claw-shaped with pointed apex, each process with basal sclerite (adjoining juxta) usually slightly longer than MALE GENITALIA (Figs 109, 167). Uncus fairly small, not process. Median process of juxta a long, narrow, pointed blade broad, no more than slightly hood-shaped; with ventral setae (twice length of main part of juxta), usually deflected to right grouped in pair of lateral patches on low cushion-like lobes (in ventral view) but occasionally straight, with a few spines at (weaker than usual in chryses-group), and small, weak, me- apex. Aedeagus without subapical spine or spinules other than dian subapical patch. Gnathos with two median processes. those on the posterior tongue. Valva with apical costal lobe rounded (not digitate) and forming a minute point posteriorly. Processes of anellus blade-like, FEMALE GENITALIA. Unknown. moderately broad except for pointed tip. Median process of juxta a narrow pointed blade approximately twice length of Diagnosis main part of juxta, curved to right (in ventral view), apical two- I. inornata is the only member of the chryses-group without fifths with spinules along one edge. Aedeagus with group of orange on the hind wing. As in all chryses-group species, males spinules (situated opposite and subapically to posterior tongue have a median tuft on the dorsum of the fore wing, and the bearing the usual spinules, which are all tiny and form a slant- only other Ischnopteris species with that character but without ing band). orange on the hind wing arefasciata, hirsuta, and obtortionis. Moths of inornata do not have such conspicuously contrasting FEMALE GENITALIA. Unknown. 62 Linda M. Pitkin

Diagnosis Diagnosis Moths ofjanzeni resemble those of chryses, but the male retin- Moths of lata are very similar to those of chryses, but they aculum ofjanzeni is larger than in chryses or any other mem- have a longer fore wing than that of chryses (comparing males; bers of the chryses-group. The male genitalia show affinities female of lata is currently unknown). There are several distinct with those of fassli in having a similar median process of the differences in the male genitalia: putative autapomorphies of juxta, and the processes of the anellus are somewhat similar, lata include the exceptionally broad uncus, and the presence, though less broad, in janzeni, but janzeni has a smaller uncus at the apex of the median process of the juxta, of only a lateral than fassli. The arrangement of the spinules on the posterior pair of spines. tongue of the aedeagus in a slanting, not longitudinal, band is putatively autapomorphic for janzeni. Distribution French Guiana and Brazil. Biological notes Moths have been found at altitudes of 1100 m and 1250 m, in Ischnopteris watsoni sp. nov. January and December. Figs 28, 29,111,169, 219

Material examined Distribution 10 , 7 (including 4 , 2 genitalia preparations) Costa Rica. Holotype , Venezuela: Aragua, Rancho Grande, 12.vii- 16.viii.1976 (Watson) (genitalia slide Geom: 21137; Ischnopteris lata sp. nov. BMNH). Figs 27,110,168 Paratypes: Venezuela: 1 , 1 , data as holotype but 28.vi- Material examined 18.vii.1974, 12.vii.1976 (BMNH); 3 , 4 , data as 3 (genitalia preparations of all) holotype but 17-20.i.1978, 25-26.i.1978, 30-31.iii.1978, 1-3.iv.1978 (Heppner) (USNM); 2 , data as holo- Holotype , Brazil: 1 , Rio Purus, Hyutanahan, iv.1922 type but 1100 m, 22-23.i.1978 (Heppner), 30-31.v.1988 (Klages) (genitalia slide LMP 420; CMNH). (Epstein) (USNM); 1 , 1 , Bol´ıvar [Province], Ptari- Paratypes: French Guiana: 2 , St Jean du Maroni (Le Moult) Tepui, 30 mi[les] N. [of] Kavanayen, 1800 m, 17- (BMNH). 19.viii.1970 (Dretz) (USNM); 1 , Cucuta (BMNH); 1 , Caracas, Los Venados (Vogl) (ZSM); 1 , Lara, Yacambu Description National Park, 13 km SE. [of] Sanare, 1460 m ('4800 ft'), ADULTS (Fig. 27). Fore wing length: ,22-23 mm. Wing pat- 4-7.iii.1978 (Heppner) (USNM). tern (only male known): fore wing with diffuse, faintly paler, Description slanting medial region, without white blotches (in the sample ADULTS (Figs 28, 29). Fore wing length: , 20-22 mm; , studied); hind wing of both sexes dark brown with apical 21-23 mm. Wing pattern: fore wing of male with or without orange region. Under-side of fore wing with dark brown band white blotch near midpoint of costa, plus other smaller diffuse (towards outer margin) and line (towards middle), and hind whitish markings; fore wing of female usually with roughly wing with dark brown line or band merging with brown ground diamond-shaped central area ranging from much paler, whit- colour and not forming a separate blotch near tornus. Outer ish, to almost concolorous. Hind wing of both sexes dark brown margin of wings evenly rounded. Male retinaculum barely en- with apical orange region. Under-side of fore wing with dark larged, and with small transparent central 'window'. Hind wing brown band (towards outer margin) and line (towards middle), of male with tuft of dense hairs; with anal margin expanded

Downloaded by [Michigan State University] at 11:29 24 December 2014 and hind wing with dark brown line distinct or merging with (not strongly) and with slight to moderate hair-scale tuft. brown ground colour, occasionally forming a separate round MALE GENITALIA (Figs 110A-B, 168). Uncus moderately blotch near tornus but in that case with line continued along- broad, slightly hood-shaped; with ventral setae grouped in dis- side blotch. Male retinaculum slightly enlarged and with small tinct well-separated pair of lateral patches on low cushion-like transparent central 'window'. Hind wing of male with anal lobes, and small, narrow, median subapical patch. Gnathos margin expanded (not strongly), without pronounced hair- with two median processes, which are as long as gnathos arms scale tuft. (measured from base of arm to point where process arises). MALE GENITALIA (Figs 111A-B, 169). Uncus slightly Valva with apical costal lobe strongly rounded (not digitate) hood-shaped, varying from moderately narrow to moderately and forming a minute point posteriorly. Processes of anel- broad; with ventral setae grouped in distinct well-separated lus broad, pointed blades. Median process of juxta a long pair of lateral patches on low cushion-like lobes, and small, blade, not pointed (twice, or slightly more than twice, length narrow, median subapical patch. Gnathos with two median of main part of juxta), curved to right (in ventral view) or processes, which are as long as gnathos arms (measured from almost straight; process with pair of postero-lateral spines. base of arm to point where process arises). Valva with apical Aedeagus with small patch of spinules (situated opposite to costal lobe rounded (not digitate) and forming a minute point posterior tongue bearing the usual spinules). posteriorly. Processes of anellus narrow and spine-like, each FEMALE GENITALIA. Unknown. with basal sclerite (adjoining juxta) not longer than process. Neotropical geometrid moths 63

Median process of juxta a pointed tongue-shaped blade, Lectotype (albiguttata), Peru: SE. Peru, Carabaya, Santo slightly longer than main part of juxta, almost straight or Domingo, 1830m('6000ft'),xi.1901 (Ockenden) (genitalia slightly curved to right (in ventral view), with a broad field slide Geom: 20997, BMNH). of many spinules across apical half. Aedeagus with triangular Lectotype (praeluteata), Peru: SE. Peru, Carabaya, Santo spine (situated opposite and subapically to posterior tongue Domingo, 1830 m ('6000 ft'), i.1903 (Ockenden) (genitalia bearing the usual spinules). slide Geom: 20754, BMNH). Paralectotypes 2 (praeluteata), data as lectotype except 1 v. 1902 (BMNH). FEMALE GENITALIA (Figs 219A, B). Lamella postvaginalis without pair of rounded depressions. Lamella antevaginalis Localities of additional material. Peru: no further data; forming a narrow band, with pair of weak lobes separated by shallow rounded median indentation. Antrum with posterior Carabaya: Oconeque; Santo Domingo; Quinton. triangular, sharply pointed, projection. Description ADULTS (Figs 30-32). Fore wing length: , 22-24 mm; , Diagnosis 25-27 mm. Wing pattern: fore wing of male dark to mid brown Moths of watsoni are very similar to those of chryses, but there mottled with olive-green, with dark streaks or dashes but not are several distinct differences in the male genitalia. Notable so distinct as in multistrigata, with or without whitish blotch of these are the apical costal lobe of the valva, which has near midpoint of costa, plus several smaller whitish markings; only a minute extension, not a digitate one as in chryses, and fore wing of female brown mottled with whitish markings and watsoni has longer gnathos processes than those of chryses. As often with olive-green, with roughly diamond-shaped central in chryses, the aedeagus has a subapical triangular spine, but area paler, darker, or more or less concolorous and marked watsoni has no minute spinules accompanying it. The male out only by dark lines around it. Hind wing of both sexes genitalia closely resemble those of inornata, particularly in greyish brown, without orange region but with diffuse patch the shape of the costal lobe of the valva, but they differ in the paler brown and warmer in hue, from apex of wing to dark apex of the valva, which is not extended and tapered on the postmedial line, and as narrower band along outer margin of non-costal side in watsoni as it is in inornata. The process of wing. Dorsum of male fore wing without distinct median tuft. the juxta of watsoni differs from that of inornata and others in Male retinaculum slightly to moderately enlarged, with very the chryses-group in its form: a pointed tongue with a field of weakly developed central 'window'. Hind wing of male with spinules across it, not just along one edge or at apex. The juxta anal margin strongly expanded, forming distinct flap, with process of chryses is occasionally similar but less pointed. pronounced hair-scale tuft. In the female genitalia, watsoni is the only species of the chryses-group known to have a pointed posterior projection of MALE GENITALIA (Figs 112A-B, 170). Uncus no more the antrum. than slightly hood-shaped; broadly digitate, with much smaller digitate apex straight (i.e. not bent over). Gnathos with two median processes. Valva with digitate apical costal process. Biological notes Processes of anellus blade-like, with narrow pointed tip. Me- Moths have been found at altitudes of 1100 m to 1800 m, in dian process of juxta similar in length to main part of juxta, January, and March to August. straight, spinulose almost to base. Aedeagus with a few triangular spinules (2 spinules present in specimens examined) on posterior tongue; vesica with long, narrow, pointed and finely Distribution spinulose sclerite, not on distinct caecum. Venezuela.

Downloaded by [Michigan State University] at 11:29 24 December 2014 FEMALE GENITALIA (Figs 220A-C). Lamella postvaginalis with pair of small rounded depressions indistinct or absent. Species not placed to group Lamella antevaginalis forming a broad band without distinct Ischnopteris albiguttata Warren lobes or median indentation, but with strong wrinkles on sur- Figs 30-32,112,170, 220 face adjoining antrum, including slightly asymmetrical pair of wrinkled patches near antrum, one on each side. Antrum Ischnopteris albiguttata Warren, 1904a: 117. LECTOTYPE with posterior digitate projection. Bursa copulatrix long and cf, PERU (BMNH), here designated [examined]. slender; signum a short oval with base half, or more than half, Ischnopteris praeluteata Warren, 1904b: 558. LECTO- length of signum. TYPE , PERU (BMNH), here designated [examined]. [Synonymy cited by Parsons et al., in Scoble (1999).] Ischnopteris praeluteata ab. albirupta Warren, 1904b: 558. Diagnosis [Infrasubspecific name, nomen nudum.] The hind wing pattern described above for albiguttata is distinctive and putatively autapomorphic for the species. The species is defined, in the male genitalia, by the juxta process having Material examined spinules almost to its base, (which does not occur in other spe- 15 , 17 (including 4 , 2 genitalia preparations) (BMNH, cies that have the processes of the anellus blade-like and the USNM). gnathos with two median processes). I. albiguttata and brehmi 64 Linda M. Pitkin

have extremely similar female genitalia, differing from those apical costal process, usually without lobe where this process of other species by having characteristic strong wrinkles on forms an angle with the costal margin but occasionally with the lamella antevaginalis, which is a broad band, not indented extremely slight lobe. Processes of anellus narrow and spine- medially. The two species differ from each other in the size of like with blunt apex, each process with broader, curved base, the base of the signum, which is larger in albiguttata (compare extending over anellus as large sclerotized region from pos- Figs 220C and 223). terior end of which spine arises. Median process of juxta short and evenly tapered (much shorter than main part of juxta), with Biological notes spinules minute or absent. Aedeagus with a group of several Moths have been found at altitudes of 1530-2140 m, in Janu- short spinules and one broad flat triangular spine (varying in ary, April to July, and October to December. size) on posterior tongue; vesica with long spinulose sclerite, not rounded at tip, not on distinct caecum.

Distribution FEMALE GENITALIA (Figs 221A, B). Lamella postvaginalis Peru. with pair of rounded depressions indistinct or absent. Lamella antevaginalis forming a narrow band without pair of lobes or Ischnopteris aurudaria (Schaus) median indentation. Antrum without distinct posterior projec- Figs 33, 34. "3.171, 221 tion. Signum approximately disc-shaped, with base almost as large as signum. Ischnopteryx aurudaria Schaus, 1901: 249. LECTOTYPE , BRAZIL (USNM), here designated [examined]. Ischnopteris aurudaria (Schaus); Parsons et al., in Scoble, Diagnosis 1999: 519. The hind wing of aurudaria has an orange region that is extensive, but less so than in catocalata or stenoptila. The hind wing pattern is variable and occasionally resembles that of Material examined some specimens of pronubata, except that the dark region 12 , 6 (including 7 , 2 genitalia preparations) (AMNH, does not extend from the dorsum up the outer margin as it BMNH, CMNH, ZSM, USNM, VOB). does in pronubata (compare Figs 33 and 34 with 57 and 58). The orange region of aurudaria is usually more extensive than Lectotype Brazil: SE. Brazil, Sao Paulo (Type No. 12469, that of the chryses-group, or of palmeri, but there is a pos- genitalia slide 60137, USNM). sibility of confusion in a few cases. The dorsum of the male fore wing of aurudaria differs from that of the chryses-group Localities of additional material. Brazil: Parana: Castro; in that the median tuft is weak or absent, and the male retin- Curitiba; R[io Grande do]S[ul]: S[ao] Jose d. Ausentes, aculum is larger than in members of the chryses-group, with Silveira; R[io de]J[aneiro]: Itatiaia; Santa Catarina ('S. Cath.', the exception ofjanzeni. The genitalia are not at all like those 'SC'): Bom Jardim da Serra; Rio Vermelho; Sao Paulo: no of the chryses-group or palmeri. Instead aurudaria is close to further data; Sao Paulo: Boraceia; Alto da Serra: no further chloroclystata in male and female genitalic features, although data; Alto da Serra: Santos. it is easily distinguished from the latter by the presence of an orange region on the hind wing. The form of the processes of Description the anellus (as described above) is a putative synapomorphy ADULTS (Figs 33, 34). Fore wing length: , 17-22 mm; , of aurudaria, chloroclystata, and fasciata. I. aurudaria dif- 20-24 mm. Wing pattern: fore wing mottled dark brown, or fers from the other two species in the distal valva shape (the dark brown with pale brown areas, and (in a minority of males) valva does not have a clearly developed lobe on the costa at

Downloaded by [Michigan State University] at 11:29 24 December 2014 with white blotch near midpoint of costa. Hind wing dark or the base of the digitate process, as is present in chloroclystata mid brown in dorsal half, with large apical orange region of- and fasciata). There are also differences in the aedeagus: the ten extending along most of costal half of wing, sometimes larger spine amongst the spinules on the aedeagus is a broad with wavy dark brown lines running from midpoint of costa triangular shape in aurudaria, but not in chloroclystata, and towards tornus. Under-side of fore wing with slanting trans- aurudaria has a larger sclerite of the vesica of the aedeagus verse yellowish or orange band, bordered by dark brown band than fasciata. Female genitalia of aurudaria and chloroclys- or patch (towards outer margin) and line (towards middle of tata are very similar, and share the character of the signum wing); hind wing with wavy dark brown line (or lines) and having a large base (but not in the centre of the signum as is a blotch near tornus. Dorsum of male fore wing with weak the case in pronubata). median tuft, or without tuft. Male retinaculum distinctly enlarged and with large transparent central 'window'. Hind wing of male with anal margin slightly to moderately expanded and Biological notes with moderate hair-scale tuft. Moths have been found at altitudes of 800 m to 2100 m, in January, February, April, October, and December. MALE GENITALIA (Figs 113, 171). Uncus only slightly hood-shaped; broadly digitate, with much smaller digitate apex straight (i.e. not bent over). Gnathos with two median pro- Distribution cesses, with hooked or weakly curved tip. Valva with digitate Brazil. Neotropical geometrid moths 65

Ischnopteris bifurcata sp. nov. Diagnosis Figs 35, 36,114, 115,172, 222 The moths of bifurcata are not very distinctive but the species is unique in its male genitalia structure. I. bifurcata is the Material examined only species that has the two prongs of the bifurcate processes Form A: 4 , 2 (genitalia preparations of all) of the anellus close together and of narrow spine-like propor- Holotype , Bolivia: [Cochabamba,] Chapare, San Jacinto, tions (other species with bifurcate processes of the anellus are xi. 1996 (Thony¨) (genitalia slide LMP 377; ZSM). chlorata, which has the prongs well apart, and hirsuta, which Paratypes: 1 , data as holotype (ZSM); 2 , 1 , Cochabamba has one of the two prongs very short and squat in shape). The (Steinbach) (CMNH). Peru: 1 , Chanchamayo, 1912 female genitalia of bifurcata resemble those of chlorata in (Schuncke) (BMNH). having a broad lamella antevaginalis without indentation, but Form B (excluded from the type series): 2 (genitalia prepar- the signum of bifurcata is shorter than that of chlorata. ations of both) Variation Localities. Bolivia: 1 , La Paz, Alto Rio Beni, 1600 Only a small number of specimens are known, but these fall m, 10-11.i.1976 (Pena˜) (AMNH). Peru: 1 , Cushi, Huanuco, into two forms (referred to here as A and B) showing a number 1820 m, 1904 (Hoffmanns) (BMNH). of differences. I am provisionally treating the two forms as conspecific, but I consider it best to restrict the type series to Description specimens of form A. ADULTS (Figs 35, 36). Fore wing length: (form A) , 23- Biological notes 25 mm, , 24 mm; (form B) , 21 mm. Wing pattern: fore wing Moths have been found at altitudes of 1600 m and 1820 m of both sexes mottled dark to mid brown with some grey-green (data available only for form B), and they have been collected or olive, and with short striations, mainly on the costa; fore in January (form B) and November (form A). wing of female (single specimen known) with paler central area, roughly diamond-shaped but not distinctly demarcated. Hind wing brown, sometimes greyish, with darker but in- Distribution distinct postmedial line and submarginal or marginal band; Bolivia and Peru. without orange region; postmedial line of female weakly angled and almost touching dark submarginal band at or near Ischnopteris bipectinata sp. nov.

vein M3. Dorsum of male fore wing without median tuft. Male Figs 37,116,173 retinaculum strongly enlarged and with large transparent cent- Material examined ral 'window'. Hind wing of male with anal margin slightly 9 (including 4 genitalia preparations) expanded, or fairly strongly expanded and forming distinct flap with hair-scale tuft. Holotype , Ecuador: Loja [Province], Vilcabamba, 1500 m, 14.ii.1993 (Mery & Attal) (genitalia slide LMP MALE GENITALIA (Figs 114,115,172). Uncus only slightly, 369; ZSM). or not, hood-shaped, broadly digitate, with much smaller digit- Paratypes: Ecuador: 1 , no further data (Buckley) (BMNH); ate apex straight or only slightly bent over, this apex either long 7 , Manabi, near Machalilla, Agua Blanca, 50 m, 23.v.1996 and narrow (form A) or very short (form B). Gnathos with two (Hillman) (CMNH). median processes, with curved tip. Valva with digitate apical costal process. Processes of anellus bifurcate with narrow, Description tapered prongs close together, form A with another smaller ADULTS (Fig. 37). Fore wing length: ,15-16 mm. Male an-

Downloaded by [Michigan State University] at 11:29 24 December 2014 spine at base of each pair of processes; both forms with patch tenna strongly bipectinate. Wing pattern (only male known): of minute spinules situated posteriorly on sclerotized region at fore wing mottled pale brown to dark greyish brown, with dark base of processes of anellus. Median process of juxta slender, brown spot near tornus; usually with fine dark lines, slanting varying in length (form A: twice or almost twice length of main and transverse, indistinct and irregular, but sometimes with part of juxta, form B: slightly more or slightly less than 1.5 strong markings, notably a pale basal wedge contrasting with times length of main part of juxta); form A: process straight dark marking bordering it. Hind wing usually more or less or slightly deflected to right (in ventral view), form B: process concolorous with fore wing, brown, sometimes greyish, with more strongly deflected to the right; process of both forms with darker band at or near outer margin usually diffuse and some- spinules at least towards apex. Aedeagus with two groups of times indistinct, and with broken indistinct postmedial line, short spines or spinules, one group on posterior tongue and the without orange region. Under-side of wings often with post- other group situated opposite; vesica with very slight, narrow, medial and other lines more distinct than on upper side. Dor- spinulose sclerite, not rounded at tip, not on distinct caecum. sum of male fore wing without median tuft. Male retinaculum FEMALE GENITALIA (Fig. 222). Lamella postvaginalis moderately well enlarged and with moderate-sized transparent without distinct pair of rounded depressions. Lamella anteva- central 'window'. Hind wing of male with anal margin little or ginalis broad and rounded, without pair of lobes or median not expanded, without pronounced hair-scale tuft. Male abdo- indentation. Antrum without well-defined projection. Signum men less narrow and less elongate than usual in Ischnopteris; a moderately small short oval to disc-shape. sternite 3 with patch of setae very weak or absent. 66 Linda M. Pitkin

MALE GENITALIA (Figs 116, 173). Uncus a narrow, 3.xii.1999 (Brehm), 1 , 2110 m, 13.v.1999 (ID#12495) pointed, tongue-shape, not hood-shaped; with only sparse (Brehm), 1 , 2130 m, (ID#3507) 14.iv.1999 (Brehm, ventral setae, not a tuft. Gnathos with two well-separated and Fiedler, Sussenbach¨), 1 , 2130 m, 5.v.1999, (ID#3508) diverging median processes, with irregularly wavy anterior (Brehm), 2 , 3°58.85'S 79°05.01'W, 2112 m, 2.i.2000 margin. Valva with small, triangular apical costal lobe; vent- (ID#13078,13079) (Sussenbach¨); 2 , data as holotype but ral process of sacculus minute, occasionally absent. Processes Rio San Francisco, 1850 m, 3.xii.1999 (Bartsch & Hauser¨); of anellus claw-like, evenly curved, tapered and pointed. Me- 1 , 1 , data as holotype but forest near Rio San Fran- dian process of juxta slightly curved to right (in ventral view), cisco, 3°58.45'S 79°04.73'W, 1800 m, larva on Escal- nearly 1.5 times length of main part of juxta; with spinules lonia paniculata, 23.x.2000, pupated 1.xii, moth emerged in apical two-thirds. Aedeagus with spinose region situated 28.xii.[2000], larva on Tabebuia chrysantha, 15.x.2000, opposite and subapically to posterior tongue, which has no pupated 26.xi, moth emerged 28.xii.[2000] (Brehm) (all of spinules; vesica with row of short spines on sclerite, which is above SMNS, except as follows: 2 to BMNH, 1 to not on distinct caecum. AMNH, 1 to CMNH, 1 to USNM); 1 , data as holotype but 2112 m, Site 6a (4), 03°58.85'S 79°05.01'W, FEMALE GENITALIA. Unknown. 6.xi.1999, DNA Tax 3061, (Brehm) (ZSM); 1 , 2212 m, Site 7b (5), 03°58.72'S 79°04.44'W, 6.xi.1999, DNA Tax Diagnosis 3238 (Sussenbach¨) (SMNS). Peru: 1 , Tambillo (Jelski) Extensive variability in the wing markings of bipectinata (BMNH). means that there are no obvious reliable distinguishing features there, but the species is notable in being the only mem- Excluded from the type series: 1 , Bolivia: Cochabamba ber of Ischnopteris with strongly bipectinate male antennae. (CMNH); this additional specimen differs in some characters I. bipectinata is atypical in some other respects: moths are on and it is not certain that it is conspecific. average the smallest in the genus, and in the male genitalia the uncus has only sparse ventral setae, not a tuft. The structure of the gnathos is putatively autapomorphic for the species, Description in having the combination of features: median processes well ADULTS (Figs 38, 39). Fore wing length: , 21-25 mm; , separated, diverging, hook-tipped, and with irregular anterior 25 mm. Wing pattern: fore wing of both sexes mottled dark to edge. The only other species in which paired processes di- mid brown, with diffuse patches of green, and some whitish verge as much is bifinita (a markedly different species, in the flecks, male usually with faintly paler medial region, some- fabiana-group), which does not have hook-tipped processes. times more distinct and forming a slanting transverse band, and occasionally with whitish blotch near midpoint of costa, plus several smaller whitish markings; fore wing of female (single Biological notes specimen known) with roughly diamond-shaped central area Moths have been found at altitudes of 50 m and 1500 m, in paler. Hind wing of both sexes greyish brown, with apical or- February and May. ange or yellow region sharply angled along margin with brown region of wing; female with dark brown streaks and blotch Distribution clearly visible against greyish ground, and extending from Ecuador. margin of orange/yellow region to near tornus. Under-side: fore wing with slanting transverse orange band, usually dull in Ischnopteris brehmi sp. nov. male but bright in female, and with white blotch at midpoint Figs 38, 39,117,174, 223 of outer margin (similar but less conspicuous blotch, whitish or yellowish, usually on hind wing); wings with dark brown, Downloaded by [Michigan State University] at 11:29 24 December 2014 Ischnopteris sp. near chryses Druce: Brehm, 2003: Table 1, approximately medial, line, more distinct on fore wing; hind and Plate 1, Fig. 3 (larva). wing greyish brown with straw-coloured apical patch. Dorsum of male fore wing without distinct median tuft. Male retinacu- Material examined lum usually slightly to moderately enlarged, and with small 36 , 1 (including 4 , 1 genitalia preparations) transparent central 'window'. Hind wing of male with anal margin strongly expanded, forming distinct flap, with hair- Holotype , Ecuador: Zamora-Chinchipe, Estacion Cientifica scale tuft. San Francisco, 3°58'S 79°04'W, 1780 m, 26.xii.1999 (ID # 3504) (Sussenbach¨) (genitalia slide LMP 460; SMNS). MALE GENITALIA (Figs 117A-B, 174). Uncus only slightly Paratypes: Ecuador: 29 , data as holotype but: 1 , hood-shaped, or not hood-shaped; broadly digitate, with much 1780 m, 26. xii.1999 (ID#3524) (Sussenbach¨), 3 , smaller digitate apex straight (i.e. not bent over). Gnathos 1845 m, 9,11.x.1999 (Brehm), 9 , 1910 m, 20.iv.1999 with two median processes. Valva with digitate apical costal (ID#3509), 30.xii.1999 (ID#3503, 3505, 3506, 3516, process. Processes of anellus blade-like, with narrow pointed 3517, 3519, 3521, 3522) (Sussenbach¨), 5 , 1960 tip. Median process of juxta similar in length to main part m, 13.v.1999 (ID#3518, 3523), 10.x.1999, 30.xii.1999 of juxta, curved to right (in ventral view), spinulose only at (ID#3520) (Brehm), 5 , 2040 m, 18.v.1999 (ID#3510, and around apex. Aedeagus with a few triangular spinules (2— 3511, 3512, 3513, 3525) (Sussenbach¨), 1 , 2040 m, 4 spinules present in specimens examined) on posterior tongue; Neotropical geometrid moths 67

vesica with long, narrow, pointed, finely spinulose sclerite, not Material examined on distinct caecum. 6 (including 4 genitalia preparations) (AMNH, BMNH, MNHN, ZSM, USNM). FEMALE GENITALIA (Fig. 223). Lamella postvaginalis with pair of small rounded depressions indistinct or absent. Lamella Holotype (catocalata), Brazil: no further data (MNHN) antevaginalis forming a broad band without distinct lobes or [photograph examined, abdomen missing]. median indentation, but with strong wrinkles on surface adjoin- Holotype (prognata), Bolivia: Rio Songo, 750 m (Fassl) ing antrum. Antrum with posterior digitate projection. Bursa (Type No. 32547, genitalia slide 60135, USNM). copulatrix long and slender; signum a short oval with base less than half length of signum. Localities of additional material. Peru: Carabaya, La Union, R. Huacamayo. ´Bolivia: [Cochabamba:] Chapare, San Jacinto; Yungas de Palmar; Rio Songo. Locality not traced: Diagnosis Esp. Santo. Ischnopteris brehmi is closely related to albiguttata, and the two species have very similar genitalia. But brehmi is markedly Description different in wing pattern, and bears a superficial resemblance ADULTS (Fig. 40). Fore wing length: , 20-23 mm. Wing to some members of the chryses-group, with an orange region pattern (only female known): fore wing mottled pale to dark present on the hind wing, and on the under-side of the fore brown, with central area, roughly diamond-shaped, either paler wing. Where the orange is dull, moths of brehmi may resemble but not well defined, or darker; hind wing mainly orange, with those offassli, but the colour of the under-side of the hind wing dark brown bar from tornus to middle of outer margin of wing, (predominantly greyish brown in brehmi and mainly yellowish with short dark brown line adjacent to inner margin of bar and straw in fassli) usually distinguishes those two species, which sometimes partly merging with it, and with no more than a trace are in any case allopatric. I. chryses, a similar-looking species of dark brown at apex of wing; without discal spot. Under-side which is sympatric with brehmi, differs in usually having a of fore wing with large orange region bordered by slanting less sharply angled margin to the orange region on the hind subapical dark brown transverse band; brown postmedial or wing. Males of those species are readily distinguished by the medial line broken or absent. absence of a distinct median tuft on the dorsum of the fore wing in brehmi, and presence of that in the chryses-group. Moths of MALE GENITALIA. Unknown. brehmi also resemble those of palmeri (only males known) but the latter have a larger 'window' in the male retinaculum. The FEMALE GENITALIA (Figs 224A, B). Lamella postvaginalis male genitalia of brehmi are similar to those of albiguttata, without distinct pair of rounded depressions. Lamella anteva- especially in the blade-like shape of the processes of anellus, ginalis with pair of low rounded lobes separated by shallow but brehmi differs in the process of the juxta, which is curved, median indentation (occasionally very weak). Antrum longer not straight, and spinulose only near the apex. The female than broad or of similar length to breadth, without marked genitalia differ from those of albiguttata only in the size of the lateral indentation as is present in pronubata; with posterior base of the signum. This is not confirmed as a reliable character, projection digitate or tapered but not sharply pointed. Bursa as only one female of brehmi is available for examination. copulatrix sometimes fairly bulbous anteriorly but more elongated than in pronubata (compare Figs 224 and 230A); signum oval, smaller than that of pronubata. Biological notes Moths have been found at altitudes of 1780-2212 m, from Diagnosis April to June, and October to December. Larvae have been Moths of catocalata (only females known) resemble those of reared from the following hostplants: Bignoniaceae: Tabebuia pronubata and stenoptila in having a very extensive orange Downloaded by [Michigan State University] at 11:29 24 December 2014 chrysantha (Jacq.) G. Nicholson, and Grossulariaceae: Escal- region on the hind wing, but without a distinct dark blotch or lonia paniculata (Ruiz & Pav.) Roem. & Schult., by Brehm band at the apex of the wing, as is present in pronubata, and (2003). without a discal spot, as is present in stenoptila. I. catocalata, but not stenoptila, has a dark brown line alongside the dark brown bar on the hind wing. I. stenoptila has a longer fore wing Distribution than the other two species, but it is currently known only from Ecuador and Peru. a single female. The female genitalia of catocalata differ from those of pronubata in several characters as described above: the form of the lamella antevaginalis, antrum, bursa copulatrix, Ischnopteris catocalata (Guenee) and signum. Differences from stenoptila are more subtle: the Figs 40, 224 lamella antevaginalis often has more obvious lobes than in Syrtodes catocalata Guenee, [1858]: 454; pl. 14, fig. 8. Holo- that species, and the shape of the antrum and its extension (as type , BRAZIL (MNHN) [photograph examined]. described above) differ. Ischnopteris prognata Dognin, 1923: 19. Holotype , BOLIVIA (USNM) [examined]. Syn. nov. Biological notes Ischnopteris catocalata (Guenee); Parsons et al., in Scoble, Moths have been found at altitudes of 610 m to 2000 m, in 1999: 519. April and November. 68 Linda M. Pitkin

Distribution chlorata is the only species that has the processes of the anellus Peru, Brazil, and Bolivia. bifurcate with prongs well apart (the prongs are close together in bifurcata and hirsuta). The single known female specimen is Ischnopteris chlorata Hubner associated only tentatively with chlorata. The female genitalia Figs 41, 42,118,175, 225 (damaged in this specimen) are somewhat similar to those of albiguttata, but with weaker wrinkles on the lamella anteva- Ischnopteris chlorata Hubner, [1823] 1806: pl. [222], ginalis, and also to bifurcata, but with a longer signum than figs 1—4. Syntypes 2 , 2 , no further data [not traced]. the latter. Identity of species established by Rindge, 1983: 227.

Biological notes Material examined Moths have been found at an altitude of 200 m, in November 8 , 1 (including 4 , 1 genitalia preparations) (BMNH). to December. Localities. Venezuela: Merida. Guyana: no further data; confluence of Oroneque and New Rivers; Rio Demerara. Sur- Distribution inam: no further data. French Guiana: St Jean du Maroni. Northern lowland regions of South America. Ecuador: Sarayacu.

Description Ischnopteris chloroclystata (Guenee) ADULTS (Figs 41, 42). Fore wing length: , 20-23 mm; , Figs 43,44,119,176, 226 23 mm. Wing pattern: fore wing of both sexes mottled mid to Syrtodes chloroclystata Guenee, [1858]: 452. LECTOTYPE dark brown and straw colour, with faint short striations particu- , BRAZIL (BMNH), here designated [examined]. larly near the costa and outer margin; darker brown postmedial Ischnopteris chloroclystata chloroclystata (Guenee); Parsons line slanting straight from costa to vein M3 (angled or indistinct etal., inScoble, 1999:519. from there on), but usually not very distinct in males; fore wing of female with roughly diamond-shaped central area (slightly Material examined paler in single specimen examined). Hind wing brown without 7 , 3 (including 3 , 2 genitalia preparations) (BMNH, orange region, with indistinct darker brown postmedial line VOB). and submarginal band; under-side of male hind wing with darker submarginal band more distinct. Dorsum of male fore Lectotype , Brazil: no further data (genitalia slide Geom: wing without distinct median tuft. Male retinaculum strongly 21045, BMNH). Paralectotypes: Brazil: 1 , 2 , no further enlarged and with transparent central 'window'. Hind wing data (BMNH). of male with anal margin not greatly expanded and without pronounced hair-scale tuft. Male abdomen with tufts evenly Localities of additional material. Brazil: no further data; rounded. Rio de Janeiro; R[io de] J[aneiro]: NovaFriburgo; Teresopolis; Sao Paulo: Alto da Serra. MALE GENITALIA (Figs 118, 175). Uncus narrow and curved, only slightly hood-shaped, tapered to pointed apex. Gnathos with one median process, with hooked tip. Valva with Description digitate apical costal process. Processes of anellus bifurcate ADULTS (Figs 43, 44). Fore wing length: , 19-23 mm; , from base with slender rod-like prongs well apart; situated 23 mm. Wing pattern: fore wing mottled brown, mainly dark posteriorly in anellus. Median process of juxta longer than in male and with some indistinct olive-greenish markings, par-

Downloaded by [Michigan State University] at 11:29 24 December 2014 juxta proper, with spinules in apical half to two-thirds; pro- ticularly at base of wing and a diffuse, broken and irregular, cess deflected to right (in ventral view), but sometimes only approximately medial, transverse band; female with roughly slightly. Aedeagus with group of several short spinules on post- diamond-shaped central area, and outer patch, either pale to erior tongue; vesica with short spinulose sclerite, not rounded tan brown with dark brown flecks, or dark brown, other areas at tip, at end of long caecum. of wing of female contrasting somewhat, either mainly darker brown or olive-green. Hind wing mid to dark brown, almost FEMALE GENITALIA (Fig. 225). Lamella postvaginalis plain (male), or with some markings but not particularly dis- without distinct pair of rounded depressions. Lamella ante- tinctive (female), without orange region. Under-side of each vaginalis forming a broad band without lobes or median in- wing, in both sexes, with cream or whitish blotch at midpoint dentation, with light wrinkles on surface adjoining antrum. of outer margin, weaker on hind wing, and fore wing with Antrum with posterior projection bulbous and lightly sclerot- small pale apical or subapical marking. Dorsum of male fore ized. Signum a fairly long oval. wing without distinct median tuft. Male retinaculum enlarged and with moderately large transparent central 'window'. Hind Diagnosis wing of male with anal margin moderately expanded and with The fore wing pattern of chlorata is fairly plain, but male hair-scale tuft slight to moderate. moths can sometimes be recognised by the evenly rounded tufts on the male abdomen. The species is better distinguished MALE GENITALIA (Figs 119, 176). Uncus only slightly and defined by a putative autapomorphy of the male genitalia: hood-shaped; broadly digitate, with much smaller digitate apex Neotropical geometrid moths 69

straight (i.e. not bent over). Gnathos with two median pro- Biological notes cesses, with curved tip. Valva with a digitate apical costal pro- The holotype of juvencata is a female specimen that has lost its cess, with a distinct rounded lobe where this process forms an abdomen. It has a wrongly associated abdomen glued on to it; angle with the costal margin. Processes of anellus narrow and that abdomen does not have the characters of Ischnopteris or spine-like with blunt apex, each process with broader, curved any closely related genus, and it may not even be a nacophor- base, extending over anellus as a large sclerotized region from ine. No other specimens are associated with juvencata and it is the posterior end of which a shorter spine arises. Median pro- impossible to determine its identity, other than that it belongs cess of juxta short and tapered (much shorter than main part in Ischnopteris, thus I am treating it as a nomen dubium. of juxta), without spinules. Aedeagus with a group of small spinules and one large spine (not a broad triangle) on posterior tongue; vesica with a long narrow sclerite, not rounded at tip, Ischnopteris fasciata sp. nov. not on a distinct caecum. Figs 45,120,177 Material examined FEMALE GENITALIA (Figs 226A, B). Lamella postvaginalis 3 (including 2 genitalia preparations) without distinct pair of rounded depressions. Lamella antevaginalis forming a narrow band without pair of lobes or median Holotype , Brazil: R[io de]J[aneiro], Nova Friburgo, 1100 indentation. Antrum with posterior projection tapered but not m, 21.i. 1998 (Becker) (genitalia slide LMP 407; VOB). appearing very distinct. Signum approximately disc-shaped, Paratypes: Brazil: 1 , data as holotype (VOB); 1 , Santa with base almost as large as signum. Catarina, Jaragua do Sul, ix.1932 (Hoffman) (BMNH).

Diagnosis Description Male moths of chloroclystata can often be recognised by the ADULTS (Fig. 45). Fore wing length: , 19-20 mm. Wing greenish band across the fairly plain dark brown fore wing, pattern (only male known): fore wing mottled dark brown even though this is a very subtle character due to the indistinct with some diffuse olive-green markings, particularly at base appearance of the band. Genitalia characters are more defin- of wing and postmedially, with irregular white band transverse itive however. Ischnopteris chloroclystata is near aurudaria and slanting; hind wing dark greyish brown, almost plain and in male and female genitalic features, although clearly distin- without orange region. Under-side of fore wing with diffuse guished from that species by the absence of an orange region whitish blotch at midpoint of outer margin, and tiny pale subap- on the hind wing. The form of the processes of the anellus (as ical marking, but these markings often indistinct. Dorsum of described above) is a putative synapomorphy of chloroclys- male fore wing with distinct median tuft. Male retinaculum tata, fasciata, and aurudaria, but chloroclystata differs from strongly enlarged and with large transparent central 'window'. the other two species in having a large, but not broad and Hind wing of male with anal margin moderately expanded and triangular, spine amongst the spinules on the aedeagus. Also, with hair-scale tuft slight to moderate. chloroclystata differs from aurudaria in having a lobe clearly developed on the costa at the base of the digitate process of MALE GENITALIA (Figs 120,177). Uncus not hood-shaped; the valva. Female genitalia of chloroclystata and aurudaria broader than that of chloroclystata, forming right angle with are very similar, and share the character of the signum with a extremely narrow rod-like apex straight (i.e. not bent over). large base (but not in the centre of the signum as is the case in Gnathos with two median processes, with curved tip. Valva pronubata). with digitate apical costal process curved and irregular, with As I am treating chloroclystata juvencata as a nomen distinct lobe where this process forms an angle with the costal dubium (see below), there are effectively no subspecies of margin. Processes of anellus narrow and spine-like with blunt Downloaded by [Michigan State University] at 11:29 24 December 2014 chloroclystata. apex, each process with broader, curved base, extending over anellus as large sclerotized region from posterior end of which a shorter spine arises. Median process of juxta short and Biological notes tapered, triangular (much shorter than main part of juxta), Moths have been found at altitudes of 1000 m to 1100 m, in without spinules. Aedeagus with group of spinules, but without January, and September to November. a strong spine, on posterior tongue; vesica with small sclerite, not rounded at tip, not on caecum. Distribution FEMALE GENITALIA. Unknown. Brazil.

Diagnosis Ischnopteris chloroclystata juvencata (Guenee) Ischnopteris fasciata appears to be closely related to chloro- nomen dubium clystata and aurudaria but differs from them in several re- Syrtodes juvencata Guenee, [1858]: 453. Holotype $, spects. Moths of fasciata do not have orange on the hind wing BRAZIL?(BMNH) [examined]. as those of aurudaria do, and males of fasciata have a dis- Ischnopteris chloroclystata juvencata (Guenee); Parsons tinct median tuft on the dorsum of the fore wing, unlike males et al., in Scoble, 1999:519. of the other two species. The white band on the fore wing 70 Linda M. Pitkin

is conspicuous and putatively autapomorphic for fasciata, as nata. The latter species (a member of the chryses-group) is are characters of the genitalia, notably the very narrow apical the only other species with a median tuft on the dorsum of rod of the uncus, the right angle between this and the broader the fore wing and without orange on the hind wing, except part of the uncus, and the small sclerite of the vesica of the for fasciata, which is readily distinguished by the white band aedeagus. on its fore wing. The male genitalia of hirsuta and obtortionis have a feature in common with those of latijuxta, in that the Biological notes uncus has a median process between two lobes. However, only Moths have been found at an altitude of 1100 m, in January hirsuta and obtortionis share the conspicuous synapomorphy and September. of strong development of the patch of dorsal hair scales, which in these two species are capitate, on the valva. I. hirsuta is defined and distinguished from obtortionis primarily by the Distribution bifurcate processes of the anellus, as described above. Also, Brazil. hirsuta has a smaller basal curve on the median process of the uncus than obtortionis. Ischnopteris hirsuta sp. nov. Figs 46,121,178 Biological notes Material examined The single known moth was found at an altitude of 1100 m, in 1 (including genitalia preparation) January. Holotype , Bolivia: La Paz, from Caranavi to Santa Ana on Alto Rio Beni road, 1100 m, 9.i. 1976 (Pena˜) (genitalia slide Distribution 18963; AMNH). Bolivia.

Description Ischnopteris latijuxta sp. nov. ADULTS (Fig. 46). Fore wing length: ,19 mm. Wing pattern Figs 47,122,179 (only male known and in poor condition): fore wing mottled Material examined brown, with dark markings on median lobe of dorsum; hind 2 (genitalia preparations of both) wing brown without orange region, ground colour pale, with darker postmedial line and diffuse band at outer margin, and Holotype , Brazil: D[istrito] F[ederal], Planaltina, 15°35'S with pronounced dark patch at anal margin. Under-side of 47°42'W, 1000 m, 25.ix.1985 (Becker) (genitalia slide LMP wings pale but similar to upper side. Dorsum of male fore 473; VOB). wing with median lobe and tuft. Male retinaculum moderately Paratype: 1 , data as holotype but 15.ii.1982 (VOB). enlarged and with fairly large transparent central 'window'. Hind wing of male with anal margin strongly expanded form- Description ing distinct flap, and with hair-scale tuft. ADULTS (Fig. 47). Fore wing length: ,16-18 mm. Wing pat- MALE GENITALIA (Figs 121,178). Uncus not hood-shaped; tern (only male known): fore wing mottled dark brown with or moderately narrow, apex with small median process between without several small diffuse white blotches; with thin black pair of lobes; this process rod-like but sinuous with basal medial line and zigzag postmedial line; hind wing slightly curve small. Gnathos with two median processes, with slightly paler brown with dark brown medial (or postmedial) line and hooked tip. Valva with digitate apical costal process, with hair- outer margin, sometimes indistinct, without orange region. like tuft; valva with patch of dorsal hair scales very large and Dorsum of male fore wing without median tuft. Male retin- Downloaded by [Michigan State University] at 11:29 24 December 2014 dense, hair scales very long and fine but with tips capitate. aculum slightly enlarged and with small transparent central Processes of anellus bifurcate with tapered prongs close to- 'window'. Hind wing of male with anal margin little expanded gether; one prong very short and squat. Median process of and without pronounced hair-scale tuft. juxta tapered, and curved slightly to right (in ventral view), MALE GENITALIA (Figs 122, 179). Uncus only slightly approximately 1.5 times length of main part of juxta, with hood-shaped, moderately narrow; apex with small median rod- spinules mainly in apical half. Aedeagus with group of short like process almost at right angles to uncus or bent over more, spinules on posterior tongue; vesica with narrow, finely spin- situated between pair of lobes. Socii well developed, semi- ulose sclerite, not rounded at tip. rigid. Gnathos almost a simple loop, with or without pair of very slight median projections. Valva with curved and pointed FEMALE GENITALIA. Unknown. digitate apical costal process. Processes of anellus narrow and triangular. Median process of juxta large and very broad, some- Diagnosis what crescent-shaped, covered in spinules. Aedeagus with pos- Ischnopteris hirsuta is very similar to obtortionis. Both spe- terior tongue, without spinules anywhere on sclerotized part cies have characteristic wing markings (in the males) - dark of aedeagus; vesica with moderately short sclerite, rounded at markings situated on the median lobe of the dorsum of the tip, not spinulose, on very short or indistinct caecum. fore wing and the anal margin of hind wing. These are much more pronounced, against the pale hind wing, than in inor- FEMALE GENITALIA. Unknown. Neotropical geometrid moths 71

Diagnosis and a conspicuously darker submarginal band on the under- The fore wing pattern of latijuxta is variable but the species is side of the hind wing. The male genitalia have in common clearly defined by putative autapomorphies of the male gen- with those of miseliata the unusual feature of a long-spined italia. Although the uncus is similar to that of obtortionis in sclerite situated at the end of a long caecum, in the vesica of having a median process between two lobes, latijuxta is the the aedeagus, but the unique shape of the uncus, with strong only species of the genus to have a simple loop for a gnathos, subapical swelling and slender apical rod, and the processes and the broad shape of the median process of the juxta is of the anellus of lemoulti (as described above) are putatively unique. autapomorphic for the species.

Biological notes Distribution Moths have been found at an altitude of 1000 m, in February Brazil. and September. Ischnopteris miseliata (Guenee) Distribution Figs 49, 50, 99,158, 212 Brazil. Syrtodes miseliata Guenee, [1858]: 453. Holotype ?, BRAZIL Ischnopteris lemoulti sp. nov. (BMNH) [examined]. Figs 48,123,180 Ischnopteris inconspicua Warren, 1907: 288. LECTOTYPE , PARAGUAY (BMNH), here designated [examined]. Material examined [Synonymy cited by Parsons et al., in Scoble, 1999: 520.] 1 (including genitalia preparation) Ischnopteris miseliata (Guenee); Parsons et al., in Scoble, Holotype , Brazil: Minas Gerais: Uberaba, (Le Moult Coll.) 1999: 520. (genitalia slide Geom: 21487; BMNH).

Material examined Description 14 , 6 (including 5 , 3 genitalia preparations) (BMNH, ADULTS (Fig. 48). Fore wing length: , 18 mm. Wing pat- VOB, ZSM). tern (only male known): fore wing dark brown mottled with grey-green-tinged mid brown, with faint short striations par- Holotype (miseliata), Brazil: no further data (BMNH) ticularly on the costa; and with small whitish but rather diffuse [wrong abdomen glued on]. markings near apex and towards tornus. Hind wing mid brown Lectotype (inconspicua) Paraguay: Sapucay, x.1904 with faint darker brown postmedial line, and with diffuse dark (Foster) (genitalia slide Geom: 21068, BMNH). Para- band at or near outer margin but usually indistinct in male; lectotype?1 , Paraguay: Sapucay, 26.x. 1904 (Foster) without orange region. Dark discal spot distinct on upper and (BMNH). under-side of hind wing, without markings very close to it. Under-side of hind wing pale brown with contrasting dark Localities of additional material. Brazil: Edo. Espirito submarginal band. Dorsum of male fore wing without median Santo: St Leopoldino, Dorf Tirol; M[inas] G[erais]: Nova tuft. Male retinaculum moderately enlarged and with trans- Lima; Parana: Castro; Rio Grande d[o] Sul: Hamburgo parent central 'window'. Hind wing of male with anal margin Velho; Rio [de] Janeiro Prov[ince]: Laguna de Sacuarema; little expanded and without pronounced hair-scale tuft. Santa Catarina: no further data; Jaragua do Sul; Sao Paulo; S[ao]P[aulo]: Bertioga. ˜Paraguay: Sapucay; Sapucay, near MALE GENITALIA (Figs 123,180). Uncus not hood-shaped; Villarrica.

Downloaded by [Michigan State University] at 11:29 24 December 2014 uncus with narrow basal part gently broadening into rounded part, then narrowing strongly to short slender apical rod. Gnathos with one median process, with hooked tip. Valva with Description digitate apical costal process. Processes of anellus with large, ADULTS (Figs 49,50). Fore wing length: ,16-20 mm; , 21- broad, plate-like base and moderately long narrow spine-like 22 mm. Wing pattern: fore wing of both sexes mottled dark apex. Median process of juxta slightly less than 1.5 times to mid brown, with paler patches sometimes green-tinged, and length of main part of juxta, curved and deflected slightly with faint short striations particularly on the costa; male with to right (in ventral view), with spinules, many long, mainly dark brown spot, often indistinct, near tornus; fore wing of fe- in apical half. Aedeagus with numerous short spinules and male with roughly diamond-shaped central area darker, paler, one larger triangular spine at base of posterior tongue; vesica or not clearly defined. Hind wing brown without orange re- with sclerite of long slender spines and spinules, sclerite not gion, with darker brown lines or bands often distinct in female rounded at tip, situated at end of fairly long caecum. but usually indistinct in male; postmedial line of female distinctly angled and almost touching dark submarginal band at FEMALE GENITALIA. Unknown. or near vein M3. Under-side of hind wing, of males and some females, with dark discal spot, sometimes faint but with no Diagnosis other distinct markings anywhere near it. Dorsum of male fore Moths of lemoulti (a single male known) resemble those of wing without median tuft. Male retinaculum strongly enlarged miseliata, but with a smaller male retinaculum than the latter, and with transparent central 'window'. Hind wing of male with 72 Linda M. Pitkin

anal margin little expanded and without pronounced hair-scale Paz,] Sarampiuni, San Carlos; Las Juntas; Santiago del tuft. Estero. MALE GENITALIA (Figs 124, 181). Uncus usually slightly Description hood-shaped, narrow throughout its length, gently curved, only ADULTS (Figs 51,52). Fore wing length: , 21-23 mm; , 24- slightly enlarged subapically, then tapered sharply to pointed 25 mm. Metathorax dorsally with small dark brown longitud- apex. Gnathos with one median process, with hooked tip vary- inal dash. Wing pattern: fore wing dark to mid brown, mottled ing from distinct to very slight. Valva with pointed apical costal with greyish-green markings, in male with well-marked dark process. Processes of anellus claw-like, evenly curved and brown lines forming a series of longitudinal dashes mainly tapered. Median process of juxta approximately, or slightly in outer half of wing; in female with whitish markings; fe- less than, 1.5 times length of main part of juxta, deflected male with roughly diamond-shaped central area moderately slightly to right (in ventral view), angled or curved mid length, or poorly defined. Hind wing brown without orange region; with spinules in apical half, mainly along one edge. Aedeagus with dark brown postmedial line, and usually with dark band with numerous short spinules on posterior tongue; vesica with at outer margin or submarginal, usually with paler band in long sclerite of slender spines and spinules, sclerite not roun- between these. Dorsum of male fore wing without median ded at tip, situated at end of long caecum. tuft. Male retinaculum slightly enlarged, usually with weakly FEMALE GENITALIA (Figs 227 A, B). Lamella postvaginalis developed transparent central 'window'. Hind wing of male without distinct pair of rounded depressions. Lamella anteva- with anal margin strongly expanded, forming a distinct flap, ginalis with pair of low lobes separated by shallow rounded with pronounced hair-scale tuft. median indentation. Antrum with posterior projection truncate MALE GENITALIA (Figs 125, 182). Uncus sinuous later- at apex. Signum elongate and pointed, leaf-shaped. ally, strongly swollen before very much narrower but spatulate apex, which is shallowly indented at tip; base of uncus ex- Diagnosis tended and slightly overlapping tegumen dorsally, with pair Moths of miseliata are not readily distinguishable by external of patches of long hair-scales. Gnathos with two median pro- features, particularly in the male. Females may be distin- cesses, with weakly or barely curved tip. Valva with costa guished from those of some other similar-looking species by curved or angled moderately or strongly, but never quite as the angled postmedial line, almost touching the submarginal much as right-angled; with small curved or bent digitate ap- band of the hind wing. In the male genitalia, the long-spined ical costal process. Processes of anellus elongate and tapered, sclerite situated at the end of a long caecum occurs only in usually curved and subtriangular, with blunt apex. Juxta miseliata and lemoulti, but the uncus is very different, nar- without distinctly developed median process. Aedeagus with row throughout its length in miseliata but with strong subap- several spinules on posterior tongue; vesica with narrow, lon- ical swelling in lemoulti. In the female, the elongate, pointed, gitudinally striated sclerite, rounded at tip, not on distinct leaf-like shape of the signum is putatively autapomorphic for caecum. miseliata. FEMALE GENITALIA (Fig. 228). Lamella postvaginalis with pair of rounded depressions indistinct or absent. Lamella Biological notes antevaginalis without distinct lobes or median indentation. An- Moths have been found at altitudes of 5-950 m, in June, July, trum without obvious posterior projection at apex. Signum a and October to December. large disc or oval, with base much smaller than signum.

Distribution Diagnosis Downloaded by [Michigan State University] at 11:29 24 December 2014 Brazil and Paraguay. Male moths of multistrigata have stronger and more distinctive fore wing markings of dark dashes than those seen in other Ischnopteris multistrigata Warren species. Both sexes can be distinguished from moths of other Figs 51, 52,125,182, 228 species by the presence of the dark dash on the metathorax. In the male genitalia, the shape of the apical process of the Ischnopteris multistrigata Warren, 1909: 103. Holotype , uncus (spatulate with a slightly indented tip) is characteristic, [ARGENTINA] (BMNH) [examined]. autapomorphic for the species. Ischnopteris multistrigata is unusual in having no median process of the juxta; seriei is Material examined similar but multistrigata differs in that the gnathos processes 6 , 2 (including 4 , 2 genitalia preparations) (AMNH, and the costa of the valva are less curved. In the female, the BMNH, CMNH, ZSM). signum is characteristic in its large size and rounded shape with much smaller base, and the base not central as in klagesi Holotype , [Argentina, stated erroneously as 'East. or pronubata. Bolivia':] Santiago del Estero, 1905-1906 (Steinbach) (genitalia slide Geom: 21197, BMNH). Biological notes Localities of additional material. Peru: Junin, Satipo. Moths have been found at altitudes of 400 m to 1000 m, in Bolivia: Buenavista; [Cochabamba,] Chapare region; [La April, July, August, and December. Neotropical geometrid moths 73

Distribution markings situated on the median lobe of the dorsum of the Peru, Bolivia, and Argentina. fore wing and the anal margin of hind wing. These are much more pronounced against the pale hind wing than in inornata. Ischnopteris obtortionis (Prout) The latter species (a member of the chryses-group) is the only other species with a median tuft on the dorsum of the fore Figs 53, 54,126,183 wing and without orange on the hind wing, except for fasci- Ischnopterix obtortionis Prout, 1928: 57. LECTOTYPE , ata, which is readily distinguished by the white band on its BOLIVIA (BMNH), here designated [examined]. fore wing. The male genitalia of obtortionis and hirsuta have Ischnopteris obtortionis (Prout); Parsons et al., in Scoble, a feature in common with those of latijuxta, in that the un- 1999: 520. cus has a median process between two lobes. However, only obtortionis and hirsuta share the conspicuous synapomorphy Material examined of strong development of the patch of dorsal hair scales, which 3 , 1 (including 3 genitalia preparations) (BMNH, in these two species are capitate, on the valva. I. obtortionis CMNH). is distinguished from hirsuta primarily by the slender, curved spine-like processes of the anellus, which are not bifurcate as Lectotype , Bolivia: E. Bolivia, Buenavista, 750 m, in hirsuta. Also, obtortionis has a larger basal curve on the viii.1906-iv.1907 (Steinbach) (genitalia slide Geom: 21023, median process of the uncus than hirsuta. The single known BMNH). Paralectotype, 1 , data as lectotype (BMNH). female specimen of obtortionis (from Peru) is in poor condition and has lost its abdomen; it is tentatively associated with Localities of additional material. Bolivia: Buenav- this species. ista; P[rovincia] del Sara. Peru: Carabaya, La Union, R. Huacamayo. Biological notes Description Moths have been found at an altitude of 750 m, in December. ADULTS (Figs 53, 54). Fore wing length: , 18-20 mm; ,18 mm. Wing pattern: fore wing of both sexes mainly Distribution mottled brown with paler flecks, with dark marking on me- Bolivia, and probably also Peru. dian lobe of dorsum (in male); hind wing brown without orange region, ground colour pale, with darker postmedial Ischnopteris ochroprosthia (Prout) line and diffuse band at outer margin (moderately distinct in Figs 55,127,184, 229 male but faint in sole female examined), and with pronounced dark patch at anal margin (in male). Under-side of wings of Ischnopterix ochroprosthia Prout, 1929: 69. Holotype , both sexes with dark markings contrasting with pale ground BRAZIL (BMNH) [examined]. colour. Dorsum of male fore wing with median lobe and tuft. Ischnopteris ochroprosthia (Prout); Parsons et al., in Scoble, Male retinaculum slightly to moderately enlarged and with 1999: 520 [type locality cited erroneously as Panama]. small transparent central 'window'. Hind wing of male with anal margin strongly expanded, forming a distinct flap, and Material examined with hair-scale tuft. 8 , 1 (including 2 , 1 genitalia preparations) (BMNH, MALE GENITALIA (Figs 126, 183). Uncus moderately USNM, VOB). narrow, not hood-shaped; apex with small median process Holotype , Brazil: Parana, Iguac´¸u, 26.x.1921 (genitalia slide between pair of lobes; this process rod-like but strongly sinu- Downloaded by [Michigan State University] at 11:29 24 December 2014 Geom: 20995, BMNH). ous with basal curve large. Gnathos with two median processes, with slightly hooked tip. Valva with apical costal pro- Localities of additional material. Brazil: Parana: Iguac´¸u; cess somewhat digitate but with pointed apex and with hair-like Ro[ndonia], Vilhena; Santa Catarina: no further data; Jaragua tuft; valva with patch of dorsal hair scales very large and dense, do Sul; Sao Paulo. ˜Bolivia: Buenavista. hair scales long and very fine but with tip capitate. Processes of anellus long, slender, spine-like and evenly curved. Median process of juxta tapered and more or less straight, often short Description (slightly shorter to slightly longer than main part of juxta), ADULTS (Fig. 55). Fore wing length: , 19-23 mm; , 29 with spinules mainly in apical half. Aedeagus with group of mm. Wing pattern: fore wing of both sexes dark brown with short spinules at base of posterior tongue; vesica with narrow, contrasting, well-defined, straw-coloured costal band, and with finely spinulose sclerite, not rounded at tip, on short caecum. fainter short straw-coloured line slanting from just below costa to outer margin of wing; hind wing pale to mid brown, without FEMALE GENITALIA. Unknown. orange region. Dorsum of male fore wing without distinct median tuft. Male retinaculum enlarged but not greatly so, and Diagnosis with fairly small transparent central 'window'. Hind wing of Ischnopteris obtortionis is very similar to hirsuta. Both spe- male with anal margin little expanded, without pronounced cies have characteristic wing markings (in the males) - dark hair-scale tuft. in Linda M. Pitkin

MALE GENITALIA (Figs 127, 184). Uncus slightly hood- colour. Dorsum of male fore wing without distinct median tuft. shaped, broadly digitate, with very small pointed apical pro- Male retinaculum enlarged and with large transparent central jection at right angles to uncus or bent over and directed an- 'window'. Hind wing of male with anal margin expanded, teriorly. Gnathos with one median process, with hooked tip. forming a flap, with slight hair-scale tuft. Valva without distinct apical costal lobe. Processes of anel- MALE GENITALIA (Figs 128, 185). Uncus slightly hood- lus long, slender and almost straight. Median process of juxta shaped, broadly digitate, with much smaller, narrow, digitate long, straight and slender (more than 1.5 times to more than apex straight (i.e. not bent over). Gnathos with two median twice length of main part of juxta), with spinules in apical processes, with curved tip. Valva with digitate apical costal half. Aedeagus with posterior tongue, without spinules any- process. Processes of anellus slender and tapered, each with a where on sclerotized part of aedeagus; vesica with moderately median lobe in the form of a wavy-edged plate; with one to short spinulose sclerite, not rounded at apex, on very short, two patches of minute spinules on sclerotized region at base of sometimes barely developed, caecum. each process of anellus. Median process of juxta only lightly FEMALE GENITALIA (Fig. 229). Lamella postvaginalis sclerotized and without spinules, or with very few inconspicu- without pair of rounded depressions. Lamella antevaginalis ous ones; process curved to right (in ventral view), similar in with a pair of low lobes separated by shallow V-shaped me- length to main part of juxta. Aedeagus with two groups of dian indentation. Antrum without posterior projection. Bursa short spines or spinules, one group on posterior tongue and the copulatrix long and slender, becoming sclerotized anteriorly, other group situated opposite and slightly subapically to the but ending in a membranous rounded sac. Signum absent. former; vesica with very slight, narrow, spinulose sclerite, not rounded at tip, not on distinct caecum. Diagnosis FEMALE GENITALIA. Unknown. Moths of ochroprosthia are clearly distinguished from those of other species of Ischnopteris by the well-defined pale costal band of the fore wing. In the male genitalia, the processes Diagnosis of the anellus are unique in shape (long, slender and almost The moths ofpalmeri (only males known) have a wing pattern straight). The female genitalia are also unique in the genus (as similar to that of chryses and brehmi, but they differ from far as is known), in not having a signum. chryses in having a longer fore wing, without a median tuft on the dorsum (a character present in the male only), and Biological notes from both species in having a large 'window' in the male retinaculum. The male genitalia of palmeri differ from those Moths have been found at an altitude of 600 m, in April and of the chryses-group in having a wavy-edged median plate on October. the processes of the anellus. That character is unique and an autapomorphy for palmeri. Distribution Brazil and Bolivia. Biological notes Ischnopteris palmeri sp. nov. Moths have been found at altitudes of 1100 m and 2450 m, in Figs 56,128,185 September. Material examined 3 (including 2 genitalia preparations) Distribution Colombia and Ecuador. Holotype , Ecuador: E. Ecuador, Alpayacu, Rio Pastaza, Downloaded by [Michigan State University] at 11:29 24 December 2014 1100 m ('3600 ft') (Palmer) (genitalia slide Geom: 21020; Ischnopteris pronubata (Felder & Rogen hofer) BMNH). Figs 57, 58,129,186, 230 Paratypes. Colombia: 1 , San Cayetano, 2450 m ('8000 ft'), ix. 1902 (BMNH). Ecuador: 1 , data as holo- Syrtodes pronubata Felder & Rogenhofer, 1875: pl. 131, type (BMNH). figs 25,25a. Holotype , [BRAZIL?] (BMNH) [examined]. Ischnopteris pronubata (Felder & Rogenhofer); Warren, 1909: Description 102). ADULTS (Fig. 56). Fore wing length: , 22-24 mm. Wing pattern (only male known): fore wing mottled dark brown and Material examined grey-green with very faintly paler medial region, sometimes 17 , 4 (including 7 , 3 genitalia preparations) (AMNH, with a few indistinct small whitish markings but not well- BMNH, CMNH, ZSM). defined white blotches; hind wing dark brown with large apical orange region. Under-side: fore wing with slanting transverse Holotype , Brazil?: Amazonas (Bates) (BMNH) [abdomen orange band, with dark brown band (towards outer margin of missing]. wing) and line (towards middle) and with white blotch at midpoint of outer margin; hind wing with dark brown line forming Localities of additional material. Guyana: no further a blotch near tornus but merging somewhat with brown ground data; Rio Potaro; Rio Potaro, Tumatumari. French Guiana: Neotropical geometrid moths 75

no further data; Godebert-Maroni; Cayenne area, Piste has one prong much shorter than the other. The two species Coralie; Nouveau Chantier; St Jean du Maroni. Ecuador: are very different in other respects: for example pronubata Napo, Rio Jatunyacu, near Tena, 10 km SW. [of] Pano. Peru: does not have a strong development of the patch of dorsal hair SE. Peru, Rio Inambari, La Oroya. Brazil: Amazonas, Fonte scales on the male valva, which are capitate in hirsuta, and Boa; Amaz[onas], Tefe.´ Bolivia: [Cochabamba,] Chapare. hirsuta has no orange markings. Chief characteristics of the female genitalia are the strongly bulbous bursa copulatrix and Description the large signum with a large and almost central base. ADULTS (Figs 57, 58). Fore wing length: , 17-20 mm; , 19-21 mm. Wing pattern: fore wing mainly mottled dark Biological notes brown, with diffuse white markings (in male: two blotches, Moths have been found at altitudes of 400 m to 920 m, in Feb- ill-defined and often joined almost forming a slanting, postme- ruary, March, May, July, September, October, and December. dial band; in female: a central area, roughly diamond-shaped). Distribution Hind wing with extensive orange region, and with dark brown South America from Guyana to Bolivia. band or blotches at outer margin (always clearly present both at apex and extending up outer margin from dorsum), male Ischnopteris seriei (Giacomelli) also with dark brown band along dorsum. Under-side of fore Figs 59, 60,130,187, 231 wing with large orange region bordered by slanting subapical dark brown band transverse; brown postmedial or medial line Ischnopteryx seriei Giacomelli, 1911: 39. Holotype ( ?, ?), present or absent. Dorsum of male fore wing with median Argentina: Tucuman or La Rioja [not traced]. tuft, usually slight or moderate. Male retinaculum strongly en- Ischnopteris seriei (Giacomelli); Parsons et al., in Scoble, larged and with large transparent central 'window'. Hind wing 1999: 520. of male with anal margin expanded and with slight to moderate hair-scale tuft. Material examined 18 ,10 (including 3 ,2 genitalia preparations) (AMNH, MALE GENITALIA (Figs 129, 86). Uncus no more than BMNH, USNM, ZSM). slightly hood-shaped, broadly digitate, with much smaller digitate apex straight (i.e. not bent over). Gnathos with two me- Localities. Argentina: Jujuy Prov[ince]: La Almona; dian processes, with weakly curved tip. Valva with digitate, Valle Grande; Tucuman Prov[ince]: no further data; Siambon; usually curved, apical costal process. Processes of anellus Sierra de la Ramada. gently curved, blade-like or narrow, with prominent basal bulge or lobe. Median process of juxta varying from shorter than, to Description same length as, main part of juxta, angled at least slightly to ADULTS (Figs 59, 60). Fore wing length: , 19-21 mm; , right (in ventral view), with spinules mainly in apical half. 22-23 mm. Head without dark brown dorsal line (as is present Aedeagus with group of triangular spines and spinules on in xylinata). Wing pattern: fore wing of both sexes mottled posterior tongue; vesica with long, narrow, finely spinulose dark brown, female with roughly diamond-shaped central area sclerite, not rounded at tip, not on distinct caecum. paler, sometimes mainly ivory coloured, or concolorous and marked out only by dark lines around it. Hind wing of both FEMALE GENITALIA (Figs 230A, B). Lamella postvaginalis sexes without orange region; hind wing paler than fore wing without pair of rounded depressions. Lamella antevaginalis and often predominantly ivory coloured, with dark brown mar- without distinct lobes or median indentation. Antrum with ginal or submarginal band and usually, although sometimes curved indentation on left side (viewed with ventral surface faint, postmarginal line. Dorsum of male fore wing without uppermost); antrum with tapered pointed posterior projection. median tuft. Male retinaculum enlarged and with small to Downloaded by [Michigan State University] at 11:29 24 December 2014 Bursa copulatrix strongly bulbous anteriorly, and with nar- moderate transparent central 'window'. Hind wing of male rower posterior region shorter than bulbous region; signum with anal margin expanded but not strongly, with slight hair- a short oval, large, and with large base almost central in scale tuft. signum. MALE GENITALIA (Figs 130, 187). Uncus no more than Diagnosis slightly hood-shaped, rounded subapically, and with small di- The hind wing of pronubata has a more extensive orange re- gitate apex more or less straight (i.e. not much bent over). gion than in other Ischnopteris species, except catocalata and Gnathos with two median processes, tapered from broad base stenoptila (two species known only from females), and occa- and strongly curved throughout their length, not just at the sionally also aurudaria. Moths of pronubata are variable, but apex. Valva with costa strongly curved, forming an acute they always have a definite dark blotch or band at the apex of angle, and with digitate apical costal process. Processes of the hind wing, either joined with or separate from one extend- anellus bent at an angle a short distance before apex, and with ing up the outer margin from the dorsum, whereas the other a subapical lobe taking the form of a triangular plate. Juxta species referred to here do not have that combination of mark- without median process. Aedeagus with irregularly shaped ings. In the male genitalia of pronubata the prominent basal posterior end, with a single, long, narrow, anteriorly-directed lobe on the processes of anellus is characteristic, sometimes spine; vesica with long, narrow, finely spinulose sclerite, not almost approaching the bifurcate condition of hirsuta, which rounded at tip, not on distinct caecum. 76 Linda M. Pitkin

FEMALE GENITALIA (Fig. 231). Lamella postvaginalis Diagnosis without distinct pair of rounded depressions. Lamella anteva- Ischnopteris stenoptila is known from a single specimen, re- ginalis without pair of lobes or median indentation. Antrum sembling the moths of pronubata and catocalata in having a with lateral projection. Signum a moderate-sized disc, with very extensive orange region on the hind wing, but without base much smaller than signum proper. a distinct dark blotch or band at the apex of the wing, as is present in pronubata, or a dark brown line, as is present in Diagnosis catocalata, alongside the dark brown blotch at the tornus. I. The dark fore wing (except for the pale central area frequently stenoptila has a discal spot on the hind wing, not present in present in the female) and contrasting pale hind wing distin- catocalata, and stenoptila has a longer fore wing than that of guishes most specimens of seriei from other species except the other two species. The female genitalia differ from those xylinata, but there is no dark line on the head of seriei as there of pronubata in several characters, notably the more elongate is in xylinata. The male genitalia are unusual for the genus in bursa copulatrix and signum. Differences from catocalata are having no median process of the juxta; only multistrigata is more subtle: the lamella antevaginalis has weaker lobes than similar, but seriei differs in the much greater curvature of the are usually present in that species, and the shape of the antrum gnathos processes and the costa of valva, and in the uniquely and its extension (as described above) differ slightly. More shaped processes of the anellus (described above). In the specimens are needed to be certain that stenoptila is a distinct female, the rounded signum is similar to that of multistrigata species. but not so large in relation to the bursa copulatrix (compare figs 228 and 231). Distribution Colombia. Biological notes Moths have been found at altitudes of 650-1600 m, in March, Ischnopteris xylinata (Guenee) April and December. Figs 62-65,131,132,188, 233, 234 Syrtodes xylinata Guenee, [1858]: 452. LECTOTYPE ´ Distribution BRAZIL (BMNH), here designated [examined]. Argentina. Ischnopterix xylinata ockendeni Prout, 1928: 56. Holotype , PERU: (BMNH) [examined]. Syn. nov. Ischnopteris stenoptila Warren Ischnopteris xylinata xylinata (Guenee); Parsons et al., in Figs 61, 232 Scoble, 1999: 520. Ischnopteris stenoptila Warren, 1907: 289. Holotype , Material examined [COLOMBIA] (BMNH), [examined]. Typical form: 86 , 21 (including 4 , 5 genitalia preparations) (BMNH, VOB, USNM, ZSM). Material examined Lectotype (xylinata), Brazil: no further data (genitalia slide 1 specimen (including genitalia preparation) (BMNH). Geom: 21014, BMNH). Paralectotype (xylinata), Brazil: 1 Holotype , [Colombia: Magdalena,] Santa Marta, Onaca, d, no further data (BMNH) [wrongly associated abdomen e[nd of] vi - b[eginning of] viii (Engelke) (genitalia slide glued on]. Geom: 21034, BMNH). Holotype (xylinata ockendeni), Peru: SE. Peru, Cara- baya, Oconeque, 2140 m ('7000 ft'), vii.1904 (Ockenden) (genitalia slide Geom: 20993, BMNH). Paratypes (xy- Description

Downloaded by [Michigan State University] at 11:29 24 December 2014 linata ockendeni): Peru: 1 , data as holotype but ii.1905 ADULTS (Fig. 61). Fore wing length: , 24 mm. Wing pattern (BMNH); 1 ,Carabaya,Tinguri, 1039 m ('3400 ft'), LI 905 (only female known): fore wing mottled, mainly mid brown; (Ockenden) (BMNH). hind wing mainly orange, with dark brown blotch from tornus to near middle of outer margin of wing, without line adjacent Localities of additional material. Peru: Carabaya, Ocon- to blotch, and with no more than a trace of dark brown near eque. Brazil: Friburgo; Minas Gerais; M[inas] G[erais]: apex of wing; with discal spot. Under-side of fore wing with Carac¸a; Parana: Castro; Rio [de] Janeiro; [Rio de Janeiro:] large orange region bordered by slanting subapical dark brown Petropolis; Rio Derg; Sao Paulo; [S˜ao Paulo:] Salto Grande, transverse band; brown postmedial or medial line broken. Paranapanema; [Sao Paulo: Alto da Serra,] Santos. ˜Bolivia: [La Paz,] Yungas, Coroico. Paraguay: no further data. MALE GENITALIA. Unknown. Argentina: Misiones ('Missions'): Haut-Parana, San Ignacio; FEMALE GENITALIA (Figs 232A, B). Lamella postvaginalis Rio Parana. without distinct pair of rounded depressions. Lamella anteva- Small form A: 9 , 3 (including 5 , 2 genitalia prepara- ginalis with pair of very weak lobes separated by very slight tions) (USNM, VOB, ZSM). median indentation. Antrum with slight lateral indentation (not marked as in pronubata); antrum not longer than broad, but Localities. Brazil: Santa Catarina; Minas Gerais: Passa with posterior triangular extension with sharply pointed apex. Quatro; Minas Gerais: Virginia; Parana: Curitiba; Rio de Bursa copulatrix fairly elongate; signum elongate, oval. Janeiro: P[ar]q[ue] Na[ciona]l Itatiaia. Brazil?: 'Prov. Rio'. Neotropical geometrid moths 77

Small form B: 3 , 1 (genitalia preparations of all) (USNM, some specimens. The male antennae of xylinata are character- VOB). Locality: Brazil: Ba[hia]: Jequie. istic: only two species of Ischnopteris have bipectinate male antennae, xylinata and bipectinata, and the rami are always Description much shorter in xylinata. I. xylinata is unusual in that the male ADULTS (Figs 62-65). Fore wing length: (typical form) , retinaculum is not modified at all, although a few other species 19-21 mm, , 20-24 mm; (small form A) , 16-19 mm, , 20- of Ischnopteris have only a slight enlargement of it. In the male 23 mm; (small form B) , 17-18 mm, , 21 mm. Male an- genitalia, the slender, evenly curved processes of the anellus tenna bipectinate but with very short rami, usually appearing are similar to those of obtortionis, but other characters are very little more than serrate (typical form), or merely serrate and different. For example, xylinata has a single gnathos process, not bipectinate (small form A), or more strongly bipectinate and does not have a pair of lobes on the uncus as in obtortionis. (small form B), but not nearly as strongly bipectinate as in The female genitalia do not have obvious consistent defining bipectinata. Head of both sexes with dark brown dorsal line. characters, although the signum is situated further away from Wing pattern: fore wing of both sexes mottled brown with the anterior end of the corpus bursae than in most other species straw colour and some white markings, and with some dark outside of thefabiana-group. brown lines and streaks; male with dark brown zigzag postmedial line; female often with diffuse dark stripe below costal Variation band, and usually with roughly diamond-shaped central or Some specimens from Brazil show various differences from more extensive area on the wing, paler, darker, or more or the norm, and are referred to here as the small forms A and B. less concolorous and marked out only by dark lines around The male moths are often smaller than those of the typical it. Hind wing of both sexes without orange region, paler than form, and the hind wing is less pale than usual, most markedly fore wing (particularly in male, in which hind wing is typic- in small form B, which may even have the hind wing darker ally predominantly cream or whitish), with brown marginal or than the fore wing. In small form A, the dark line on the submarginal band and (usually in female but only a trace in head is sometimes indistinct. With these features, moths of the male) postmarginal line. Dorsum of male fore wing without small forms may not always be immediately recognised as xy- median tuft. Male retinaculum not enlarged and without linata, but the antennal structure of the males, neither simple transparent central 'window'. Hind wing of male with anal filiform as in most species of Ischnopteris nor as strongly margin not strongly expanded, without pronounced hair-scale bipectinate as in bipectinata, identifies them as xylinata. The tuft. degree of pectination ranges in the three forms of the species: from serrate in small form A, to weakly bipectinate (usu- MALE GENITALIA (Figs 131, 132, 188). Uncus usually ally) in the typical form, and to distinctly bipectinate in small slightly hood-shaped; broadly digitate with much smaller, form B. The genitalic characters of the two small forms are short, digitate apex, which varies from almost straight to bent mainly those of typical xylinata, but the gnathos is smaller than over. Gnathos with one median process, hook-like, strongly that of the typical form, the median process of juxta is on aver- and evenly curved or with hooked tip. Valva with digitate ap- age slightly shorter, and the digitate apical costal process of the ical costal process. Processes of anellus slender, evenly curved; valva is shorter, and in form B also broader (at least basally). with weakly developed patch of minute spinules on sclerotized The processes of the anellus are more weakly curved in form B region at base of each process of anellus. Median process of than in the other two forms. Females of small form A have a juxta tapered and more or less straight, approximately 1.5 times shorter, broader-based projection of the antrum than in the to twice length of main part of juxta, with spinules in apical other two forms; the signum of small form A is smaller than half. Aedeagus with one or more small serrate projections on usual, and is situated not quite so far away from the anterior posterior tongue, vesica with spinulose sclerite, not rounded at end of the corpus bursae as in the other forms. tip, on very short caecum. Downloaded by [Michigan State University] at 11:29 24 December 2014

FEMALE GENITALIA (Figs 233A-B, 234). Lamella postva- Biological notes ginalis with pair of small rounded depressions weak or absent. Moths have been found at altitudes of 950 m to 2140 m (typical Lamella antevaginalis with or without pair of lobes (usually form), 920 m to 2200 m (small form A), and 600m to 750 m weak) separated by median indentation. Antrum with pos- (small form B). They have been collected in January to July, terior projection tapered, with ridge lying usually approxim- and September (typical form); January, February, April, and ately across base of projection of antrum (typical form and October (small form A), and November (small form B). small form B), or without ridge (small form A). Signum oval, situated well away from anterior end of corpus bursae. Distribution Diagnosis South America from Peru and Brazil to Argentina. The pale hind wing contrasting with the darker fore wing is usually distinctive for xylinata, especially in the males, but Species excluded from Ischnopteris: seriei may appear very similar. However, that species does not 'Ischnopteris' discolor species group have a dark line on the head as is present in xylinata. Females The following three species appear to be closely related to of xylinata with a dark stripe below the costal band of the fore each other, and are here treated provisionally as the discolor- wing are easily recognised, but this character is present only in group, but their relationships within the Nacophorini and their 78 Linda M. Pitkin

correct placement to genus remain unresolved. They do not 'Ischnopteris' discolor Warren share the apomorphies by which I have defined Ischnopteris Figs 66, 67,133,189, 235 and Rucana, although they share some other similarities with Ischnopteris discolor Warren, 1904b: 557. Holotype , PERU the latter genus. The moths have the general appearance of Ischnopteris andRucana, with simple filiform antennae in both (BMNH) [examined]. sexes, elongate fore wing and slender elongate male abdomen, Ischnopteris conjugens Warren, 1905a: 59. Holotype wings mainly mottled brown, mid or dark, with or without an , PERU (BMNH) [examined]. [Synonymy cited by orange region on the hind wing. However, the male retinaculum Parsons et al., in Scoble (1999).] is not enlarged and it does not have the transparent central 'window' that is present in nearly all species of Ischnopteris; Material examined the front of the head is swollen, but not shiny and scale-free as 4 , 3 (including 2 , 1 genitalia preparations) (BMNH, is common in Rucana. The moths have a feature on the fore USNM). wing, similar to one in Rucana, consisting of two small whitish dashes, one near the tornus and another near the dorsum on the Holotype (discolor), Peru: SE. Peru, Carabaya, Santo outer side of the postmedial line; a short dark bar between the Domingo, 1990 m ('6500 ft'), x.1902 (Ockenden) (genitalia whitish markings is sometimes distinct but sometimes merges slide Geom: 21293, BMNH). with the dark ground colour of the wing. This feature occurs Holotype (conjugens), Peru: SE. Peru, Carabaya, Santo also in certain other nacophorine moths (e.g. Quillaca). Domingo, 1830 m ('6000 ft'), xi.1902 (Ockenden) (genitalia slide Geom: 21001, BMNH). In the male genitalia, the uncus is narrow, almost rod- like but slightly curved and tapered to a pointed apex, without Localities of additional material. Peru: Carabaya, Santo the distinct dorsal swelling that characterizes Rucana. The un- Domingo. Bolivia: no further data. cus does not have a hood-shaped structure, nor does it have dense ventral setae, such as are present in Ischnopteris, but Description dorsal hair-scales are occasionally fairly dense, although not ADULTS (Figs 66, 67). Fore wing length: , 19 mm; , 19- forming such a pronounced tuft as in Rucana or Quillaca. The 20 mm. Wing pattern: fore wing of male dark chocolate brown, gnathos has a single median process that is sclerotized, pointed with very pale olive-brown postmedial and antemedial lines tongue-like, and curved. The valva does not have an apical and a small subapical marking, and with two small whitish costal lobe or process (except in one undescribed species pos- dashes, one near tornus and another near dorsum on outer sibly affiliated to this group), nor does it have a patch of dorsal side of postmedial line; fore wing of female usually paler setae in the region of the sacculus, and the sacculus does not than that of male, and always with ill-defined pale central have a ventral process (a definitive character of Ischnopteris). area transected by dark zig-zag line or narrow band, overall The transtilla forms a distinct pair of bulbous lobes (as are with mottled markings of dark brown and very pale olive- often seen in Rucana and occasionally in Ischnopteris). The brown. Hind wing of both sexes mid brown, slightly greyish, paired processes of the anellus are usually similar to those without orange region. Under-side of wings paler than upper commonly occurring in Ischnopteris and Rucana, extending side (male); both sexes with apical or subapical cream band, postero-laterally from juxta, pointed and claw-like, and the sometimes diffuse or broken, and with brown postmarginal base of the processes is extended, but one undescribed species line, the latter present at least on hind wing. of the discolor-group differs in having bifurcate processes. The juxta has a V-shaped anterior end, like that of Rucana MALE GENITALIA (Figs 133, 189). Characters as described but smaller; the medio-posterior process of the juxta is long under discolor-group. (in the three described species), usually with some spinules,

Downloaded by [Michigan State University] at 11:29 24 December 2014 and similar to that of Ischnopteris and Rucana but always FEMALE GENITALIA (Fig. 235). Characters as described tongue-shaped and straight, not bent to one side as in some under discolor-group. Ischnopteris species. The vesica of the aedeagus always has cornuti in the form of a group of slender spines. Diagnosis Adults of this species differ from those off esta and festiva in The female genitalia are known for only two of the named having no orange region on the hind wing. An undescribed species. The bursa copulatrix is mainly membranous, very species is similar to discolor but with a paler, more mottled, long and slender, with a bulbous anterior end, and with a fore wing pattern, and that species is unique in this group of postero-lateral caecum. The antrum forms a small collar, not a species in having bifurcate processes of the anellus. complete ring, and has no posterior projection. These features are similar to those of Rucana, but the species of the discolor- group have a signum, which is absent in Rucana. The signum Biological notes in the discolor-group is rugose to spinose and extended to one Moths have been found at altitudes of 1830 m and 1990 m, in side of the disc-like base. October and November. The discolor-group comprises three named species, and in addition there are three (or possibly more) undescribed spe- Distribution cies in this group. Peru and Bolivia. Neotropical geometrid moths 79

'Ischnopteris' festa Dognin Localities of additional material. Colombia: E. Figs 68,134,190 Colombia, [Cundinamarca,] Medina. Ecuador: Zamora- Chinchipe, Parque Nacional Podocarpus, Rio Bombuscara, Ischnopteris festa Dognin, 1912: 148. Holotype , COLOM- N.P. station; Zamora-Chinchipe, Estacion Cientifica San Fran- BIA (USNM) [examined]. cisco. Peru: Pumayacu. Material examined 5 (including 2 genitalia preparations) (BMNH, SMNS, Description USNM). ADULTS (Fig. 69). Fore wing length: , 15-17 mm; , 17- 19 mm. Wing pattern (both sexes similar but female tends Holotype , Colombia: [Tolima,] Quindio, Rio Toche, 2400 to be paler than male): fore wing mottled dark brown with m (Fassl) (Type No. 32544, genitalia slide 60148, USNM). varying amount of paler flecks, and with pale postmedial and antemedial lines (sometimes indistinct); with two small whit- Localities of additional material. Colombia: upper ish dashes, one near tornus and another near dorsum, on outer ('Ob.'[erer]) Rio Negro. Ecuador: Zamora-Chinchipe, old side of postmedial line; postmedial line forming acute angle. road Loja-Zamora; Zamora-Chinchipe, Rio San Francisco, Es- Hind wing dark or mid brown to grey, with orange apical tacion Cientifica San Francisco. ´Peru: [Junin], El Porvenir. region. Under-side of wings similar to upper but paler and duller; fore wing with pale apical band, not strongly defined Description and sometimes indistinct or broken; hind wing with brown ADULTS (Fig. 68). Fore wing length: , 19-20 mm. Wing postmarginal line. pattern (only male known): fore wing mottled dark brown, with pale olive postmedial and antemedial lines and a few MALE GENITALIA (Figs 135, 191). Characters as described whitish flecks, particularly two small whitish dashes, one near under discolor-group. tornus and another near dorsum, on outer side of postmedial line; postmedial line forming obtuse angle. Hind wing dark FEMALE GENITALIA (Fig. 236). Characters as described brown, sometimes greyish, with orange apical region. Under- under discolor-group. side of wings paler than upper side, with paler, dull yellow and brown, apical regions sometimes forming an apical band; hind Diagnosis wing with brown postmarginal line. Moths of festiva andfesta are very similar; both have an orange region on the hind wing, but festiva moths are smaller and MALE GENITALIA (Figs 134, 190). Characters as described the postmedial line on their fore wings forms an acute angle under discolor-group. (obtuse infesta).

FEMALE GENITALIA. Unknown. Biological notes Moths have been found at altitudes of 500-1845 m, in June, Diagnosis October, and December. Moths of festa and festiva are very similar; both have an orange region on the hind wing, but festa moths are larger and Distribution the postmedial line on their fore wings forms an obtuse angle Colombia, Ecuador, and Peru. (acute in festiva). Other species excluded from Ischnopteris: Biological notes Moths have been found at altitudes of 760 m to 2400 m, in 'Ischnopteris' callistrepta (Prout) March, October, and December. not illustrated Downloaded by [Michigan State University] at 11:29 24 December 2014 Ischnopterix callistrepta Prout, 1928: 56. Holotype , PERU Distribution (BMNH) [examined]. Colombia, Ecuador, and Peru. Material examined 'Ischnopteris' festiva Warren 6 , 4 (including 1 , 1 genitalia preparations) Figs 69,135,191, 236 Holotype , Peru: SE. Peru, Carabaya, Limbani, 2900 m Ischnopteris festiva Warren, 1904b: 557. Holotype , PERU ('9500 ft'), iv.1904 (Ockenden) (BMNH). (BMNH) [examined]. Paratype 1 , data as holotype but v.1904 (BMNH).

Material examined Localities of additional material. Ecuador: Zamora- 5 , 2 (including 2 , 1 genitalia preparations) (BMNH, Chinchipe, Estacion Cientifica San Francisco (SMNS). ´Peru: SMNS, USNM). Carabaya, Agualani (BMNH).

Holotype , Peru: SE. Peru, River Slucuri, 760 m Remarks ('2500 ft'), vi.1901 (Ockenden) (genitalia slide Geom: This species does not share the apomorphies of Ischnopteris, 21002, BMNH). and it does not have any obvious affinities with that genus or 80 Linda M. Pitkin

the other two in this study. Its relationships within the Naco- central 'window' in speculifera. Dorsum of male fore wing phorini are unresolved, although it may be related to 'Cidario- without median tuft or lobe. Hind wing of male with anal phanes' albimargo (Dognin) and also possibly to other species margin usually not expanded (but expanded and with hair- misplaced in Cidariophanes. scale tuft in speculifera). Metathorax with no more than weak dorsal tuft. Male abdomen fairly elongate and slender, with one Distribution or more dorsal rows or patches of 'metallic' scales, situated Ecuador and Peru. where segments overlap, and with small black or dark brown patches of scales, sometimes slightly raised. Sternite 3 of male One other species, Ischnopteryx viriosa Dognin, is here trans- abdomen usually with triangular or inverted V-shaped patch ferred to Quillaca: of setae (absent in some specimens of cythereata).

Quillaca viridifascia (Warren) MALE GENITALIA (Figs 136-143, 192-198). Uncus nar- not illustrated row, rod-like or sometimes spatulate or tapered, usually with pointed apex, not distinctly hood shaped; one species (spe- Ischnopteris viridifascia Warren, 1904b (September): 558. culifera) with dorsal pseudouncus. Socii usually very small LECTOTYPE [stated as ], PERU (BMNH), here desig- or absent, but occasionally moderately well developed in spe- nated [examined]. culifera. Gnathos often weakly developed and with arms not Ischnopteryx viriosa Dognin, 1904 (November): 365. Holo- joined, or forming pair of projections joined by weaker median type , PERU (USNM) [examined]. Syn. nov. strip, or gnathos with single hook- or tongue-like median exten- Ischnopterisprojectata Warren, 1905a: 60. LECTOTYPE , sion. Valva with costal band (extremely broad in albipennis), PERU (BMNH), here designated [examined]. [Synonymy with or without ventral row of long spines or spine-like setae cited by Parsons et al., in Scoble (1999): 163.] at inner edge of costal band; valva without apical costal lobe or process; sacculus with neither patch of dorsal setae nor small pointed ventral process (as are characteristic of Ischnopteris). Stegotheca Warren Transtilla usually weak or absent, or present merely as lateral lobes, but lobes longer and almost meeting in speculifera. Pro- Stegotheca Warren, 1900: 204. Type species: Stegotheca cesses of anellus small or absent, except in speculifera, which amis sa Warren, 1900, by original designation. [Placed has a large pair extending to transtilla. Juxta a very large plate incorrectly in Selidoseminae by Warren, 1900: 204; (except in speculifera), usually with spinules and granules in assigned to Nacophorini by Pitkin, 2002.] posterior region; cristae often weak but sometimes moderately Salasaca Rindge, 1983: 189. Type species: Salasaca spinea prominent. Aedeagus: vesica usually with cornuti, commonly Rindge, 1983, by original designation. Group 2 of Rindge a large region of regularly arranged squat spinules or occasion- (1983). Syn. nov. ally minute denticles, but in speculifera with a group of longer Canelo Rindge, 1983: 233. Type species: Canelo spines similar to those of Rucana. constrictus Rindge, 1983, by original designation. Group 3 of Rindge (1983). Syn. nov. FEMALE GENITALIA (Figs 237-241). Bursa copulatrix mainly membranous, very long and often very slender, but Description sometimes with bulbous anterior end. Signum usually present, ADULTS (Figs 70-77, 91, 93, 94). Fore wing length: , 12- but absent in phalangifera, and female of speculifera unknown; 20 mm; , 14-21 mm. Antenna simple filiform in both signum ranging from smooth to dentate, usually fairly small sexes, sometimes fairly thick in male. Wing pattern: fore and hollow, with disc-shaped or oval base either in the centre wing brown with mottled markings, sometimes with whitish of the signum or adjacent to rim of signum. Downloaded by [Michigan State University] at 11:29 24 December 2014 flecks but these only conspicuous in females of albipennis; dark brown dash present, sometimes indistinct, near apex of Diagnosis fore wing; antemedial and postmedial lines narrow and dark Moths of Stegotheca generally resemble those of Ischnopteris, brown, postmedial strongly angled. Hind wing varying, either but are smaller on average, and never with an orange apical fairly plain brown or greyish, or white or yellowish-orange blotch on the hind wing as in some Ischnopteris species. Spe- with brown band at outer margin. Under-side of wings usu- cies of Stegotheca (with the exception of S. albipennis) are ally with marginal band (or on fore wing at least an apical less sexually dimorphic in wing pattern than is the case in marking) but markings sometimes diffuse; specimens with in Ischnopteris. Stegotheca is defined by two autapomorphies: white or yellowish-orange on upper side of hind wing have the patch(es) of 'metallic' scales on the abdomen, and the pres- those colours more diffusely on under-side of fore wing, ence of a fovea or a fovea-like structure on the fore wing of mottled with brown. Fore wing fairly elongate; with blister-like the male. Elsewhere in the tribe Nacophorini a fovea is known fovea in male, with or without crests of large scales at edge; only in Mimophyle and Plesiophyle, neither of which is con- fovea usually situated near base of wing between cubital and firmed as belonging in this tribe. Those two genera are very anal veins, but speculifera instead with fovea-like structure different from Stegotheca in appearance and in genitalic char- between subcosta and radius. Male retinaculum usually not acters (see Pitkin, 2002: Figs. 140, 141, 382, 383, 548, 662). enlarged, without transparent central 'window', but enlarged The enlargement of the male retinaculum in two species of in two species (amissa and speculifera) and with transparent Stegotheca, with a window in one and without a window in the Neotropical geometrid moths 81

other, is a synapomorphy apparently shared with Ischnopteris, — Moths larger: fore wing length 15 mm or more 7 where it is present in most species. 7. Fore wing of male with dark patch situated medially to The male genitalia of Stegotheca are characterized by the subbasally (Figs 72, 73), female with large pale central weak gnathos (the arms are not joined by sclerotization in five area costiplaga of the seven species), the large plate-like juxta with spinules - Fore wing of both sexes without any large dark or pale areas or granules posteriorly, and the weak transtilla (except in the (Fig. 76) phalangifera sp. nov. anomalous speculifera). Stegotheca species do not share the apomorphies of Ischnopteris: dense ventral setae on the uncus Species and a tiny ventral process on the sacculus of the valva. The Stegotheca albipennis (Warren) comb. nov. valva does not have an apical costal lobe or process as is Figs 70,136,192, 237 present in Ischnopteris. Ischnopteris albipennis Warren, 1904b: 556. Holotype Stegotheca was previously known from a sole specimen [stated as ], PERU (BMNH) [examined]. (a male of amissa). Examination of the scales on the abdo- Canelo albipennis (Warren); Pitkin, 2002: 206. men, and the fovea on the fore wing, now confirms Canelo Canelo constrictus Rindge, 1983:234, figs 73,92. Holotype , and Salasaca as synonyms of Stegotheca, a possibility that I BOLIVIA (AMNH) [examined]. [Synonymized by Pitkin, discussed in Pitkin (2002: 205). 2002: 206.] Biological notes Moths of Stegotheca are known from elevations ranging from Material examined 140 m to 2800 m. Nothing is known about their early stages 7 (+1 photograph) , 12 (including 3 , 2 genitalia pre- or hostplants. parations) (AMNH, BMNH, CMNH, USNM).

Distribution Holotype (albipennis), Peru: SE. Peru, Carabaya, Santo Stegotheca occurs in the Neotropical Region, where it is known Domingo, 1830 m ('6000 ft'), iv.1902 (Ockenden) (genitalia from Guatemala southwards to Bolivia. slide Geom: 20228, BMNH). Checklist of species Holotype (constrictus), Bolivia: Yungas de Palmar, 2000 m (AMNH). albipennis (Warren, 1904b) (Ischnopteris) comb. nov. [from Canelo] Localities of additional material. Colombia: E. constrictus (Rindge, 1983) (Canelo) Colombia, [Cundinamarca,] Medina; Muzo; upper amissa Warren, 1900 (Stegotheca) ('Ob.'[erer]) Rio Negro. Ecuador: no further data [digital parvula (Schaus, 1912) (Ischnopteryx) syn. nov. photograph by Gunnar Brehm]. Peru: Carabaya: Oconeque; costiplaga (Dognin, 1911) (Ischnopteris) comb. nov. Santo Domingo; Cuzco, Cosnipata, Paucartambo; Tinguri; [from Ischnopteris] SE. Peru, Rio Inambari, La Oroya. Bolivia: Cochabamba, spinea (Rindge, 1983) (Salasaca) syn. nov. Yungas del Espiritu Santo. cythereata (Guenee, [1858]) (´Syrtodes) comb. nov. [from Canelo] Description phalangifera sp. nov. ADULTS (Fig. 70). Fore wing length: , 17-20 mm; , 19- speculifera (Bastelberger, 1908c) (Ischnopteris) comb. nov. 21 mm. Wing pattern: fore wing mottled brown, with slanting [from Ischnopteris] median patch bounded by postmedial line and ranging from indistinct to distinctly darker; apical region of female fore wing Key to species dappled with white; hind wing white, with greyish brown band 1. Hind wing with brown marginal band contrasting with pale

Downloaded by [Michigan State University] at 11:29 24 December 2014 at outer margin. Under-side of fore wing faintly to strongly inner region (Figs 70, 74) 2 white-tinged, with brown margin. Raised scales at edge of — Hind wing without contrasting pattern 4 fovea (on male fore wing) little differentiated from surround- 2. Hind wing with white inner region (Fig. 70)... albipennis ing scales. Male retinaculum not enlarged, without transparent — Hind wing with yellow or dull orange inner region central 'window'. Abdomen with one slightly swollen patch of (Fig. 74) cythereata (typical form) 'metallic' scales (at anterior end of segment 4); tip of abdomen 4. Male with retinaculum enlarged; female genitalia as in yellow. Fig. 238 or female as yet unknown 5 — Male with retinaculum not enlarged; female genitalia as in MALE GENITALIA (Figs 136, 192). Uncus spatulate with Figs 239, 241, or unknown 6 pointed apex. Gnathos with arms joined by short weaker me- 5. Fore wing of male with elongate fovea-like structure dian strip, each arm ending with spine-like projection. Valva between veins subcosta and radius; retinaculum with trans- with extremely broad costal band, flanked with row of long parent central 'window' [female unknown] ... speculifera spines or spine-like setae (including some multi-tipped). Pro- — Fore wing of male with fovea between cubital and anal cesses of anellus absent. Juxta posteriorly forming two lobes veins; retinaculum without transparent central 'window'; separated by median indentation; triangular spinules and often female genitalia as in Fig. 238 amissa faint granules present on lobes. Aedeagus with tapered pos- 6. Moths small: fore wing length of male 11 mm [female terior end; vesica often with faint scaly appearance of surface unknown] cythereata (brown form) but without distinct cornuti; vesica without caecum. 82 Linda M. Pitkin

FEMALE GENITALIA (Fig. 237). Lamella postvaginalis greatly enlarged but without transparent central ' window '. Ab- strongly wrinkled medially; sternite 8 with posterolateral pair domen with two patches of 'metallic' scales (a band at anterior of low lobes close together, with indentation between them. end of segment 4, and a small patch at posterior end of seg- Lamella antevaginalis broad, with slight median indentation; ment 3), situated between two spots of black raised scales, one breadth of posterior end of antrum similar to distance between on segment 3 and the other on segment 4. apophyses anteriores; antrum vase-like, no more than weakly MALE GENITALIA (Figs 137, 193). Uncus rod-like, slightly sclerotized. Bursa copulatrix very long and slender, mainly curved, pointed. Gnathos with weakly developed arms not membranous. Signum a smooth rounded pocket. joined medially by sclerotized section. Valva with medio- ventral ridge well separated from costa; costa straight; valva Diagnosis without row of spines or spine-like setae. Processes of anellus Several characters are distinctive and putatively autapo- present as pair of small inconspicuous lobes situated postero- morphic for albipennis. A particularly obvious feature is the laterally to juxta. Juxta with minute spinules and granules in hind wing colouration, white with a brown marginal band, but posterior portion, and with broad, rounded, posterior margin. the genitalia also have characters that are unique in the genus, Aedeagus: vesica with large, broad region of evenly spaced, including the presence of multi-tipped spines or setae on the short and stout spines; vesica with short caecum. male valva (costiplaga also has spines but not multi-tipped), and the smooth, rounded, pocket-shaped signum in the female. FEMALE GENITALIA (Fig. 238). Lamella postvaginalis lightly sclerotized and weakly wrinkled, varying in size from large to small; sternite 8 with posterolateral pair of low lobes Biological notes well apart but only weakly developed and without distinct Moths have been found at altitudes of 500 m to 2140 m, in margin. Posterior end of antrum fairly broad but breadth less January to April, September, and November. than distance between apophyses anteriores; antrum cup- or vase-shaped and only lightly sclerotized. Bursa copulatrix Distribution mainly membranous, with wrinkles and minute spinules in Western South America from Colombia to Bolivia. very slender posterior half; anterior half varying: long and narrow or bulbous. Signum a small hollow ovoid with base more Stegotheca amissa Warren or less central; with several wrinkles or ridges and weakly Figs 71, 94,137,193, 238 serrate edge. Stegotheca amissa Warren, 1900: 205. Holotype , Diagnosis ECUADOR (BMNH) [examined]. Ischnopteryx parvula Schaus, 1912: 427. LECTOTYPE , Stegotheca amissa is the only species in the genus with a male retinaculum that is enlarged but does not have a transparent COSTA RICA (USNM), here designated [examined]. Syn. central 'window'. In the male genitalia the juxta shape, with nov. its broad rounded margin, is distinctive and putatively autapomorphic for amissa. In the female, the form of the signum, as Material examined described above, differs from that of other Stegotheca species. 3 , 2 (including 2 , 2 genitalia preparations) (BMNH, SMNS, USNM). Biological notes One moth was found at an altitude of 525 m; moths have been Holotype (amissa), Ecuador: [Rio?] Cachab´ı, low country, found from May to July and in November. xi. 1896 (Rosenberg) (genitalia slide Geom: 19311, BMNH). Lectotype (parvula), Costa Rica: Carillo (Type No.

Downloaded by [Michigan State University] at 11:29 24 December 2014 Distribution 17605, genitalia slide 60140, USNM). Paralectotype, 1 , Costa Rica, Panama, and Ecuador. Costa Rica: Carillo, vii (USNM). Stegotheca costiplaga (Dognin) comb. nov. Localities of additional material. Costa Rica: Heredia Figs 72, 73,138,194, 239 Province, Braulio Carrillo N[ational] P[ark], W. Rio Peje, Puesto Ceibo. Panama: Chiriqui. Ecuador: Cachab´ı. Ischnopteris costiplaga Dognin, 1911: 179. Holotype , COLOMBIA (USNM) [examined]. Description Salasaca spinea Rindge, 1983: 190, figs 31, 41, 51. Holotype ADULTS (Figs 71,94). Fore wing length: , 13-15 mm; ,17- , ECUADOR (AMNH) [examined]. syn. nov. 18 mm. Wing pattern (both sexes): fore wing mottled brown or Material examined charcoal, usually without but occasionally (one female) with 5 , 1 (including 4 , 1 genitalia preparations) (AMNH, large dark median patch (contrasting strongly with ground CMNH, SMNS, USNM). colour) slanting from costa towards dorsum and bounded by postmedial line; hind wing brown, grey or charcoal, without Holotype (costiplaga), Colombia: Cali, San Antonio (Fassl) distinct border but with narrow darker postmedial line. Raised (Type No. 31185, genitalia slide 60142; USNM). scales at edge of fovea hair-like, sparse and deciduous; Holotype (spinea) Ecuador: Loja Province, Loja to fovea situated directly above male retinaculum, which is Zamora, 2800 m, 26-27.x.1977 (Pena˜) (genitalia slide Neotropical geometrid moths 83

19043A; AMNH). Allotype , Ecuador: Loja [Province], nor some spines multi-tipped as in albipennis. The female Macara to Catacocha, 650 m, 13—14.viii.1977 (´Pena˜) genitalia differ from those of other species of Stegotheca in (AMNH). having the lamella antevaginalis with a pair of lobes, similar to that of Ischnopteris species. However, there is no definite proof Localities of additional material. Colombia: no further that the only two known females truly belong to costiplaga. data; Cali, San Antonio. Ecuador: Esmeraldas, Cotacachi- Cayapas Reserve, Rio de Cristal; Zamora-Chinchipe Province, Biological notes Estacion Cientifica San Francisco. Moths have been found at altitudes of 650 m to 2800 m, in May, August, and October. Description ADULTS (Figs 72, 73). Fore wing length: , 15-18 mm; , Distribution 18 mm. Wing pattern: fore wing mottled brown and straw, Colombia and Ecuador. sometimes with patches of olive green; male with large dark median to subbasal patch (contrasting strongly or slightly Stegotheca cythereata (Guenee) comb. nov. with ground colour) slanting from costa towards dorsum and bounded by postmedial line, female with large pale central area Figs 74, 75, 91, 139,140,195,196, 240 (cream, or according to Rindge (1983) greyish white). Hind Syrtodes cythereata Guenee, [1858]: 452. Holotype ´ , wing pale to dark, white-tinged brown or grey, with indistinct [BRAZIL] (BMNH) [examined]. darker brown border, with or without narrow dark postme- Canelo cythereata (Guenee); Pitkin, 2002: 206. dial line. Under-side of wings with dark patches and whitish patches at outer margin, varying from very distinct and strong Material examined to considerably fainter. Raised scales at edge of fovea little dif- Typical form: 43 , 18 (including 5 , 3 genitalia prepar- ferentiated, not much larger than surrounding scales. Male ret- ations) (AMNH, BMNH, CMNH, SMNS, USNM, VOB, inaculum not enlarged, without transparent central 'window'. ZSM). Abdomen with one slightly swollen patch of brown 'metallic' Holotype , Brazil: Para (genitalia slide Geom: 20232, scales, posteriorly adjoining dark brown spot. BMNH).

MALE GENITALIA (Figs 138, 194). Uncus tapered, with Localities of additional material. Guatemala: pointed apex. Gnathos with single tongue-like median exten- Cayuga. Costa Rica: Cartago, 2 km E. [of] Moravia de sion. Valva with costal band (only moderately broad); ventral Chirripo.´ Trinidad: Caparo. Colombia: western Colombia, ridge present along inner edge of costal band, with short row Valle, Calima Dam [digital photograph by Bo Sullivan]; E. of long spines or spine-like setae (none multi-tipped) situated Colombia, [Cundinamarca,] Medina. Venezuela: San Esteban. apically, but well before apex of valva. Processes of anellus Surinam: Marowijne Valley, Areowarwa Kreek. Brazil: Para; absent. Juxta posteriorly forming two lobes pressed together, R[io de]J[aneiro], Mangaratiba. French Guiana: no fur- with marginal band of triangular spinules and sometimes with ther data; Cayenne; Godebert-Maroni; St Jean du Maroni. faint granules. Aedeagus with tapered, tongue-like, posterior Ecuador: Loja 'environs'; Napo, Misahualli; Napo-Pastaza, extension; vesica with very faint scaly appearance of surface Dureno, S. Rio Aguarico; Zamora-Chinchipe, Parque but without distinct cornuti; vesica without caecum. Nacional Podocarpus, [Rio] Bombuscara. Peru: Carabaya, R. Huacamayo, La Union; Chanchamayo; Hu´anuco, Pozuzo, FEMALE GENITALIA (Fig. 239). Lamella postvaginalis not [Puno,] Chaquimayo. Bolivia: E. Bolivia, Buenavista; sclerotized except for some median wrinkles; sternite 8 without [Cochabamba,] Chapare, Chapare area, upper Rio Chipirire; posterolateral pair of lobes (such as are present in albipennis Santa Cruz, Prov[incia del] Sara. Downloaded by [Michigan State University] at 11:29 24 December 2014 and phalangifera). Antrum forming a collar, narrow at posterior end where it adjoins the broad lamella antevaginalis, Brown form: 2 ; genitalia preparations of both (BMNH). which has at its posterior margin a pair of lobes with a median Brazil: Rio de Janeiro, [Serra dos Orgaos] Organ Mts, near indentation. Bursa copulatrix long, with light sclerotization Tijuca. and striations in more slender posterior half, and with membranous, slightly bulbous, anterior half. Signum a small ovoid, Description hollow and extended to one side of base, with a few denticles ADULTS (Figs 74, 75, 91). Fore wing length: (typical form) and dentate edge. cf, 12-14 mm; , 14-16 mm; (brown form) , 11 mm. Wing pattern (both sexes): fore wing brown mottled with straw or Diagnosis yellow-ochre, a few specimens with apical half of wing darker Stegotheca costiplaga can be recognised, in the male, by the brown; apex of fore wing with blackish marking. Hind wing: combination of a dark patch situated medially to subbasally typical form yellow to dull orange, with well-defined brown on the fore wing, and the retinaculum not being enlarged as band at outer margin, and sometimes with faint brown postme- in amissa. In the male genitalia, the form of the juxta, with dial line; brown form fairly pale brown mottled with yellowish lobes pressed together and not separated by an indentation as straw, with darker postmedial line and with indistinct brown in albipennis, is autapomorphic for costiplaga. The spines on band at outer margin. Under-side of fore wing often yellow- the valva are distinctive, not situated well away from the costa tinged, with darker brown at margin. Raised scales at edge 84 Linda M. Pitkin

of fovea (on male fore wing) large and blade-shaped. Male cythereata vary in the shape and size of the antrum, and in the retinaculum not enlarged, without transparent central 'win- form of the sclerotized region of the bursa copulatrix. dow'. Abdomen with two bands of silvery 'metallic' scales Two male specimens from Brazil, referred to here as the in both sexes (at anterior end of segments 3 and 4, posterior brown form, differ in having brown hind wing colour, instead band sometimes indistinct); bands situated between three black of yellow or orange. Their small size and general similarities of spots; narrow black band or scattered black scales at posterior the genitalia associates them most closely with cythereata, al- end of segment 2. though there are minor genitalic differences, described above. They might eventually prove to be a distinct species, but at the MALE GENITALIA (Figs 139, 140, 195, 196). Uncus rod- present time I consider it best to treat them under cythereata, like, apex pointed, tapered, or very slightly swollen. Gnathos since that species encompasses considerable variation, which with weakly developed arms, often not joined. Valva without has not yet been fully investigated. row of spines or spine-like setae; valva usually with short, weak, ventral ridge near costa, but ridge absent in two specimens (from Surinam and Colombia), and situated medio- Biological notes ventrally in the brown form. Processes of anellus present as Moths have been found at altitudes of 150 m to 1150 m, in pair of small lobes, usually bearing a few setae, situated pos- January, February, May to July, and September to December. terolaterally to juxta (but not clearly defined in specimen from Colombia). Juxta with tapered posterior portion usually end- Distribution ing in truncate apex; usually with large region of granules Typical form: Central and South America from Guatemala to in posterior region, and a line of minute spinules at margin, Brazil and Bolivia. Brown form: Brazil. but granules absent in two specimens (from Surinam and Colombia), spinules absent in another specimen, and both Stegotheca phalangifera sp. nov. granules and spinules absent in the brown form. Aedeagus Figs 76,141,197, 241 with posterior end more or less truncate; vesica with large Material examined and broad region of cornuti (usually evenly spaced and often (3 , 2 ; genitalia preparations of all). consisting of a patch of small triangular spinules adjacent to a patch of larger spines, but subject to variation, in brown Holotype , French Guiana: Piste Coralie 17.xi.1989 ('Sene- form consisting of well-spaced spines interspersed with small caux rec.') (genitalia slide LMP 372; ZSM). spinules); vesica with caecum. Paratypes: Peru: 1 , [Loreto,] Contamana, Rio Ucay- ali, x-xii (BMNH); 1 , Madre de Dios, Avispas, 20- FEMALE GENITALIA (Fig. 240). Lamella postvaginalis 30.ix.1962 (Pena˜) (AMNH). Brazil: 1 , Ro[ndonia], lightly wrinkled; sternite 8 without distinct pair of posterolat- Cacaulandia, 140 m, 13-31.xii. 1997 (Becker) (VOB); eral lobes (such as are present in albipennis andphalangifera). 1 , data as before but xi. 1994 (VOB). Breadth of posterior end of antrum much less than distance between apophyses anteriores; antrum bulbous or vase-like Description and sclerotized, sometimes only lightly. Bursa copulatrix with ADULTS (Fig. 76). Fore wing length: ,15-16 mm; ,17 mm. some sclerotization, striations and/or wrinkles, but with mem- Wing pattern (both sexes): fore wing mottled brown or grey- branous, bulbous anterior end. Signum a dentate hollow disc, brown, without any large dark or pale areas; hind wing brown with base central. or grey without distinct border; with darker but fine postmedial line. Under-side of hind wing with darker margin, sometimes Diagnosis also on under-side of fore wing but less distinct. Raised scales

Downloaded by [Michigan State University] at 11:29 24 December 2014 The most obvious characteristic of cythereata is the hind wing at edge of fovea hair-like, deciduous. Male retinaculum not colouration, yellow or yellowish with contrasting brown outer enlarged, without transparent central ' window '. Abdomen with marginal band (except in the brown form - see under 'Vari- blackish band or broken markings with some dark 'metallic' ation'). The male genitalia of cythereata do not have any ob- scales in both sexes (some of these markings at anterior end vious features that are both distinctive and consistent, but the of segment 3, and situated between two or three tiny blackish female's dentate signum with the base central is unique in the spots. genus, as far as is known; costiplaga has a dentate signum but MALE GENITALIA (Figs 141, 197). Uncus a pointed rod, the base is at the edge. very slightly constricted before broadening to base. Gnathos weakly to moderately well developed, with arms joined by Variation median plate that is occasionally only lightly sclerotized. Valva One male from Surinam and another from Colombia show with weak medio-ventral ridge apically approaching costa; some minor differences in the male genitalia, described above, costa slightly curved; valva without row of spines or spine-like and the Colombian specimen differs from the norm (i.e. setae. Processes of anellus ill-defined or absent. Juxta large but Fig. 139) in having the valva broad and with a sinuous costa. lightly sclerotized, with posterior margin tapered to rounded Another Colombian male, examined in a digital photograph or blunt apex and only faintly granular. Aedeagus: vesica with provided by Bo Sullivan, is unusual in having a median indent- long narrow strip of evenly spaced, short and broad spines; ation at the posterior end of the juxta. The female genitalia of vesica without caecum. Neotropical geometrid moths 85

FEMALE GENITALIA (Fig. 241). Lamella postvaginalis a to subbasal patch; hind wing greyish brown, almost plain, narrow sinuous transverse sclerite; sternite 8 with posterolat- without orange region. Under-side of wings greyish brown, eral pair of low lobes well apart. Breadth of posterior end of fore wing with cream apical spot, sometimes indistinct. Fore antrum similar to distance between apophyses anteriores; an- wing with pronounced elongate fovea-like transparent region, trum broad, semicircular, moderately well-sclerotized. Bursa situated between subcosta and radius and lying over retin- copulatrix very long and slender, anterior half to two-thirds aculum, flanked on costal edge by double comb-like row of membranous, but with a sclerotized patch situated more pos- modified scales on underside of wing. Male retinaculum en- teriorly; signum absent. larged and with transparent central 'window'. Male frenulum elongate, slightly swollen subapically. Hind wing of male with Diagnosis anal margin expanded and with darker hair-scale tuft. Abdo- Stegotheca phalangifera is closely related to amissa but men elongate and slender, with one to two dorsal patches of differs in having a normal male retinaculum (not enlarged as in 'metallic' scales, situated at anterior end of segments 3 and 4; amissa), male genitalia with the apical end of the medio-ventral abdomen with up to four very small deciduous tufts of dark ridge of the valva approaching the costa, a tapered juxta (this brown raised scales. has a broad margin in amissa), and a different arrangement of MALE GENITALIA (Figs 142A-B, 143, 198). Uncus some- spines in the vesica. The last character, a distinctive narrow what rod-like but divided into two tapered parts: a ventral strip of regularly-spaced spines, is a putative autapomorphy process, and a dorsal pseudouncus curved and bearing a dorsal for phalangifera. Further putative autapomorphies are seen in tuft of fine setae or hair-like scales. Gnathos with single me- the female genitalia: the narrow sinuous lamella postvaginalis, dian extension, hook-like or pointed and tongue-like. Valva and phalangifera is the only Stegotheca species (as far as is with costal band not broad, without row of spines or spine-like known) without a signum. setae; valva with small dense patch of dorsal setae situated towards base, in sacculus region, similar to that of Ischnop- Biological notes teris but not quite so well defined; setae deciduous, very long Moths have been found (some at an altitude of 140 m) in and fine. Transtilla lobes almost meeting medially (longer than September, November, and December. in other species of Stegotheca). Processes of anellus well developed, situated posterolaterally to juxta and extending to Distribution transtilla, claw-shaped, curved and tapered to pointed apex. French Guiana, Peru, and Brazil. Juxta smaller than in other species of Stegotheca, but with long medio-posterior furca-like process; process sometimes Stegotheca speculifera (Bastelberger) comb. nov. spinulose or rugose. Aedeagus with two lightly sclerotized Figs 77, 93,142,143,198 and tongue-like posterior extensions, one usually finely spinulose and the other without spinules; vesica with group of spines Ischnopteris speculifera Bastelberger, 1908: 62. Holotype , forming sclerotized strip along caecum. PERU (SMF) [examined]. FEMALE GENITALIA. Unknown. Material examined Form A: 5 ; including 4 genitalia preparations (AMNH, Diagnosis CMNH, SMF, ZSM). Stegotheca speculifera is readily recognized and defined by Holotype , Peru: [Huanuco,] Pozuzo, 800 m (type specimen several characters, notably the modified fovea-like structure of number SMFL 284, genitalia slide number 1685/04, SMF). the fore wing, and the modified frenulum and retinaculum. The

Downloaded by [Michigan State University] at 11:29 24 December 2014 Localities of additional material. French Guiana: Saut male genitalia of speculifera are very different from those of Caouenne. Peru: Cuzco, Quincemil; [Huanuco,] Pozuzo. other species of Stegotheca in having a pseudouncus (as in Bolivia: Prov[incia] del Sara. Rucana mediosecta), and well-developed processes of the anellus (as in Ischnopteris andRucana). Despite its anomalies, Form B: 1 ; including genitalia preparation (BMNH). Loc- the species is placed in Stegotheca because it has 'metallic' ality: Panama: no further data. scales on the abdomen, the presence of which are autapomorphic for the genus. Form unknown: 1 (INBio) [digital photograph]. Locality: Costa Rica: Alajuela Prov[ince], Upala, Albergue Hel- iconias, 700 [m], 3–4.xi.2000 (Delgado). Variation One male examined differs in some genitalic features, and Description comes from Panama, whereas the others are from South ADULTS (Figs 77, 93). Fore wing length: , 15-16 mm. America. Until more specimens are available for examina- Front of head projecting at lower edge; scale covering not tion I am treating these as two forms of speculifera. Form B dense, even in specimens in good condition. Wing pattern (the specimen from Panama) has longer processes of the anel- (only male known): fore wing mid brown, mottled, with whit- lus and a more distinctly spinulose process of the juxta. There ish flecks, and with four brownish orange longitudinal streaks is variation also within form A, particularly in the size of the towards outer margin, between veins; without dark median dorsal pseudouncus. 86 Linda M. Pitkin

Biological notes ulatrix mainly membranous or with varying degree of sclerot- Moths have been found at altitudes of 450 m to 800 m, in ization (most pronounced in abnormipalpis); sometimes very January?, August, and November. long and slender, or sometimes fairly bulbous; signum absent.

Distribution Diagnosis Form A: French Guiana, Peru, and Bolivia. Form B: Panama. Moths of Rucana are like smaller versions of Ischnopteris but Form unknown: Costa Rica. without a modified male retinaculum (except in the anomalous species mediosecta). Rucana moths can often be recognized by the shiny and swollen front of head, but, in the expanded Rucana Rindge concept of Rucana given here, not all species share this character. The fore wing marking of a small dark wedge between Rucana Rindge, 1983: 216. Type species: Ischnopteris whitish streaks near the tornus is similar to that seen in certain abnormipalpis Warren, 1904a, by original designation. other, generally larger, nacophorine moths, e.g. the 'Ischnop- Group 3 of Rindge (1983). teris' discolor-group, and Quillaca. Moths of two species of Rucana (chaconi and mediosecta) bear a superficial resemb- Description lance to certain Neotropical larentiine moths that have sim- ADULTS (Figs 78-90). Fore wing length: , 12-21 mm; , ilar orange wing markings: Spargania augustaria (Herrich- 13-17 mm. Antenna simple and filiform in both sexes, thicker Schaffer) and Euphyia crispa (Druce). in male. Front of head swollen, shiny and free of scales in Rucana is defined here primarily by autapomorphic char- many but not in all species. Labial palpus often with dark acters of the male genitalia, notably the dorsally swollen uncus. brown and yellow, or sometimes plain yellow, inner face; third The uncus is shaped like a duck's head, particularly where the segment extremely long and slender in abnormipalpis, very extreme apex of the uncus is flattened like a bill. The only ex- short in all other species. Wing pattern: fore wing mottled ception is the provisionally placed species mediosecta, which dark brown with red flecks, with olive-green and sometimes has a pseudouncus but otherwise has genitalia typical of Ru- white markings in irregular transverse bands; with small dark cana. A pronounced dorsal tuft of slender scales on the uncus brown wedge between whitish streaks at dorsum, near tornus; occurs in Rucana, but also in Quillaca, although the latter hind wing plainer, grey-brown or whitish with grey-brown genus is readily recognized by its unique and highly distinct- outer marginal markings, or with orange region. Fore wing ively modified valva (as in Pitkin, 2002: fig. 372). Rucana's fairly elongate, without fovea. Male retinaculum usually not bulbous lobes of the transtilla are conspicuous, but similar ones enlarged, but slightly enlarged and with transparent central occur in the 'Ischnopteris' discolor-group and occasionally in 'window' in one species provisionally placed here: medio- Ischnopteris. secta. Dorsum of male fore wing without median tuft or lobe. In the female, Rucana differs from Ischnopteris and Hind wing of male with anal margin usually not expanded (but Stegotheca in never having a signum. The pair of pocket-like moderately expanded and with hair-scale tuft in bisecta, and depressions of the lamella antevaginalis, often seen in Rucana, weakly also in mediosecta). Metathorax with dorsal tuft, abdo- might be indicative of a relationship with Ischnopteris as sim- men with longitudinal row of weak dorsal tufts. Male abdomen ilar structures are common in thefabiana-group of that genus. usually long and very slender; sternite 3 with triangular patch The transfer of three species to Rucana in Pitkin (2002: of setae. 224), brunneoviridis, degener and fidelis, has now been con- MALE GENITALIA (Figs 144-153,199-208). Uncus swollen firmed by dissection. dorsally but with apex somewhat flattened (except for medio-

Downloaded by [Michigan State University] at 11:29 24 December 2014 secta, which has a pseudouncus); uncus bearing dense dorsal Biological notes tuft of slender or hair-like scales, which are often slightly Moths of Rucana are known from elevations ranging from capitate. Gnathos small and not strongly developed, with one 200 m to 3000 m. Nothing is known about their early stages narrow and pointed median extension. Valva with costal band, or hostplants. but without processes. Transtilla often forming distinct pair of bulbous lobes. Processes of anellus situated between juxta Distribution and transtilla; processes narrow, except for broad base, with Rucana occurs in South America, as far south as Peru and slightly curved, pointed apex; juxta diamond-shaped, with Brazil. moderately elongate, tapered or occasionally tongue-shaped medio-posterior process, often covered with minute spinules. Aedeagus with posterior end lightly sclerotized and wrinkled; Checklist of species vesica usually with cornuti, a group of slender spines. abnormipalpis (Warren, 1904a) (Ischnopteris) bisecta (Dognin, 1914) (Ischnopteris) comb. nov. [from FEMALE GENITALIA (Figs 242-249). Lamella postva- Chrysomima] ginalis often with pair of pocket-like depressions; lamella brunneoviridis (Warren, 1905b) (Ischnopteris) antevaginalis forming a curved or V-shaped band, or occasion- chaconi sp. nov. ally semicircular or heart-shaped. Antrum forming collar, not degener (Warren, 1905b) (Ischnopteris) a complete ring, and without posterior projection. Bursa cop- fidelis (Warren, 1904a) (Ischnopteris) Neotropical geometrid moths 87

marmorata (Dognin, 1913) (Ischnopteris) comb. nov. [from fore wing (Fig. 83). Male genitalia with spines of vesica Ischnopteris] situated well away from sclerotized part of the aedeagus mathani sp. nov. (Figs 203, 208); female genitalia with long, narrow bursa mediosecta (Warren, 1909) (Ischnopteris) comb. nov. [from copulatrix (Fig. 246), or unknown 9 Ischnopteris]. Placement in Rucana provisional. 9. Labial palpus plain yellow, except sometimes at tip. Male sclerovesica sp. nov. genitalia with prominent spinose sclerite in vesica of aedeagus, in addition to usual group of spines (Fig. 208).. Key to species sclerovesica sp. nov. 1. Hind wing with orange region 2 - Labial palpus mottled with brown. Male genitalia with — Hind wing without orange region 4 usual group of spines in vesica of aedeagus, but without 2. Orange marking on hind wing confined to a large apical additional spinose sclerite (Fig. 203) degener blotch (Fig. 79) bisecta (form A) 10. Fore wing of both sexes without distinct postmedial line, — Orange marking extending across middle and to base of hind wing of male two-tone: white or cream in costal one- wing (Figs 82, 88) 3 third or more and greyish brown in dorsal half (Fig. 86). 3. Orange region covering costal half of hind wing, with no marmorata more than small dark brown marking at apex (Fig. 82).. - Fore wing with distinct whitish postmedial line, hind wing chaconi sp. nov. not two-tone (Fig. 80) bisecta (form B) — Orange region covering middle of hind wing, with large dark brown marking at apex (Fig. 88) mediosecta Species 4. Front of head shiny, free of scales (e.g. Fig. 90) 5 Rucana abnormipalpis (Warren) — Front of head not shiny, not free of scales although some- Figs 78, 90,144,199, 242, 243 times with thin covering 10 5. Labial palpus with third segment extremely long and Ischnopteris abnormipalpis Warren, 1904a: 116. LECTO- slender abnormipalpis TYPE , PERU (BMNH), here designated [examined]. — Labial palpus with third segment very short 6 Rucana abnormipalpis (Warren); Rindge, 1983: 217. 6. Fore wing dark and fairly plain, hind wing much paler but with contrasting dark band at outer margin (Fig. 81). Male Material examined genitalia with median process of the juxta a squarish plate Typical form: 12 , 18 (including 3 , 4 genitalia prepar- (Fig. 146B); female genitalia with fairly bulbous bursa ations) (BMNH, SMNS, USNM). copulatrix constricted medially (Fig. 244) Lectotype , Peru: SE. Peru, Carabaya, Santo Domingo, 1830 brunneoviridis m ('6000 ft'), xii.1901 (Ockenden) (genitalia slide Geom: — Fore wing various, with pronounced whitish patch (as in 21538, BMNH). Paralectotype, 1 , data as lectotype, but Fig. 84) or without; hind wing various, with or without xi. 1902 (BMNH). band at outer margin, but band usually diffuse. Male genitalia with median process of the juxta narrow; female Localities of additional material. Ecuador: Zamora- genitalia with bursa copulatrix either narrow, or bul- Chinchipe, Parque Nacional Podocarpus, [Rio] Bombuscara. bous but not constricted medially (Figs 246, 247, 249).. Peru: Carabaya: Santo Domingo; Rio Inambari, La Oroya; 7 Tinguri; Chirimayo. Brazil: [Amazonas] ('Upp. Amazon'), 7. Fore wing with postmedial line strongly angled towards Fonte Boa. outer margin, hind wing in both sexes not much paler than fore wing (Fig. 87). Male genitalia with median process of Form B: 2 (genitalia preparations of both) (ZSM). Downloaded by [Michigan State University] at 11:29 24 December 2014 the juxta extending well beyond processes of the anellus (Fig. 151); female genitalia as in Fig. 249 Localities. Bolivia: [La Paz,] Sarampiuni, San Carlos. mathani sp. nov. — Fore wing with postmedial line not strongly angled, or if Description strongly angled then hind wing much paler than fore wing, ADULTS (Figs 78,90). Fore wing length: , 12-14 mm; , 13- at least in males. Male genitalia with median process of 15 mm. Front of head swollen and shiny, free of scales. Labial the juxta not extending beyond processes of the anellus palpus with third segment extremely long and slender. Wing (Figs 148, 149); female genitalia as in Figs 246, 247.... pattern: fore wing of both sexes mottled dark brown with red 8 flecks, and with olive-green postmedial, antemedial and sub- 8. Fore wing of male with pronounced whitish patch on marginal bands and other markings; postmedial line slightly outer side of postmedial line; hind wing of both sexes angled towards outer margin of wing or almost straight (male), not strongly paler than fore wing (Fig. 84). Male genitalia or moderately angled (female). Hind wing paler than fore wing, with the vesica spines situated not far from sclerotized part sometimes only slightly; brown or grey with darker but dif- of the aedeagus (Fig. 204); female genitalia with elongate fuse band at outer margin; and faint postmedial line; without oval bursa copulatrix (Fig. 247) fidelis orange region. Markings on underside of wings varying, those — Fore wing usually without pronounced whitish patch; if on hind wing sometimes more, sometimes less, distinct, than present then hind wing of both sexes much paler than on upper side. 88 Linda M. Pitkin

MALE GENITALIA (Figs 144, 199). Uncus with slight Forest Res[erve]. Ecuador: Zamora-Chinchipe, Rio San subapical swelling, without pseudouncus. Processes of anellus Francisco, Estacion Cientifica San Francisco. evenly tapered, claw-shaped. Median process of juxta a narrow Form B: 1 specimen (including genitalia preparation) blade, covered with minute spinules except at pointed apex, ex- (ZSM). Locality: Peru: [Junin], Rio Oxabamba, 'Hda Mo- tending beyond processes of anellus. Aedeagus without spines sela'. or spinules at posterior end; vesica with group of long slender spines situated well away from sclerotized part of aedeagus. Description FEMALE GENITALIA (Figs 242,243). Lamella postvaginalis ADULTS (Figs 79, 80). Fore wing length: , 17-21 mm. Front with pair of pocket-like depressions situated directly postero- of head swollen but not free of scales. Labial palpus with dorsally to pair of small lobes (typical form), or instead with third segment very short. Wing pattern: fore wing dark brown large single pocket, weakly bilobed where it adjoins ostial re- mottled with straw and olive-green fine striations, without red gion (form B). Lamella antevaginalis a curved band, without flecks; with whitish postmedial line slightly wavy but other- spinules (typical form), or with a large field of minute spin- wise almost straight or slightly curved in direction of base of ules spaced evenly and close together, not forming short rows wing; form A with diffuse pale band (white and grey-green) (form B). Bursa copulatrix with long, narrow, partly sclerot- on outer side of that line; form B plain. Hind wing almost plain ized posterior section, and with shorter, bulbous, membranous, brown or grey, as dark as fore wing or slightly paler, with or- anterior sac (typical form), or bursa copulatrix a long mem- ange apical region (form A), without orange region (form B). branous sac with postero-lateral bulge (form B). Under-side of wings: fore wing of form A with yellowish markings (with brown flecks in them) forming postmedial band and Diagnosis apical blotch; fore wing of form B and hind wing of both forms The elongated third segment of the labial palpus is distinctive grey-brown dappled with pale, straw or yellowish, markings, and autapomorphic for abnormipalpis. In the male genitalia, but with darker postmedial and submarginal lines or markings; the median process of the juxta extends beyond the processes hind wing of form A additionally with yellow, brown-flecked, of the anellus as in mathani and mediosecta, but of the three apical region. species, only abnormipalpis has spines in the vesica of the aedeagus. The female genitalia of abnormipalpis are charac- MALE GENITALIA (Figs 145, 200). Uncus with large, or terized in the typical form by an autapomorphy of the bursa moderately large, subapical swelling, without pseudouncus. copulatrix: its long, partly sclerotized, posterior section. The Processes of anellus curved, with broad base narrowing ab- genitalia of form B are very different, as described above. That ruptly to moderately long, narrow, posterior spine. Median form is known only from two females, from Bolivia; it may process of juxta tongue-shaped and tapered, with weakly de- prove to be a different species, but study of more material veloped minute spinules; not approaching apices of processes (particularly males) is needed to determine its status. of anellus. Aedeagus without spines or spinules at posterior end, almost smooth; vesica with strip of spines situated fairly near to sclerotized part of aedeagus. Biological notes Moths have been found at altitudes of 310-1990 m, throughout FEMALE GENITALIA. Unknown. the year. Diagnosis Distribution Rucana bisecta has the largest moths of the genus, and bi- Typical form: Ecuador, Peru, and Brazil. Form B: Bolivia. secta is the only species to have an orange apical blotch on the hind wing. Because of their size and colouring, the moths re- Downloaded by [Michigan State University] at 11:29 24 December 2014 Rucana bisecta (Dognin) comb. nov. semble some species of Ischnopteris (e.g. the chryses-group), Figs 79, 80, 145, 200 but they do not have the enlarged modified male retinaculum that is characteristic of Ischnopteris. Moths of bisecta are Ischnopteris bisecta Dognin, 1914: 407. Holotype , more likely to be confused with those of 'Ischnopteris' festa COLOMBIA (USNM) [examined]. or festiva, but in bisecta the postmedial line of the fore wing Chrysomima bisecta (Dognin); Parsons et al., in Scoble, 1999: is not bowed towards the outer margin of the wing, as it is 161. in 'Ischnopteris' festa or festiva. In form B of bisecta (which does not have an orange blotch), the almost straight postmedial Material examined line, and otherwise more or less plain wings, are distinguishing Form A: 5 (including 3 genitalia preparations) (SMNS, features. In the male genitalia, bisecta is the only Rucana spe- USNM, ZMUJ). cies with the processes of the anellus narrowing abruptly to a Holotype , Colombia: E. Colombia, [Cundinamarca,] Med- long, narrow, posterior spine, an autapomorphy for this species. ina, 500 m (Fassl) (Type No. 32545, genitalia slide 60150, R. bisecta was previously known only from the holotype. USNM).

Localities of additional material. Colombia: E. Biological notes Colombia, [Cundinamarca,] Medina; Cundinamarca, Moths have been found at altitudes of 500 m to 1850 m, in Monterredondo; Western Cordillera, [Cauca,] Tambito February, April, May, and December (form A), July (form B). Neotropical geometrid moths 89

Distribution margin), but the species is better distinguished and defined by Form A: Colombia and Ecuador; form B: Peru. genitalic characters. It is the only Rucana species with the median process of the juxta as a squarish plate, an autapomorphy Rucana brunneoviridis (Warren) for brunneoviridis. In the female genitalia the shape of the Figs 81,146, 201, 244 bursa copulatrix, being fairly bulbous and constricted medially, differs from those of other members of the genus; however, Ischnopteris brunneoviridis Warren, 1905b: 359. LECTO- only one female specimen of brunneoviridis is known. TYPE , PERU (BMNH), here designated [examined]. Rucana brunneoviridis (Warren); Pitkin, 2002: 224. Biological notes Moths have been found at altitudes of 200 m to 1990 m, in Material examined April, November, and December. 2 , 2 (genitalia preparations of all) (AMNH, BMNH).

Lectotype , Peru: SE. Peru, Carabaya, Santo Domingo, 1990 Distribution m ('6500 ft'), xii.1902 (Ockenden) (genitalia slide Geom: Peru. 20742, BMNH). Paralectotype, 1 , data as lectotype but iv. 1902 (BMNH). Rucana chaconi sp. nov. Localities of additional material. Peru: Carabaya: Santo Figs 82,147, 202, 245 Domingo; Madre de Dios, Tambopata Reserve, Explorers' Material examined Inn. 6 , 3 (including 2 , 1 genitalia preparations)

Description Holotype , Costa Rica: San Jose Province, P[ar]k Nac[ional] ADULTS (Fig. 81). Fore wing length: , 14-15 mm; , 15- Braulio Carrillo, Estacion Carrillo, 700 m, vii.1984 (´I. & A. 17 mm. Front of head swollen and shiny, free of scales. Chacon) (genitalia slide LMP 413; INBio). Labial palpus either plain yellow except at tip, or mottled with Paratypes: Colombia: 1 , Muzo, 400-800 m (Fassl) brown, with third segment very short. Wing pattern: fore wing (BMNH). Costa Rica: 1 , A[la]j[uela Province], of both sexes mainly mottled dark brown with some red flecks, Bijagua, 700 m, 3–4.xi.2000 (Becker) (VOB); 1 [digital with olive-green postmedial and antemedial bands and other photograph], Alajuela Prov[ince], Upala, Albergue markings not strongly pronounced; postmedial line slightly to Heliconias, LN 423500-300250, 700 m, 3-4.xi.2000 moderately angled towards outer margin of wing (male), or (Delgado) (INBio); 1 , Cartago Prov[ince], Para´ıso, more strongly angled (female). Hind wing much paler than P[ar]que Na[ciona]l Tapanti, Est[acion] Queb[rada] Se- fore wing, whitish, mottled with grey-brown, contrasting with gundo, 100 [m?] N., 100 [m?] W, start of Send[ero] fairly well-defined grey-brown band at outer margin; without La Pava, LN 194459-560500, 1400 m, vii.1999 (Del- orange region. Under-side of hind wing with more dark mark- gado) (INBio); 1 [digital photograph], Guanacaste ings than on upper side. Prov[ince], 9 km S. of Santa Cecilia, Estacion Pitilla, LN 329950-380450, 700 m, iv.1995 (Moraga) (INBio); MALE GENITALIA (Figs 146A-B, 201). Uncus with large 1 [digital photograph], data as before but LN 330200- subapical swelling, without pseudouncus. Processes of anellus 380200, 700 m, 21-29.xi.1992 (Moraga) (INBio); 1 , evenly tapered, claw-shaped. Median process of juxta a large, Heredia Province, Braulio Carrillo N[ational] P[ark], angular, squarish plate, covered with minute spinules; apex 10°16.1'N 84°05.1'W, 1070 m, 22.v.2003 (Brehm) truncate or shallowly excised, not reaching apices of processes (BMNH). 1 , data as holotype (INBio). of anellus. Aedeagus with a group of short spinules, and with Downloaded by [Michigan State University] at 11:29 24 December 2014 one triangular spine, at posterior end; vesica with a group of long slender spines situated not far from sclerotized part of Description aedeagus. ADULTS (Fig. 82). Fore wing length: , 13-14 mm; , 15 mm. Front of head swollen; not free of scales although with thin FEMALE GENITALIA (Fig. 244). Lamella postvaginalis with coverage. Labial palpus with third segment very short. Wing pair of pocket-like depressions distinct or weak, situated dir- pattern (both sexes): fore wing mottled mid to dark brown or ectly postero-dorsally to pair of lobes. Lamella antevaginalis blackish, with varying amount of pale yellowish grey-green a curved or V-shaped band, with large field of minute spinules markings including postmedial line; the latter strongly angled forming short rows. Antrum short. Bursa copulatrix fairly long, towards outer margin but often indistinct; area between ante- narrowing medially, between fairly bulbous posterior and an- medial and postmedial lines usually red-tinged dark brown, terior parts; posterior part with large, lightly sclerotized region, particularly towards dorsum of wing. Hind wing two-tone, or- anterior part entirely membranous. ange in costal half and mid to dark brown or blackish in dorsal half, two zones clearly demarcated by an almost straight or Diagnosis slightly curved line; dark brown or black spot present or ab- Moths of brunneoviridis can usually be recognized by their sent at apex in orange region. Under-side of fore wing with combination of wing markings (fore wing dark and fairly plain, basal half to two-thirds largely orange but sometimes with hind wing much paler but with dark, distinct band at outer some brown flecks or larger brown area towards dorsum, 90 Linda M. Pitkin

occasionally orange region much reduced and wing then pre- Cochabamba, Chapare, Paracti; La Paz, Unduavi — Coroico; dominantly dark brown; hind wing orange except for dark Rio Songo; Yungas, Forestal. brown blotch near tornus and sometimes dark brown spot at apex. Description ADULTS (Fig. 83). Fore wing length: , 12-16 mm; , 12- MALE GENITALIA (Figs 147, 202). Uncus with large subap- 16 mm. Front of head swollen and shiny, free of scales. La- ical swelling, without pseudouncus. Processes of anellus claw- bial palpus mottled with brown, with third segment very short. shaped, tapering strongly from broad base. Median process of Wing pattern: fore wing of both sexes mottled mid to dark juxta narrow, tapered, and short, not approaching apices of brown and olive-green, usually with some whitish flecks and processes of anellus, not curved; without spinules. Aedeagus often some reddish flecks or patches, with pale olive-green without spines or spinules at posterior end; vesica with group or whitish postmedial and antemedial bands, male sometimes of slender spines situated well away from sclerotized part of with whitish marking on outer side of postmedial line but aedeagus. rarely as pronounced as in fidelis; postmedial line angled to- FEMALE GENITALIA (Fig. 245). Lamella postvaginalis with wards outer margin of wing, usually slightly to moderately in pair of pocket-like depressions, without pair of lobes situated male, but occasionally strongly, and often strongly angled in postero-anteriorally to these. Lamella antevaginalis moderate- female. Hind wing usually much paler than fore wing, whit- sized (compare Fig. 245 with Fig. 248 of marmorata) and ish, with grey-brown margin and submarginal band varying semicircular, covered with minute spinules spaced evenly and from weak to distinct, but wing sometimes not that pale, and close together, not forming short rows. Antrum short. Bursa occasionally as dark as fore wing; hind wing without orange copulatrix extremely long, narrow, mainly membranous. region. Under-side of wings mottled grey-brown and whitish, fore wing markings generally fainter than on upper side, and Diagnosis under-side of hind wing not very pale. Rucana chaconi is closely related to marmorata but differs in MALE GENITALIA (Figs 148, 203). Uncus with slight to having a wing pattern (as described above) that is unique in the moderate subapical swelling, without pseudouncus. Processes genus and an autapomorphy for the species. The new species of anellus evenly tapered, claw-shaped. Median process of also differs slightly from marmorata in several genitalic char- juxta moderately narrow, tapered or tongue-shaped, covered acters. In the male, the uncus is more swollen, the processes with minute spinules; sometimes reaching apices of, but not of the anellus are less strongly curved, and the aedeagus does extending beyond, processes of anellus. Aedeagus usually with not have a patch of spinules (as is present in marmorata). The posterior end forming two lobes close together, but not always female genitalia of both species are mainly membranous with with both lobes clearly defined; often with spinules on one of an extremely long bursa copulatrix, but chaconi has a smaller the lobes. Vesica without bulbous caecum; vesica with group lamella antevaginalis than marmorata. of spines situated well away from sclerotized part of aedeagus, rarely without spines. Biological notes Moths have been found at altitudes of 400-800 m, to 1400 m, FEMALE GENITALIA (Fig. 246). Lamella postvaginalis with in April, May, July, and November. pair of pocket-like depressions fairly distinct or occasionally weak, with distinct or weak pair of lobes situated postero- Distribution anteriorally to these. Lamella antevaginalis a curved band, or Costa Rica and Colombia. heart-shaped, weakly covered with minute spinules usually forming short rows but occasionally evenly spaced or absent. Rucana degener (Warren) Antrum fairly short. Bursa copulatrix long, narrow, mainly Downloaded by [Michigan State University] at 11:29 24 December 2014 membranous but with sclerotized patch halfway (very approx- Figs 83,148, 203, 246 imately) along its length; with membranous postero-lateral Ischnopteris degener Warren, 1905b: 360. Holotype , PERU bulge, usually small but occasionally large. (BMNH) [examined]. Rucana degener (Warren); Pitkin, 2002: 224. Diagnosis Moths of degener vary considerably in wing pattern, and may Material examined be confused with other species. However, there are some subtle 25 , 8 (including 11 , 4 genitalia preparations) (AMNH, distinctions: the majority of degener moths have a paler hind BMNH, CMNH, SMNS, USNM, ZSM). wing than those of fidelis, more white flecks on the fore wing than in brunneoviridis, and a more brown-flecked labial palpus Holotype , Peru: SE. Peru, Carabaya, Santo Domingo, 1990 than in sclerovesica. The male genitalia resemble those of m ('6500 ft'), x.1902 (Ockenden) (genitalia slide Geom: fidelis and sclerovesica in having evenly tapered claw-shaped 21543, BMNH). processes of the anellus, with a tapered or tongue-shaped and Localities of additional material. Ecuador: Prov[ince] spinulose juxta process that does not extend beyond the anellus Zamora-Chinchipe, Estacion Cientifica San Francisco. ´Peru: processes. The uncus of degener is swollen less than that of Carabaya, Santo Domingo; Carabaya, La Oroya; N. fidelis, but usually slightly more than that of sclerovesica. More Peru, Chaupe. Bolivia: Cochabamba, Yungas de Corani; obviously, in degener, the spines of the vesica are situated Neotropical geometrid moths 91

further away from the sclerotized part of the aedeagus than orange region. Under-side of both wings with markings fainter they are in fidelis, and neither species has an additional spinose than on upper side. sclerite as in sclerovesica. The female genitalia of degener MALE GENITALIA (Figs 149, 204). Uncus with large do not have very distinctive features, but the bursa copulatrix subapical swelling, without pseudouncus. Processes of anel- differs clearly from that offidelis in being narrower and lacking lus evenly tapered, claw-shaped. Median process of juxta the postero-lateral sclerotization present in fidelis. tongue-shaped with pointed apex, narrow or moderately broad, covered with minute spinules; not extending beyond processes Variation of anellus. Aedeagus with distinctly bilobed posterior end, with Two males examined from Ecuador: Zamora-Chinchipe have spinules along one of the lobes; vesica with group of spines genitalic characters similar to those of fidelis and degener, but situated not far from sclerotized part of aedeagus. appear somewhat intermediate and are darker than usual in degener. A female from the same locality might be associated FEMALE GENITALIA (Fig. 247). Lamella postvaginalis with them; this specimen has genitalia not at all like those without pair of pocket-like depressions. Lamella antevaginalis of fidelis and rather resemble those of degener. Other than a wrinkled V-shaped band, without spinules. Bursa copulatrix mentioning these three specimens here, I have not included an elongate oval, with large postero-lateral caecum; sclerot- them under either species; their identity remains uncertain. ized and ridged region extending over posterior end of bursa and caecum. Biological notes Moths have been found at altitudes of 750-3000 m, in January, Diagnosis February, May, June, and September to December. Moths of fidelis can usually be recognized by their wing pattern, with a whitish patch (pronounced in males) on the dark Distribution fore wing, and with the hind wing fairly dark (at least in males). Ecuador to Bolivia. The male genitalia are similar to those of degener, but differ in having a more swollen uncus, and in having the spines of the vesica situated closer to the sclerotized part of the aedeagus. Rucana fidelis (Warren) In the female genitalia the large, partly sclerotized postero- Figs 84, 85,149, 204, 247 lateral caecum of the bursa copulatrix region is distinctive and Ischnopteris fidelis Warren, 1904a: 117. LECTOTYPE , a putative autapomorphy for the species. PERU (BMNH), here designated [examined]. Rucana fidelis (Warren); Pitkin, 2002: 224. Biological notes Moths have been found at altitudes of 500 m to 1990 m, in Material examined January, April, and September to November. 7 , 3 (including 3 , 2 genitalia preparations) (BMNH, USNM). Distribution Colombia and Peru. Lectotype , Peru: Carabaya, Santo Domingo, 1830 m ('6000 ft'), xi. 1901 (Ockenden) (genitalia slide Geom: 21541, BMNH). Rucana marmorata (Dognin) comb. nov. Figs 86,150, 205, 248 Localities of additional material. Colombia: E. Colombia, [Cundinamarca,] Medina. Peru: Carabaya, Santo Ischnopteris marmorata Dognin, 1913: 32. Holotype ,

Downloaded by [Michigan State University] at 11:29 24 December 2014 Domingo (specimens include 1 paralectotype of brunneovi- COLOMBIA (USNM) [examined]. ridis, abdomen missing, but moth appears to be fidelis); Carabaya, La Oroya; Rio Inambari, La Oroya. Material examined Typical form: 6 , 1 (including 4 , 1 genitalia preparations) (AMNH, USNM, ZSM). Description Holotype , Colombia: E. Colombia, [Cundinamarca,] Med- ADULTS (Figs 84,85). Fore wing length: ,14-15 mm; , 14- ina, 500 m (Fassl) (Type No. 31184, genitalia slide 60141, 15 mm. Front of head swollen and shiny, free of scales. Labial USNM). palpus with third segment very short. Wing pattern: fore wing of both sexes mottled dark brown with olive-green markings, Localities of additional material. Colombia: E. and with a few reddish flecks or occasionally large patches, Colombia, [Cundinamarca,] Medina; Cundinamarca, with postmedial line white; this line in male slightly angled Monterredondo. Bolivia: Cochabamba, Chapare, Cristal towards outer margin of wing, and with pronounced whitish Mayo; La Paz, from Caranavi to Santa Ana on Alto Rio Beni patch on outer side of line; line more strongly angled in female, road. and whitish patch more diffuse and at least partly on inner side of line. Hind wing almost plain brown, usually paler than fore Form B: 2 (genitalia preparations of both) (AMNH). wing but not strongly so; sometimes with darker band at outer Localities. Peru: Cuzco, Cosnipata, Paucartambo. margin but not well-defined or strongly contrasting; without Bolivia: [Cochabamba,] Chapare. 92 Linda M. Pitkin

Description Rucana mathani sp. nov. ADULTS (Fig. 86). Fore wing length: , 13-15 mm; Figs 87,151, 206, 249 ,15 mm. Front of head swollen; not free of scales but with Material examined thin covering. Labial palpus with third segment very short. 5 , 2 (including 4 , 1 genitalia preparations) Wing pattern (both sexes): fore wing dark brown, mottled with varying amount of pale greyish olive markings; postmedial and Holotype , Ecuador: Bol´ıvar Province, Balzapamba, ix. 1893 other lines indistinct. Hind wing of male two-tone but some- ii.1894 (deMathan) (genitaliaslideGeom: 21540; BMNH). times weakly demarcated, white or cream in costal one-third Paratypes: Colombia: 1 , Choco, San Juan, La Selva, or more, and greyish brown, sometimes pale, in dorsal part; 1410 m ('4600 ft'), ix.1909, (Joicey Bequest) (BMNH). hind wing of female greyish brown shading paler costally but Ecuador: 2 , 1 , data as holotype (BMNH); 1 , Pichin- not demarcated into two zones; without orange region. Under- cha Prov[ince], Quito to Santo Domingo, 1200 m, 25.ii.1965 side of both sexes with markings fainter than on upper side, (Pena) (AMNH). except for dark apex and outer margin of fore wing, and often submarginal and postmedial lines on hind wing. Excluded from the type series: 1 [abdomen missing], MALE GENITALIA (Figs 150A-B, 205). Uncus with fairly Ecuador: Pichincha, 59 km W. [of] Quito, 8 km W. of slight to large subapical swelling, without pseudouncus. Pro- Chiriboga, Las Palmeras, 24.x. 1988 (Miller) (AMNH). cesses of anellus evenly tapered, strongly curved and claw- shaped. Median process of juxta very narrow and spine- Description like, curved, with pointed apex; sometimes with, but often ADULTS (Fig. 87). Fore wing length: , 14-15 mm; , 15- without, minute spinules; moderately long but not extending 16 mm. Front of head swollen and shiny, free of scales. La- beyond processes of anellus (typical form), or small (form B). bial palpus with third segment very short. Wing pattern (both Aedeagus with or without field of spinules near posterior end; sexes): fore wing mainly dark brown with some red flecks and vesica with group of spines situated well away from sclerotized patches, with olive-green and whitish postmedial and ante- part of aedeagus. medial bands and other markings fairly pronounced (more so in males than in females), but not forming pronounced patch; FEMALE GENITALIA (Fig. 248). Lamella postvaginalis with postmedial line strongly angled towards outer margin. Hind pair of pocket-like depressions. Lamella antevaginalis large wing almost plain grey-brown, not much paler than fore wing; (compare Fig. 248 with Fig. 245 of chaconi) and heart-shaped, without orange region. Under-side of both wings usually fairly covered with minute spinules spaced evenly and close together, plain although fore wing with three small pale brown markings not forming short rows. Antrum fairly short. Bursa copulatrix near outer margin. extremely long, narrow, mainly membranous. MALE GENITALIA (Figs 151, 206). Uncus with slight to moderate subapical swelling, without pseudouncus. Processes Diagnosis of anellus fairly broad, narrowing abruptly to small curved Male moths of marmorata can usually be recognised by their pointed apex. Median process of juxta very narrow and spine- two-tone (whitish and grey-brown) hind wing pattern, but like, covered with minute spinules; very long, extending well females are less distinctive. Neither sex has the scale-free, beyond processes of anellus. Aedeagus without spines or spin- shiny front of head common in Rucana, and the only other ules at posterior end, almost smooth; vesica without spines. Rucana moths without this character differ from marmorata in having orange on the hind wing, or (bisecta (form B)) a FEMALE GENITALIA (Fig. 249). Lamella postvaginalis distinct whitish postmedial line on the fore wing. In the male without pair of pocket-like depressions. Lamella antevaginalis genitalia, the median process of the juxta is more narrow and a curved band, without spinules. Bursa copulatrix long, narrow Downloaded by [Michigan State University] at 11:29 24 December 2014 spine-like in marmorata than in other species of Rucana, with but with bulbous anterior end, mainly membranous. the exception of mathani, in which the juxta process is longer and extends beyond the processes of the anellus. [See under Diagnosis chaconi for further other differences.] The female genitalia of Moths of mathani are fairly dark-winged, resembling those of marmorata are similar to those of chaconi in that both are fidelis, but with the postmedial line on the fore wing angled mainly membranous with an extremely long bursa copulatrix, more strongly, at least in males, and without the pronounced but marmorata has a larger lamella antevaginalis than that of white patch of fidelis males. The male genitalia of mathani are chaconi. distinctive and definitive in having a very narrow and spine- like median process of the juxta, extending well beyond the processes of the anellus (unlike that of marmorata), putatively Biological notes autapomorphic for mathani. The female genitalia do not have Moths of the typical form have been found at altitudes of 500 any obvious apomorphies, but the species differs from others of m to 1420 m, in January, September, and December. Moths of Rucana in the absence of sclerotization in the bursa copulatrix, form B have been found in August and November. and in the absence of a pair of pocket-like depressions in the lamella postvaginalis. The only other exception is form B of Distribution abnormipalpis, and that has a postero-lateral bulge in the bursa Colombia to Bolivia. copulatrix, which is not present in mathani. Neotropical geometrid moths 93

Variation with a brown band extending along both dorsum and outer mar- One female specimen differs slightly from the description gin. Also unique in the genus is the enlargement of the male above in having stronger markings on the under-side of the retinaculum, with a transparent central 'window'. The male wings, including a pale band at the outer margin of the fore genitalia are highly distinctive: mediosecta is the only Rucana wing in place of smaller markings. This specimen, which has species with a pseudouncus. In that respect (and also in hav- lost its abdomen, is not included in the type series. ing an Ischnopteris-like retinaculum) mediosecta resembles Stegotheca speculifera, a species without orange hind wings. Biological notes The systematic position of these species is problematic as both Moths have been found at altitudes of 1200 m to 1960 m, in have Rucana-like features of their male genitalia, but I have February, September, and October. placed speculifera in Stegotheca because it has the apomorphy most clearly defining that genus, one or more dorsal patches Distribution of 'metallic' scales on the abdomen, a character not present in Colombia and Ecuador. mediosecta.

Rucana mediosecta Warren, comb. nov. Biological notes Figs 88,152, 207 Moths have been found at altitudes of 610 m and 900 m, in August and December. Ischnopteris mediosecta Warren, 1909: 102. Holotype , PERU (BMNH) [examined]. Distribution Material examined Ecuador and Peru. 2 (genitalia preparations of both) (BMNH, USNM). Rucana sclerovesica sp. nov. Holotype , Peru: Carabaya, Rio Huacamayo, La Union, 610 Figs 89,153, 208 m ('2000 ft'), xii.1904 (Ockenden) (genitalia slide Geom: 21015, BMNH). Material examined 4 (genitalia preparations of all) Locality of additional material. Ecuador: Napo, Simon Bol´ıvar, Coca River Canyon. Holotype , Ecuador: Zamora-Chinchipe, old road Loja- Zamora, 04°01.46'S 79°01.68'W, 1380 m, 30.x.2000 Description (ID#8328) (Sussenbach¨) (genitalia slide LMP 471; SMNS). ADULTS (Fig. 88). Fore wing length: , 16-17 mm. Front of Paratypes: Venezuela: 1 , Merida, Pedregosa, 3000 m head swollen but not free of scales. Labial palpus with third (Briceno) (BMNH). Ecuador: 1 , Prov[ince] Zamora- segment very short. Wing pattern (only male known) : fore wing Chinchipe, Estacion Cientifica San Francisco, 3°58'S mottled dark brown, with scattered small whitish markings 79°04'W, 1850m, 11.v.1999 (ID#3530) (Sussenbach¨) mainly from indistinct postmedial line to outer margin of wing; (SMNS); 1 , Zamora-Chinchipe, Estacion Cientifica San hind wing with large central orange region, and with broad Francisco, 3°58'S 79°04'W, 1845 m, 11.x.1999 (Brehm) dark brown band around margin (including a large dark brown (SMNS). marking at apex of wing). Under-side of fore wing mainly brown but some with diffuse orange markings, particularly at Additional specimen excluded from the type series: Ecuador: costa and in region of postmedial line; under-side of hind wing 1 [identity unconfirmed, abdomen missing], E. Ecuador, Rio similar to upper side. Male retinaculum slightly enlarged and Pastaza, El Rosario, 1500 m ('4900 ft') (Palmer) (BMNH).

Downloaded by [Michigan State University] at 11:29 24 December 2014 with transparent central 'window'. Description MALE GENITALIA (Figs 152, 207). Uncus divided into two ADULTS (Fig. 89). Fore wing length: , 13-15 mm. Front parts: a ventral process, and a dorsal pseudouncus bearing a of head swollen and shiny, free of scales. Labial palpus plain dorsal tuft of fine setae or hair-like scales. Processes of anellus yellow, except sometimes at tip, with third segment very short. claw-shaped, curved and tapered strongly from broad base to Wing pattern (only male known): fore wing mottled dark pointed apex. Median process of juxta very narrow but with brown and olive-green, with some white flecks and reddish base more bulbous; densely covered with minute spinules; very flecks or patches, with pale olive-green postmedial and other long, extending well beyond processes of anellus. Aedeagus diffuse transverse lines, without whitish marking on outer side smooth, without spines or spinules at posterior end; vesica of postmedial line as in fidelis and some specimens of de- without spines. gener; postmedial line slightly to moderately angled towards FEMALE GENITALIA. Unknown. outer margin of wing. Hind wing slightly paler than fore wing, grey or brown, shading to dark margin and diffuse submarginal Diagnosis band; hind wing without orange region. Under-side of wings The hind wing colouration of mediosecta is unique in the mottled grey-brown, straw and whitish, fore wing markings genus and an autapomorphy for this species; some other generally fainter than on upper side, and under-side of hind Rucana species have orange and brown hind wings, but not wing with dark postmedial line and submarginal band. 94 Linda M. Pitkin

MALE GENITALIA (Figs 153, 208). Uncus with very slight BASTELBERGER, M.J. 1908. Weitere Geometriden aus meiner subapical swelling, without pseudouncus. Processes of anellus Sammlung. Entomologische Zeitschrift, Frankfurt a. M. 22, 58-59, fairly evenly tapered, claw-shaped. Median process of juxta 61-63. BREHM, G. 2003. Host-plant records and illustrations of the larvae of tongue-shaped or tapered, varying in width, covered with 19 geometrid moth species from a montane rainforest in Ecuador. minute spinules; not reaching apices of processes of anel- Nachrichten des Entomologischen Vereins Apollo 24, 29-34. lus. Aedeagus with posterior end tapered, not forming two BROWN, K.S. 1979. Ecologia geografica e evoluçao nas florestas lobes close together, sometimes wrinkled but without spin- neotropicais. Apendice I. 21 pp. Sao Paulo. ules. Vesica forming bulbous caecum with strongly wrinkled DELFÍN-GONZÁLEZ, H. & BEUTELSPACHER, C.R. 1986. Catalogo de la coleccion Roberto Muller (Lepidoptera) del Museo de Historia area, lightly or well sclerotized, extending from sclerotized Natural de la Ciudad de Mexico. XV. Famila [sic] Geometridae. part of aedeagus to a prominent sclerite bearing three to four Anales del Instituto de Biologia, Universidad Nacional Autonoma sturdy spines; rest of vesica narrow and with group of smaller de México (Serie Zoologia)´ 57 (2), 425-478. spines situated well away from sclerotized part of aedeagus. DIAS, M.M. 1998. Consideraçoes taxonomicas sobre o genero Cund- inamarca Rindge (Lepidoptera, Geometridae) e descriçao de uma FEMALE GENITALIA. Unknown. nova especie. Revista Brasileira de Zoologia 15, 951-958. DOGNIN, P. 1904. Hétérocères nouveaux de l'Amérique du Sud. An- nales de la Société entomologique de Belgique 48, 115-134; 358- Diagnosis 369. Moths of sclerovesica resemble those of degener, but the two DOGNIN, P. 1911. Hétérocères nouveaux de l'Amérique du Sud. Mem- species can be distinguished by their labial palpi: plain yel- oires de la Société entomologique de Belgique 18, 151-188. low except sometimes at the tip in sclerovesica, as opposed DOGNIN, P. 1912. Hétérocères nouveaux de l'Amérique du Sud. to flecked with brown in degener. The two species (and also Memoires de la Société entomologique de Belgique 19, 121- 177. fidelis) have similar male genitalia, although the uncus of de- DOGNIN, P. 1913. Hétérocères nouveaux de l'Amérique du Sud. Mem- gener is least swollen in sclerovesica, but sclerovesica has a oires de la Societé entomologique de Belgique 22, 1-54. very obvious and unique apomorphy in the aedeagus struc- DOGNIN, P. 1914. Hétérocères nouveaux de l'Amérique du Sud. Án- ture, the spinose sclerite that is additional to the usual group nales de la Societé entomologique de Belgique 57, 380-417. of spines on the vesica. DOGNIN, P. 1923. Hétérocères nouveaux de l'Amérique du Sud 21, 38 pp. Rennes. DRUCE, H. 1891-1900 (1893). Biologia Centrali-Americana. Lepid- Biological notes optera - Heterocera 2, 1-622; 3, 101 pls. Moths have been found at altitudes of 1380 m to 3000 m, in FELDER, R. & ROGENHOFER, A.F 1875. Reise der osterreichischen May and October. Fregatte Novara um die Erde (Zool.) 2 (Abt.2) (5): pls 121-140. Wien. FLETCHER, D.S. 1979. In: NYE, I.W.B., Ed., The Generic Names of Distribution Moths of the World 3, Geometroidea. xx + 243 pp. London. Venezuela and Ecuador. GIACOMELLI, E. 1911. Lepidopteros riojanaos noevos opoco con- scidos. Anales de la Sociedad Cientifica Argentina 72, 19- 40. Acknowledgements GUENéE, A. [1858] (dated 1857). Uranides et Phalenites 2. In: I am most grateful for the loan and provision of a wealth of BOISDUVAL, J.B.A.D. & GUENÉE, A., Eds., Histoire naturelle des material (including digital images), essential for this work, and Insectes, Species général des Lépidoptéres 10, 1-584; Atlas; pls for much useful information from the following: Vitor Becker, 1-22. Brazil; Gunnar Brehm, Universitat Bayreuth, Bayreuth, Germany HOLLOWAY, J.D. [1994], dated 1993. The moths of Borneo: Family [Gunnar Brehm's collection accrued from recent Neotropical stud- Geometridae, subfamily Ennominae. Malayan Nature Journal 47, ies is now deposited mainly in the SMNS]; Niels Christensen, 1-309, 593 figs, 19 colour pls. ZMUC; Bernardo Espinoza and Isidro Chacon, INBio; Patricia HÜBNER, J. 1816—[1825]. Verzeichniss bekannter Schmettlinge [sic]. Gentili-Poole and Alma Solis, USNM; Axel Hausmann, ZSM; 431 pp. Augsburg. Downloaded by [Michigan State University] at 11:29 24 December 2014 Wolfram Mey, MNHU; Wolfgang Nassig, SMF; Eric Quinter, HÜBNER, J. [1823] [1820-1826] (dated 1806). Sammlung exotischer AMNH; John Rawlins, CMNH, Bo Sullivan, USA, and Janusz Schmetterlinge. 2, 4 pp. (index), 225 pls. Augsburg. Wojtusiak, ZMUJ. Roger Bristow, Devon, U.K., was a helpful cour- KLOTS, A.B. 1956. Lepidoptera. pp. 97-111 in TUXEN, S.L., Ed., ier of specimens to and from the AMNH. My visit to the ZMUC Taxonomist's Glossary of Genitalia in Insects. Copenhagen. was supported by a COBICE grant. Daniel Janzen, University of MCQUILLAN, P.B., YOUNG, C.J. & RICHARDSON, A.M.M. 2001. A re- Pennsylvania, Philadelphia, USA, generously provided the valuable vision of the Australian moth genus Paralaea Guest (Lepidoptera: addition of original data on biology, and the use of his photographs of Geometridae: Ennominae). Invertebrate Taxonomy 15, 277-317. larvae, from DHJ and the Janzen and Hallwachs Caterpillar Rearing MöSCHLER, H.B. 1882. Beitrage zur Schmetterlings-Fauna von Database at http://janzen.sas.upenn.edu/. Surinam. IV. Verhandlungen der Zoologisch-Botanischen Gesell- Special thanks are due to Shayleen James (BMNH) for the schaft in Wien 31, 393-442, pls 17, 18. tremendous technical assistance she has provided, with dissections PARRA, L.E. & BEECHE, M.A. 1986. Omaguacua longibursae n. sp.: and with illustrations and preparation of plates, and also to Tara Lewis nuevo geometrido para Chile (Lepidoptera: Geometridae). Boletin (BMNH volunteer), for all the work she did on illustrations. I also wish de la Sociedad de Biologia de Concepcion 57, 137-143. to thank Malcolm Scoble and other colleagues at the BMNH, and also PARSONS, M.S., SCOBLE, M.J., HONEY, M.R., PITKIN, L.M., & the referees, for helpful comments on the manuscript. PITKIN, B.R. 1999. The Catalogue. In: SCOBLE, M.J., Ed., Geometrid Moths of the World: a Catalogue (Lepidoptera: Geometridae). xxv, 1016 pp. (2 volumes). CSIRO Publishing, References Collingwood. PITKIN, L.M. 1993. Neotropical Emerald moths of the genera AGASSIZ, L. 1847. Nomenclator Zoologicus. Fascicle 12 (Index uni- Nemoria, Lissochlora and Chavarriella, with particular refer- versalis). Zoological Society of London. ence to the species of Costa Rica (Lepidoptera: Geometridae, Neotropical geometrid moths 95

Geometrinae). Bulletin of The Natural History Museum (Ento- STOLL, C. [1780-1782]. In: Cramer, P., De uitlandsche Kapellen 4, mology) 62, 39-159, pl. 1. 29 + 252 pp., pls 289–400. Amsterdam & Utrecht. PITKIN, L.M. 2002. Neotropical ennomine moths: a review of the gen- VALDECASAS, A.G., BECERRA, J.M. & MARSHALL, D. 1997. Extend- era (Lepidoptera: Geometridae). Zoological Journal of the Linnean ing the availability of microscopic type material for taxonomy and Society 135, 121–401. research. Trends in Ecology and Evolution 12, 211-212. PITKIN, L.M., MORA, R.A, & SCOBLE, M.J. 1996. A checklist to the WALKER, F. 1854-1866. List of the specimens of lepidopterous insects Ennominae (Geometridae) of Costa Rica: taxonomy for a national in the collection of the British Museum. 1862, part 25: 1281-1477; biodiversity inventory. Gayana Zoologia 60, 121-155. 1866, part 35: 1535-2040. London. PROUT, L.B. 1928. New Geometridae. Novitates Zoologicae 34, 53- WARREN, W. 1895. New species and genera of Geometridae in the 70. Tring Museum. Novitates Zoologicae 2, 82-159. PROUT, L.B. 1929. New species and subspecies of Geometridae. Novit- WARREN, W. 1900. New genera and species of American Drepan- ates Zoologicae 35, 63-77. ulidae, Thyrididae, Epiplemidae, and Geometridae. Novitates RINDGE, F.H. 1983. A generic revision of the New World Nacophorini Zoologicae 7, 117-225. (Lepidoptera, Geometridae). Bulletin of the American Museum of WARREN, W. 1904a. New American Thyrididae, Uraniidae, and Geo- Natural History 175, 147-262, figs 1-124, tables 1-9. metridae. Novitates Zoologicae 11, 1-173. SCHAUS, W. 1901. New species of Geometridae from tropical WARREN, W. 1904b. New American Thyrididae, Uraniidae, and Geo- America. Part II. Transactions of the American Entomological metridae. Novitates Zoologicae 11, 493-582. Society 27, 241-276. WARREN, W. 1905a. New Thyrididae, Uraniidae, and Geometridae SCHAUS, W. 1912. New species of Heterocera from Costa Rica. XV. from South and Central America. Novitates Zoologicae 12, 41-72. Annals and Magazine of Natural History 9, 423–433. WARREN, W. 1905b. New American Thyrididae, Uraniidae, and Geo- SCHAUS, W. 1939. New Neotropical Lepidoptera of the family Noto- metridae. Novitates Zoologicae 12, 307-379. dontidae. Annals of the Carnegie Museum 27, 321-348, pls 31-33. WARREN, W. 1907. American Thyrididae, Uraniidae, and Geomet- SOMMERER, M.D. 2002. To agree or not to agree: the question of ridae in the Tring Museum. Novitates Zoologicae 14, 187-323. gender agreement in the International Code of Zoological Nomen- WARREN, W. 1909. New American Uraniidae and Geometridae in the clature. Nota lepidopterologica 25, 191-204. Tring Museum. Novitates Zoologicae 16, 69-109.

Index (Accepted names are in roman and synonyms are in italics. Principal references are in bold.)

abnormipalpis 15, 86-88, 87, 92 chlorosata 54 albiguttata 48, 63, 64, 67, 68 choritis 54 albimargo 80 chryses 17,46, 58-60, 62, 63, 66, 67, 74 albipennis 80-84, 81 chryses-group 48,49, 57-64, 67, 70, 73, 74, 88 albirupta 63 Chrysomima 15, 17 Amblurodes 15, 17 Cidariophanes 15, 80 amissa 15, 80-83, 82, 85 commixta 17, 51, 52 Anischnopteris 15, 17 conjugens 78 apicemaculata 54 constrictus 80, 81 augustaria 86 costiplaga 81-84, 82 aurudaria 47, 58-60, 64, 69, 75 crispa 86 Cundinamarca 15 beckeri 48, 49 cythereata 15, 80, 81, 83, 84 bermejona 15 Downloaded by [Michigan State University] at 11:29 24 December 2014 bifinita 17,46,47,49, 51-53, 55, 66 Dasciopteryx 15 bifurcata 47, 48, 65, 68 degener 86, 87, 90, 91, 93, 94 bipectinata 17, 48, 65, 66, 77 discolor 78 bisecta 86-88, 92 discolor-group 77-79, 86 brehmi 16,46,47, 59, 60, 63, 66, 67, 74 dispar 59 brunneoviridis 86, 87, 89, 90 Erilophodes 15 bryifera 17, 47, 51, 52, 54 Euphyia 86 callistrepta 79 fabiana 16, 48, 49, 53, 54, 57 Canelo 13, 15, 80 fabiana-group 17, 48, 49, 51, 53, 57, 66, 77, 86 catocalata 47, 64, 67,75,76 fasciata 47, 61, 64, 57, 70, 73 Ceratonyx 15 fassli 47, 57-60, 59, 62, 67 chaconi 16, 86, 87, 89, 90, 92 festa78, 79, 88 Charca 15, 17 festiva 78, 79, 88 chavesi 16, 46, 47, 57-60 fidelis 86, 87, 90-92, 91-94 chlorata 16, 17, 48, 65, 68 chloroclystata 16,48, 64, 68, 69 hirsuta 47, 61, 65, 68,70,73,75 chlorophaearia 17, 48, 51, 53-56 hoffmani 47, 59, 60 96 Linda M. Pitkin

inornata 48, 57, 61, 63, 70, 73 parvula 82 illineata 48, 54, 55 pexatata 49, 51,53 inconspicua 71 phalangifera 80, 81, 83-85, 84 Ischnopteris 13-17, 16, 46-81, 83, 85, 86, 88, 93 Plesiophyle 15, 80 Ischnopterix 15, 16 praeluteata 63 Ischnopteryx 16, 80 prognata 67 projectata 80 janzeni 47, 57, 59, 61, 62, 64 pronubata 17, 47, 56, 64, 67, 69,72,74-76 juvencata 69 Quillaca 15, 17, 78, 80, 86 klagesi 17,48,49, 55-57, 72 rostellaria 48, 53, 55-57, 56 lata 47, 59, 62 Rucana 13-17, 78, 80, 85, 86-93 latijuxta17, 48, 70, 71,73 lemoulti 48, 71, 72 Salasaca 13, 15, 80 sclerovesica 87, 90, 91, 93, 94 Mallomus 14 seriei 48, 72, 75, 76, 77 Marcodava 15 Spargania 86 marmorata 87, 90-92, 91 speculifera 15, 80, 81, 85, 93 mathani 87, 88, 92 spinea 80, 82 mediosecta 15, 85-88, 93 Stegotheca 13-15, 17, 80-86, 93 Mimophyle 15, 80 stenoptila 47, 64, 67, 75, 76 miseliata 48, 71, 72 Syrtodes 16 multistrigata 47, 49, 57, 63, 72,76 trimaculata 54 Nazca 15 Trichostichia 15, 17 obfuscata 53, 54 velledata 16, 51, 52 obtortionis 47, 61,70,71, 73, 77 versipennis 17 ochroprosthia 17, 47, 73, 74 viridifascia 80 ockendeni76 viriosa 80 Oratha 15 watsoni 47, 59-63, 62 Pagrasana 15 xylinata 17, 48, 49, 75-77, 76 palmeri 46,47, 59, 64, 67, 74 Parasynegia 15 Yashmakia 15 Downloaded by [Michigan State University] at 11:29 24 December 2014