A Revision of the Neotropical Beetle Genus Nordus ... - Senckenberg

Entomologische Abhandlungen 62 (1): 3–64

© Museum für Tierkunde Dresden, ISSN 0373-8981, 30.06.2004

A Revision of the Neotropical Beetle Genus Nordus Blackwelder (Insecta: Coleoptera: Staphylinidae: Xanthopygina) STYLIANOS CHATZIMANOLIS Division of Entomology, Natural History Museum and Biodiversity Research Center, and Department of Ecology and Evolutionary Biology, The University of Kansas, Lawrence, KS, 66045-7523, USA [[email protected]] Abstract. The staphylinid genus Nordus Blackwelder, 1952 is revised. Lectotypes are designated for 19 species: Brachydirus aeneipennis Bernhauer, B. antennatus Sharp, B. banghaasi Bernhauer, B. batesi Sharp, B. championi Sharp, B. cribricollis Sharp, B. densiventris Bernhauer, B. divisus Sharp, B. erosus Bernhauer, B. fungicola Sharp, B. gebieni Bernhauer, B. maculiceps Sharp, B. major Bernhauer, B. occipitalis Bernhauer, B. simplex Sharp, B. solitarius Sharp, B. styloceros Sharp, B. xanthocerus Nordmann, and Staphylinus testaceus Fabricius. Neotypes for B. picticornis Solsky and B. zischkai Scheerpeltz are designated. Eleven species are described as new: N. amphivolos sp.n., N. cherylae sp.n., N. dichromos sp.n., N. elegans sp.n., N. elytisi sp.n., N. omios sp.n., N. periergos sp.n., N. portokalis sp.n., N. seferisi sp.n., N. semelius sp.n., and N. stomachoponos sp.n. Brachydirus chapareanus Scheerpeltz is placed into synonymy with B. nigroscutellatus Bernhauer; B. divisus Sharp and B. gracilis Sharp with B. terminalis Sharp; B. ecuadorensis Scheerpeltz and B. occipitalis Bernhauer with B. solitarius Sharp; B. erosus Bernhauer and B. schadei Scheerpeltz with S. velitaris Erichson; and B. schuberti Wendeler with B. maculiceps Sharp. Distribution maps, identification keys and illustrations of structural features including aedeagi are provided for the known species. The presence of color morphs in some species and secondary sexual features are discussed. Key words. Staphylininae, Staphylinini, polymorphic species, coloration, secondary sexual structures.

Contents 1. Introduction 1.1. General 1.2. Taxonomic history 2. Materials and methods 3. Abbreviations 4. Descriptions of taxa 4.1. Nordus Blackwelder 4.2. Nordus aeneiceps (Sharp) 4.3. Nordus aeneipennis (Bernhauer) 4.4. Nordus amazonicus (Sharp) 4.5. Nordus amphivolos sp.n. 4.6. Nordus antennatus (Sharp) 4.7. Nordus banghaasi (Bernhauer) 4.8. Nordus batesi (Sharp) 4.9. Nordus championi (Sharp) 4.10. Nordus cherylae sp.n. 4.11. Nordus cribricollis (Sharp) 4.12. Nordus densiventris (Bernhauer) 4.13. Nordus dichromos sp.n. 4.14. Nordus diversiventris (Bernhauer) 4.15. Nordus elegans sp.n. 4.16. Nordus elytisi sp.n. 4.17. Nordus facialis (Erichson) 4.18. Nordus fungicola (Sharp) 4.19. Nordus gebieni (Bernhauer)

1.

3 3 5 5 6 6 6 8 8 9 10 11 12 13 14 16 17 18 19 20 21 22 23 24 26

5. 6. 7. 8.

4.20. Nordus longipes (Sharp) 4.21. Nordus maculiceps (Sharp) 4.22. Nordus major (Bernhauer) 4.23. Nordus nigroscutellatus (Bernhauer) 4.24. Nordus omios sp.n. 4.25. Nordus periergos sp.n. 4.26. Nordus picticornis (Solsky) 4.27. Nordus portokalis sp.n. 4.28. Nordus seferisi sp.n. 4.29. Nordus semelius sp.n. 4.30. Nordus simplex (Sharp) 4.31. Nordus solitarius (Sharp) 4.32. Nordus speciosus (Bernhauer) 4.33. Nordus stomachoponos sp.n. 4.34. Nordus styloceros (Sharp) 4.35. Nordus terminalis (Sharp) 4.36. Nordus testaceus (Fabricius) 4.37. Nordus velitaris (Erichson) 4.38. Nordus xanthocerus (Nordmann) 4.39. Nordus zischkai (Scheerpeltz) 4.40. Keys to species of Nordus Designation of lectotypes and neotypes and synonymies Discussion Acknowledgements References

27 27 29 30 31 31 32 33 34 35 36 37 39 40 40 42 43 44 45 46 47 51 51 52 52

Introduction

1.1. General The subtribe Xanthopygina was proposed by SHARP (1884) as the ‘group’ Xanthopygina and originally included the genera Tympanophorus Nordmann, Selma Sharp (= Elmas), Xanthopygus Kraatz, Lampropygus Sharp (= Xanthopygus), Brachydirus Nordmann (= Nordus), Plociopterus Sharp, Philothalpus Kraatz, Gastrisus Sharp, Isanopus Sharp, Styngetus Sharp, Ocyolinus Sharp, Phanolinus Kraatz, Nausicotus Sharp, and Glenus Kraatz, all of them distributed in the New World tropics. Since then, Xantho-

pygina have been treated as a subtribe by most specialists (HERMAN 2001; NAOMI 1982, 1983a, 1983b; NEWTON & THAYER 1985; NEWTON et al. 2001; SMETANA 1995; SMETANA & DAVIES 2000), but also as a tribe (HAYASHI 1997) or a subfamily (MOORE 1964). Today, the subtribe Xanthopygina is comprised of 37 genera which include a total of approximately 385 species. Adult xanthopygines are among the largest in body size and the most strikingly colored of all staphylinids. Many are over 2.5 centimeters

4

CHATZIMANOLIS: Revision of Nordus

Figs. 1–8. Representative species of Nordus. 1: N. dichromos black morph, 10 mm. 2: N. dichromos golden-orange morph, 10 mm. 3: N. velitaris golden-orange/black morph, 10.5 mm. 4: N. speciosus, 12 mm. 5: N. elegans, 9 mm. 6: N. testaceus, 9.5 mm. 7: N. semelius, 9 mm. 8: N. zischkai, 9.5 mm.

long and have bodies dominated by shining metallic colors or by dramatically contrasting reds, blacks, and yellows. Most of them have a bright yellow tip at the end of the

abdomen (segment VIII and sometimes VII). This feature is the source of the name Xanthopygina from the Greek words ‘␰␣␯␪ó␵’ (blond) and ‘/␷␥␫␦␫␱␯’ (narrow rump).

5

Entomologische Abhandlungen 62 (1)

Despite the striking appearance of adult Xanthopygina, their natural history and behavior are almost completely unknown; there are behavioral observations only for the genera Xanthopygus (QUEZADA et al. 1969), Nordus (CHATZIMANOLIS 2000, 2003), and Xenopygus. Members from these genera are found on rotting mushrooms and fruits, but detailed observations have been made for very few species. Species of most xanthopygine genera are found in the neotropics, but a few are distributed in the Oriental, Ethiopian, Palearctic and Australian regions. The few Oriental genera were revised by NAOMI (1982, 1983a, 1983b) and HAYASHI (1997), but these authors often included genera that now have been transferred to other subtribes (see HERMAN 2001 for the current classification). A phylogenetic treatment of Xanthopygina is lacking (but is now being completed by Chatzimanolis), and this taxon is probably not monophyletic as it is currently recognized. Nonetheless, its members can be identified by the structure of the pronotum: the superior line of the pronotal hypomeron is continued to the anterior edge of the pronotum without deflection and it is completely visible from above. In addition, the known larvae tend to have very elongate urogomphi (A.F. Newton Jr. pers. comm.). The genus Nordus is distributed from Mexico to Argentina and is unusual in the subtribe Xanthopygina in that males have strikingly developed secondary sexual characters on the posterior abdominal sterna, which show great variation among species. In addition, the punctuation pattern of the head and the variability of the shape of the antennomeres are unique among genera of the xanthopygines. SHARP (1876, 1884) hypothesized that Nordus is closely related to Plociopterus and Elmas, but these relationships have not been rigorously tested. No further speculations can be made until there is a phylogenetic hypothesis for the subtribe. CHATZIMANOLIS (2000, 2003) studied the natural history and behavior of Nordus fungicola (Sharp). The larva of the same species is presently being described (CHATZIMANOLIS in press a), while no other larvae are known for this genus.

1.2. Taxonomic history NORDMANN (1837) first described the genus Brachydirus to accommodate the species B. xanthocerus. ERICHSON (1839) did not consider it a valid genus and transferred it to the ‘Fam. III’ of the genus Staphylinus, along with two new species that he described and one described earlier by FABRICIUS (1801). KRAATZ (1857) re-validated the genus in a footnote of his monograph on the German beetle fauna. Additional species were described by BERNHAUER (1905, 1917a, 1917b, 1926), SCHEERPELTZ (1971), SHARP (1876, 1884), SOLSKY (1872), and WENDELER (1926). BLACKWELDER (1952) recognized that ‘Brachydirus Nordman [sic], 1837’ was a junior homonym of ‘Brachydirus Smith Woodward, 1811’ (a genus in Pisces) and proposed the replacement name ‘Nordus’. SCHEERPELTZ (1971) provided a key to the species of Nordus (as Brachydirus Nordmann) that was inaccurate and almost impossible to use. Recently, CHATZIMANOLIS & ASHE (2004) synonymized N. horni

(Bernhauer) with Plociopterus variegatus Sharp. In this revision, I synonymize eight previously erected species names, describe 11 new species, and provide keys and distribution maps for all known species of Nordus. The phylogenetic relationships among species of Nordus, along with a study of the evolution of the coloration in these beetles, will be presented elsewhere (CHATZIMANOLIS in press b).

2. Materials and methods Dried specimens were first relaxed in warm soapy water, then apical abdominal segments containing the aedeagus were dissected from the abdomen. Mouthparts of selected individuals were also dissected for study and scanning electron micrography. The apical abdominal segments were cleared using 10% KOH, and the aedeagus removed from the inside of the abdomen. Body structure, mouthparts, and aedeagi were studied using an Olympus SZ-60 dissecting microscope. Illustrations were made with a camera lucida on an Olympus SZX9 dissecting microscope. SEM photographs were taken using a LEO 1550 Field Scanning Electron Emission Microscope. Photographs were prepared using a Microptics ML-1000 Digital Imaging System. Relative width of the gula is given as a ratio of least width (narrowest transverse distance between the sutures) to length (the distance along the gula midline between the line connecting the posterior tentorial pits and the anterior margin of the gula). Density of punctures on the head, pronotum, and elytra is expressed in terms of the average number of punctures in a transverse linear distance of 0.3 mm measured (with an eyepiece micrometer) in the middle of the anterolateral right quadrant of the appropriate body part (epicranium of head, pronotum, or right elytron); in this procedure each puncture touched by the line is counted. In the ‘Diagnosis’ sections emphasis has been given for a selection of characters that will identify both sexes. However, males from most species can be easily identified simply by examining the antenna (Figs. 22–57), the aedeagus (Figs. 122–255), and sternum VIII (Figs. 69–103). Comments on the specimen labels are included in brackets [ ] in the citations of the specimen labels in the ‘Material’ sections. Distribution maps were created based on a careful review of the available locality data (label data cited in the ‘Material’ sections below). They do not include specimens for which only the country or collector was mentioned on the label. SCHEERPELTZ (1971) gave distribution records for some species that are incorrect, no specimens exist for these localities in his collection or in any other collection that I examined. HERMAN (2001) in his catalogue of the Staphylinidae mentioned these distributions from SCHEERPELTZ (1971). Terms for structural features follow BLACKWELDER (1936), SMETANA & DAVIES (2000), and ASHE & CHATZIMANOLIS (2003). The evolutionary species concept outlined by WILEY & MAYDEN (2000) was used in this revision.

6


3. Abbreviations Specimens used in this revision were from the following collections: AMNH American Museum of Natural History, New York, USA BMNH The Natural History Museum, London, Great Britain CASC California Academy of Sciences, San Francisco, USA CNCI Canadian National Collection, Ottawa, Canada CZUG Centro de Estudios en Zoología CUCBA, Universidad de Guadalajara, Zapopan, Mexico DEI Deutsches Entomologisches Institut, Eberswalde, Germany FMNH Field Museum of Natural History, Chicago, USA INBC INBio, Santo Domingo de Heredia, Costa Rica ISNB Institut Royal des Sciences Naturelles de Belgique, Brussels, Belgium MCZC Museum of Comparative Zoology, Harvard University, Cambridge, USA NMW Naturhistorisches Musem Wien, Vienna, Austria SEMC Snow Entomological Collection, Natural History Museum/Biodiversity Research Center, University of Kansas, Lawrence, USA UNCB Museuo de Historia Natural, Instituto de Ciencias Naturales, Universidad Nacional de Colombia, Bogota, Colombia UNSM University of Nebraska State Museum, Lincoln, USA USNM National Museum of Natural History, Washington, DC, USA ZMUC Zoological Museum, University of Copenhagen, Copenhagen, Denmark ZMHB Museum für Naturkunde der Humboldt-Universität, Berlin, Germany Collecting methods: FIT

Flight intercept trap

4. Description of taxa 4.1. Nordus Blackwelder, 1952 (Figs. 1–8) Brachydirus Nordmann, 1837: 131 [preoccupied, non Brachydirus Smith Woodward, 1811 (Pisces)]; KRAATZ (1857: 538). Staphylinus; ERICHSON (1839: 353). Nordus Blackwelder, 1952: 262, new name for Brachydirus Nordmann, 1837. Type species. Brachydirus xanthocerus Nordmann, 1837 fixed by BLACKWELDER (1952) by objective synonymy of Nordus Blackwelder with Brachydirus Nordmann, for which B. xanthocerus had already been established as the type species by monotypy. Description. L e n g t h . 6.7–13 mm. H e a d . With short microsetae and longer black macrosetae along margins. Postclypeus in same plane as frons. Epicranium continuously covered with large punctures (as in Fig. 17); on anterior part punctures smaller, very dense and uniformly distributed, on posterior part larger and less dense. Medial area between antennae flat. Eyes large and centered in middle part of lateral margin of head. Postmandibular carina present, complete and prominent to base of head; remainder of head beneath eyes rounded and sparsely irregularly punctuate. Gula with transverse impression very well developed. Antennae 1.5–2ⴛ length of head. Antennomeres short but shape variable (Figs. 16, 22–57); 1st as thick as 2nd, 11th

with 2 circumferential rows of prominent macrosetae, one near middle and one near apex; 7th–10th transverse, not appearing ‘serrate’ (anterior and posterior margins of antennomere subequal in length). Labrum as in Fig. 12. Mandibles typical of Staphylininae (BLACKWELDER 1936), prominent, with setose prostheca and a single small tooth, as in Fig. 10. Maxilla as in Fig. 11, with 4 palpomeres; 4th subcylindrical, pointed apically, 1.5–2ⴛ length of 3rd, with sparse setation; 3rd with numerous setae apically; 2nd about 2ⴛ length of 3rd, strongly curved. Stipes with multiple macrosetae; galea elongate, galea and lacinia with dense brush-like setae. Labium as in Fig. 9; with 3 palpomeres; 3rd subcylindrical, pointed apically, with sparse setation, about 1.5ⴛ length of 2nd; 2nd with long macrosetae mesally, 2ⴛ length of 1st. Ligula very short, membranous, with short emargination medially. Mentum and submentum of equal width, submentum slightly longer than mentum. Hypopharynx well developed. T h o r a x . Pronotum subquadrate to quadrate; lateral margins convex to concave in dorsal aspect; lateral parts more or less flat and not curved downwards; punctures large (as in Fig. 15), not uniformly distributed (except in N. major); punctuation variable among species; disc with impunctate area medially, which extends length of pronotum and is usually broadest anteriorly; pronotum with long black macrosetae along margins. Micropunctures on metepimeron absent. Scutellum prominent, with yellow microsetae and medium-sized punctures. Mesobasisternum without pair of prominent setae. Elytra slightly longer than pronotum, with medium-sized to large punctures, uniformly punctuated; microsetae directed posterolaterally; macrosetae present along margins. Hind wings typical of Staphylininae (BLACKWELDER 1936), fully developed. L e g s . Protibia with a series of large subapical spurs and many macrosetae (as in Figs. 13, 18), meso- and metatibia with shorter subapical spurs. Tarsal formula 5-5-5. Protarsus with enlarged tarsomeres, 1st–4th uniformly covered with spatulate setae ventrally, 1st smaller than 2nd. Metatarsus longer than mesotarsus, otherwise similar, tarsomeres 2–4 ventrally lobed. In mesotarsomeres 1st elongate, 2ⴛ length of 2nd (except in N. speciosus), 5th elongate similar to 1st. Pretarsal claws very slender and sharply curved near base. Legs completely covered with long macrosetae. A b d o m e n . Paired prototergal glands present on tergum I. Terga III and IV with tergal basal carina and subbasal (arch-like) carina; V–VII with tergal basal carina only, but in some species V with subbasal carina in addition. Setation and punctuation patterns on terga variable. Tergum IX forming an acutely rounded apex, with long black prominent macrosetae near apex. Sterna III–V with moderate transverse impressions, VI with faint transverse impression, shining. In most species segment VII with black iridescent setae at least in anterolateral quarters. Cerci long and either straight or slightly curved, covered with long black macrosetae. Male and female genitalia. Typical of Staphylininae (BLACKWELDER 1936). Female: Spermatheca not sclerotized; sternum IX of female divided into two lobes, each with large basal part of coxa IX, distal part of coxa IX, and very small apical stylus. Male: Aedeagus with large tube-like median lobe and single (dorsomedially fused) paramere. Paramere with various combinations of sensory spinules (= peg-like setae, see HAMMOND 1972 for photographs and discussion) and apical setae.


7

Figs. 9–12. Typical mouthparts of a Nordus species, here of N. amazonicus. 9: Labium and hypopharynx (hp), dorsal aspect, scale 0.2 mm. 10: Left mandible, with tooth (to) and prostheca (ps), dorsal aspect, scale 0.1 mm. 11: Right maxilla, ventral aspect, galea (gl), lacinia (lc), stipes (st), scale 0.2 mm. 12: Labrum, dorsal aspect, scale 0.1 mm. Figs. 13–18. 13: Right protibia, dorsal aspect, of N. amazonicus, scale 0.3 mm. 14: Right mesotibia and tarsus of N. speciosus, scale 0.6 mm. 15: Pronotum of N. cribricollis, scale 0.6 mm. 16: Antenna of N. major, showing differences in microtrichae between antennomeres 5 and 6, scale 0.2 mm. 17: Head in dorsal aspect, showing punctuation, here in N. aeneipennis, scale 0.3 mm. 18: Left protibia and tarsus of N. fungicola, scale 0.3 mm.

S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (Figs. 61–65) with an emargination in most species; hind margin of sternum VIII (Figs. 69–103) variable; sternum IX as in Figs. 104–112, with apical processes of varied length, with or without median emargination. Female: with no apparent secondary sexual structures except in some species hind margin of sternum VIII exceeding lateral margins, forming a moderately rounded to pointed extension (Figs. 113–121). Diagnosis. Nordus differs from (nearly) all xanthopygines in the following features: large prominent eyes that are at least 0.6ⴛ length of the head (similar only in Allostenopsis, Elmas, Philothalpus and some species of Plociopterus); head anteriorly with large, dense and contiguous punctures (almost all other xanthopygines with a medial area lacking punctures; Elmas has dense, contiguous punctures but these are smaller and have the same density anteriorly and poster-

iorly); antennae short (2ⴛ length of head; usually 3ⴛ in other xanthopygines); antennomeres 3–5 without microtrichae (antennomeres 1–2 without microtrichae in all xanthopygines, in xanthopygines other than Nordus microtrichae starting on antennomere 4 or 5); antennomere 11 with 2 circumferential rows of prominent macrosetae, one near middle and one near apex (other xanthopygines with several circumferential rows of prominent macrosetae near middle and one near apex); pronotum with lateral parts more or less flat and not curved downwards as in other xanthopygines; micropunctures on metepimeron absent; abdominal terga III–VII with setation not uniform as in other xanthopygines. Habitat. Known from elevations of 0–2300 m in wet tropical lowland and cloud forests. Specimens have been collected primarily in flight intercept traps, malaise traps or by sweeping the low forest vegetation. CHATZIMANOLIS

8

(2003) observed N. fungicola feeding on small drosophilid flies that were gathered on rotting palm fruits, rotting fungi, or dung. It is hypothesized that at least the species found in lowlands have similar habits. Distribution. Known from southern Mexico to northern Argentina (Figs. 19–21, 58–60, 66–68).

4.2. Nordus aeneiceps (Sharp, 1876) (Figs. 122–125) Brachydirus aeneiceps Sharp, 1876: 115. Nordus aeneiceps (Sharp); HERMAN (2001: 3579). Description. L e n g t h . 6.7 mm. C o l o r a t i o n . Head golden-orange with anterior 3/4 of epicranium metallic green; mandibles and labrum goldenorange; antennomeres all golden-orange. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII and posterior 1/2 of VII golden-orange. Coxae and femora dark brown to black; tibiae and tarsi dark orange; pro- and mesotrochanters dark orange, metatrochanter dark brown; legs orange around articulations of podomeres. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior 3/4 of epicranium relatively large, about 7.6 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior 1/4 about 4.5 punctures / 0.3 mm, distance between punctures as wide as 0.5 punctures. Length of eyes ca. 0.8ⴛ length of head, distance between eyes less than 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.09), gular sulci widely separated anteriorly and almost contiguous at midlength and posteriorly. Antennomeres according to SHARP (1876): 1st gradually thickened, club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 6th almost quadrate, 7th–10th wider than greatest length. T h o r a x . Pronotum quadrate, width : length = 0.94; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in at least 3 very dense, contiguous longitudinal rows, punctures of a row contiguous; large, slightly convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow and black microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with large punctures, about 3.2 punctures / 0.3 mm, punctures contiguous, microsetae mostly yellow and partly black. Legs with yellow and black macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with few punctures on anteromedial part of disc, without punctures on posterior half; V with many punctures laterally and anteriorly, without punctures posteriorly; VI–VII extremely punctuated; IV–V and VIII covered with yellow microsetae throughout, VI and VII with black microsetae and with yellow ones laterally. Paratergites VI–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: VI–VII on antero-


lateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–VIII with yellow microsetae on each side. Segment VII with black iridescent microsetae throughout. Cerci straight, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII with broad and deep emargination medially; sternum VIII with evenly curved ventral surface, hind margin with broad and deep U-shaped emargination medially; sternum IX with a pair of moderate apical processes and a moderate U-shaped emargination medially. Female: unknown. M a l e g e n i t a l i a . As in Figs. 122–125. Paramere in dorsal outline consistently converging to a pointed tip, apically subequal in length to median lobe and narrower; dorsally convex in lateral outline. Sensory spinules (only 6) arranged in an irregular band and restricted to tip. Median lobe with subtruncate or slightly emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus aeneiceps and N. simplex can be distinguished from other golden-orange species by the metallic green anterior part of the epicranium. They differ from each other as follows: punctures on anterior part of epicranium dense and not longitudinally elongate in N. aeneiceps but very dense and longitudinally elongate in N. simplex; cerci straight in N. aeneiceps but slightly curved in N. simplex; structure of aedeagus as shown in Figs. 122–125, 226–229; paramere tip in dorsal outline pointed in N. aeneiceps but rounded in N. simplex. Distribution. Ega, Brazil (Fig. 19). No specimens are known from Bolivia and Colombia as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Habitat. Unknown. Remarks. Only holotype known. Antennomeres 4–11 are missing and the description above was based on SHARP (1876). Material. Holotype, ‚ with labels ‘Type’, ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus aeneiceps ‚ Type D. S.’ (BMNH). SHARP (1876) states in the original description that he had available only a single specimen, which therefore is the holotype.

4.3. Nordus aeneipennis (Bernhauer, 1917) (Figs. 17, 22, 69) Brachydirus aeneipennis Bernhauer, 1917: 115. Nordus aeneipennis (Bernhauer); HERMAN (2001: 3579). Description. L e n g t h . 8.0–10.0 mm. C o l o r a t i o n . Head light red-brown; mandibles and labrum light brown; antennomeres 1–2 brownish-yellow, 3–7 brown, 7–11 yellowish-brown, 10 and basal half of 11 clouded with brown. Pronotum light red-brown, elytra brown, scutellum light brown. Abdomen light brown except: middle of tergum V clouded with dark brown; tergum VI dark brown to black but anterolateral angles and narrow posterior margin red-brown; tergum VII black but extreme anterolateral angles and posterior 1/4 goldenorange; paratergites dark brown-black; segment VIII golden-orange. Legs light brown except distal parts of pro- and mesofemora and -tarsi orange, anterior surface of protibiae and ventral surface of protarsi dark brown. H e a d . (Fig. 17) Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium large, about 5.3 punctures / 0.3 mm, longitudinally elongate, margins of

9


punctures confluent; in posterior half about 3.6 punctures / 0.3 mm, distance between punctures as wide as 0.5 punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 22): 1st thickened only in distal half, but narrower near distal end, longer than 2nd plus 3rd, 2nd subequal in length to 3rd, 5th subequal in length to 4th, 1st– 3rd longer than greatest width, 4th–6th subquadrate, 7th– 10th wider than greatest length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 0.94; anterolateral angles not prominent, subcariniform; lateral margins very concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in 1–2 sparse longitudinal rows separated by distance as wide as 0.5–1 punctures, punctures of a row not contiguous, but distance varies; large, convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayishyellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.6 punctures / 0.3 mm, interspaces as wide as 0.5–1 punctures, microsetae yellow throughout. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII well punctuated (V–VI less so medially); IV with yellow microsetae medially and posterolaterally; V with yellow microsetae throughout, VI with yellow ones laterally and brown ones medially, VII with brown ones. Paratergites IV–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–VI and VIII with yellow microsetae, VII with brown ones. Segments IV–VII with black iridescent microsetae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 69) with evenly curved ventral surface, hind margin with two moderate triangular lobes (tips pointed) and a moderate U-shaped emargination medially; sternum IX (identical to that in Fig. 110) with two moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a slightly pointed extension (identical to that in Fig. 115). M a l e g e n i t a l i a . Identical to those in Figs. 134–137. Paramere in dorsal outline consistently converging to a rounded tip, apically slightly shorter than median lobe and narrower; completely straight in lateral outline. Sensory spinules arranged in two very short longitudinal rows restricted to tip. Median lobe with a broad subacute apex in

dorsal outline, slightly upcurved apically in lateral outline, forming a small dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus antennatus, N. banghaasi, N. picticornis, N. aeneipennis and N. zischkai can all be distinguished from other species of Nordus by the following characters: punctures on anterolateral corners of sterna VI and VII less dense than on the rest of these sterna; punctures on anterior part of epicranium longitudinally elongate; and shape of sternum VIII in males (shape varies among these species, but overall is distinctive for this group of species). N. antennatus and N. banghaasi can be distinguished from N. picticornis, N. aeneipennis, and N. zischkai by antennomere 4 being longer than wide (subquadrate in other species). In N. picticornis head, pronotum and elytra are golden-orange, whereas in N. aeneipennis and N. zischkai head and pronotum are reddish-brown to black and elytra are dark brown to black. The very similar N. picticornis and N. aeneipennis are well distinguished by the shapes of the aedeagus (Figs. 210–213, 134–137) and sternum VIII (Figs. 97, 69). N. aeneipennis can by distinguished from N. zischkai by the following characters: color of head and pronotum reddishbrown and elytra dark brown (all black in N. zischkai); punctures on anterior part of the epicranium denser in N. aeneipennis than in N. zischkai; punctures on posterior part of epicranium shallow in N. aeneipennis and deep in N. zischkai; and in males the shape of the aedeagus is different (Figs. 134–137 for N. aeneipennis and Figs. 252–255 for N. zischkai). Distribution. Known from elevations of 330–1850 m in Bolivia and Peru (Fig. 19). Habitat. Unknown. Remarks. Collected in flight intercept traps. Material. Lectotype, ‚, here designated, from PERU, with labels ‘‚’, ‘1800 m, 11o3´S 75o17´W [in Peru] Green W. N. Iconnicoff 14.xi.1906’, ‘aeneipennis Bern. Typus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Lectotype Brachydirus aeneipennis Bernhauer des. Chatzimanolis 2003’ (in FMNH). – Paralectotype, „, from PERU, with labels ‘1000 m 11o3´S 75o17´W [in Peru] Green W. N. Iconnicoff 14.ix.1906’, ‘aeneipennis Bern. Cotypus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Paralectotype Brachydirus aeneipennis Bernhauer des. Chatzimanolis 2003’ (in FMNH). – BOLIVIA: ex coll. Klima (2 in NMW). PERU: Callanga (2 in FMNH, 2 in NMW); Chanchamayo (1 in BMNH); 1955 Karl Bransik leg. (1 in FMNH); Pasco Dept., Villa Rica, 5 km SSW Villa Rica Rd., Rio de la Sal 1850 m, 10o45´6´´S 75o18´12´´W, 15– 17.x.1999, R. Brooks leg. FIT (1 in SEMC); Pasco Dept., Villa Rica Rd., 1475 m, 10o47´5´´S 75o18´54´´W, 15–18.x.1999, R. Brooks & D. Brzoska leg. FIT (1 in SEMC); Tingo Maria Huan, 20.v.1947, 2200 ft [730 m], J.C. Pallister leg. (2 in AMNH); Villcanota (1 in FMNH).

4.4. Nordus amazonicus (Sharp, 1876) (Figs. 13, 23, 61, 70, 126–129) Brachydirus amazonicus Sharp, 1876: 113. Nordus amazonicus (Sharp); HERMAN (2001: 3579). Description. L e n g t h . 9.5–11 mm. C o l o r a t i o n . Head golden-orange with anterior 1/3 of epicranium shining black; mandibles and labrum dark brown to black; antennomeres 1–7 yellow, 8–10 dark brown to black, 11 black basally and yellow distally. Pronotum, scutellum, and elytra golden-orange. Abdomen black except segments I, II, VIII and posterior 1/3 of VII goldenorange. Legs golden-orange. Very few individuals (both males and females) without shining black anterior part of head, instead head completely golden-orange.

10

H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium moderately dense, about 4.9 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 2.7 punctures / 0.3 mm, but punctures almost confluent. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.03), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 23): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 3rd longer than 2nd, 5th subequal in length to 4th, 1st–6th longer than greatest width, 7th–9th subquadrate; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 0.95; anterolateral angles not prominent, subcariniform; lateral margins strongly concave in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in at least 2–3 dense, contiguous longitudinal rows, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayish-yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with mediumsized punctures, about 3.6 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; V–VII extremely punctuated; IV–V and VIII covered with yellow microsetae, VI with yellow ones laterally and black ones medially, VII with black ones. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–IV and VIII with yellow microsetae. Segments V–VII with black and yellow iridescent microsetae throughout. Cerci straight, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (Fig. 61) with a small lobe extending across midline, with series of black prominent setae at its base; sternum VIII (Fig. 70) not with evenly curved ventral surface but rather elevated along middle, forming a carina, distal end of carina continuing into a single, narrow, discrete median lobe upon the hind margin, lobe terminating into flat hook like process; sternum IX (identical to that in Fig. 105) with two small apical processes and a shallow emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a slightly pointed extension (identical to that in Fig. 115). M a l e g e n i t a l i a . As in Figs. 126–129. Paramere in dorsal outline almost parallel-sided from middle to subapical region, then forming a rounded tip, apically subequal in length to median lobe and narrower; slightly dorsally convex in lateral outline. Sensory spinules (very few) arranged in a regular band and restricted to tip. Median lobe with


truncate-emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Among golden-orange species N. amazonicus and N. batesi can by identified by the combination of the following characters: antennomeres 8–10 and posterior half of 11 dark brown to black; epicranium with very dense punctuation anteriorly; punctuation on pronotum along medial impunctate area not dense; lateral margins of pronotum strongly concave; and in males sternum VIII forming a single elongate lobe at hind margin, without any median emargination or cleft. N. amazonicus can be distinguished from N. batesi by the following characters: antennomeres 4–7 longer than wide (4–6 subquadrate, 7 wider than long in N. batesi); antennomere 8 subquadrate (wider than long in N. batesi); antennomere 5 longer than 6 (subequal in N. batesi); and by the differences in sternum VIII (Figs. 70, 74) and the aedeagi (Figs. 126–129, 142–145). Distribution. Known from elevations of 200–300 m in Brazil, Bolivia, Colombia, Ecuador and Peru (Fig. 19). Habitat. Unknown. Remarks. Collected in flight intercept and malaise traps. Material. Holotype, „ with labels ‘Type’, ‘Ega’, [upside down] ‘Sharp Coll. 1905-313’, ‘S. America Brazil’, ‘Brachydirus amazonicus „ Type D. S.’ (in BMNH). SHARP (1876) states in the original description that that he had available only a single specimen, which therefore is the holotype. – BOLIVIA: Cochabamba, 67.5 km NE Est. Biol. Valle de Sajita, of Univ. de San Simon, 300 m, 17o6´33´´S 64o47´52´´W, 7–9.ii.1999, F. Genier leg. FIT (1 in SEMC). COLOMBIA: Putomayo Santa Rosa Kofan Indian village, 2– 24.x.1970, B. Malkin leg. (1in FMNH). ECUADOR: Napo, Limoncocha, 5.vi.1977, M.E. Steine leg. (1 in USNM); Napo, Limoncocha, 250 m, 9–16.iii.1976, J.M. Campell leg. (1 in CNCI); Napo, Tiputini Research station, 220 m, 5–25.ix.2000, D.J. Inward & K.A. Jackson leg. (1 in BMNH); Napo, Yuturi Lodge, Rio Napo, 270 m, 0o32´45´´S 76o2´18´´W, 20– 21.v.1999, R. Brooks & D. Brzoska leg. FIT (1 in SEMC); Pastaza Prov., Ashuara Indian village on Rio Macuma nr. Rio Morona, B. Malkin leg. (1 in FMNH); Sucumbios, Sacha Lodge, 270 m, 0o28´14´´S 76o27´35´´W, 21–24.v.1999, R. Brooks leg. FIT (10 in SEMC); Sucumbios, Sacha Lodge, 270 m, 0.5oS 76oW, 24.iii.–3.vi.1994, Hibbs leg. malaise trap (5 in SEMC). PERU: Monsou Valley Tingo Maria, 2.xii.1954, E.I. Schlinger & E.S. Ross leg. (1 in CASC); Loreto Dept., 1.5 km N Teniente Lopez, 2o35´S 76o06´W, 18.vii.1993, 210– 240 m, R. Leschen leg. FIT (3 in SEMC); Tambopata Prov., Madre de Dios Dpto., 15 km NE Puerto Cusco Amazonica, 12o33´S 69o03´W, 200 m, Maldonado Reserva, 13.vi.1989, J.S. Ashe & R.A. Leschen leg. FIT (1 in SEMC).

4.5. Nordus amphivolos sp.n. (Figs. 24, 71, 130–133) Description. L e n g t h . 8.5–10 mm. C o l o r a t i o n . Head golden-orange with anterior 2/3 of epicranium shining black; mandibles and labrum light brown; antennomeres light brown to black. Pronotum, scutellum, and elytra golden-orange. Abdomen black except segments I, II, VIII and posterior 1/3 of VII golden-orange. Legs yellow, darker orange in few specimens. H e a d . Microsetae brown. Anteclypeus very small. Punctures in anterior half of epicranium moderately dense, about 4.9 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 4.0 punctures / 0.3 mm, punctures separated by a distance as wide as 0.5 punctures. Length of eyes ca. 0.8ⴛ length of head, distance between eyes ca. 1ⴛ length of eye. Middle of epicranium strongly convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow

11


near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 24): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th subequal in length to 4th, 1st–6th longer than greatest width, 7th subquadrate, width of distal margin of 9th and 10th at least 1.5ⴛ length; 7th–11th densely covered by microtrichae, 6th only with very few microtrichae, 4th– 10th with single circumferential row of prominent macrosetae near middle, 1st–3rd with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.03; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial im-punctate area broadest anteriorly (width 1.5–2 punctures); punctures flanking this area in at least 3 dense, contiguous longitudinal rows, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with brown microsetae. Mete-pimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow setae and punctures covering entire surface. Elytra not depressed near scutellum, with large punctures, about 3.1 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; V–VII extremely punctuated, with slightly less punctures medially; V–VII with yellow microsetae laterally and brown iridescent ones medially. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; III–IV and VIII with yellow microsetae. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII with broad and shallow emargination medially (identical to that in Fig. 63); sternum VIII (Fig. 71) with evenly curved ventral surface, hind margin with broad and moderately deep U-shaped emargination medially; sternum IX (identical to that in Fig. 110) with two moderate apical processes and a moderate U-shaped emargination. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 130–133. Paramere in dorsal outline very slender, almost parallel-sided from middle to subapical region, then forming a sub-acute tip, apically subequal in length to median lobe and narrower; slightly dorsally convex in lateral outline. Sensory spinules (many) arranged in a pair of regular bands and restricted to tip. Median lobe with broad subacute apex in dorsal outline, slightly upcurved apically in lateral outline, forming apically two widely separated, short dorsal lobes but no dorsomedial carina basal to these. Diagnosis. N. amphivolos is very similar to N. longipes and N. stomachoponos. Only these species are golden-orange with the anterior 3/4 of the head shining black and have antennomeres 2–10 light brown to black. N. stomachoponos can be distinguished from the other two by the very shallow and sparse punctuation on the pronotum, the very dense punctuation on the anterior part of the head, and

antennomere 7 being wider than long (subquadrate in the other two species). In N. longipes antennomeres 5 and 6 are quadrate, while in N. amphivolos they are longer than wide; in N. longipes the last antennomere is brown-black while in N. amphivolos it is pale yellow. Distribution. Known from an elevation of 150 m in Brazil and Venezuela (Fig. 21). Habitat. Unknown. Derivatio nominis. The specific epithet is derived from the Greek word ‘␣␮␸␫␤␱␭␱␵’ (dubious) and refers to the problems discussed below. Remarks. The status of N. amphivolos is somewhat problematic since there are no males known for N. longipes, and females are slightly different (both species found in Pará, Brazil). Based on the available morphological structures (see diagnosis of N. longipes) I recognize N. amphivolos as a species separate from N. longipes pending the discovery of N. longipes males. Material. Holotype, ‚ with labels ‘Brazil, Para, Jacareacanga, xii.1968, M. Alvarenga’, HOLOTYPE Nordus amphivolos Chatzimanolis des. Chatzimanolis 2003’ (in AMNH). – Paratypes, 76 specimens: same locality data and collector (3 in AMNH, 1 in SEMC); ‘Brazil, Mato Grosso, Vila Vera, 12o46´S 55o30´W, x.1973, M. Alvarenga’ (2 in AMNH); ‘Brazil, Sinop, M. Grosso, x.1975, M. Alvarenga’ (48 in CNCI, 2 in FMNH, 5 in SEMC); same locality data and collector, x.1976 (13 in CNCI); ‘Venezuela: Amazonas Rio Mavaca Camp, 2o2Ń 65o06´W, 150 m, 16–27.iii.1989’ (2 in AMNH). All paratypes with label ‘Paratype Nordus amphivolos Chatzimanolis des. Chatzimanolis 2003’.

4.6. Nordus antennatus (Sharp, 1876) (Figs. 25,72, 134–137) Brachydirus antennatus Sharp, 1876: 110. Nordus antennatus (Sharp); HERMAN (2001: 3579). Description. L e n g t h . 8.5–10.5 mm. C o l o r a t i o n . Head black; mandibles and labrum light brown; antennomeres 1–5 dark yellow, 6–9 whitish-yellow, 10–11 brown. Pronotum, elytra, and scutellum black. Abdomen black except segments VIII and posterior 1/3 of VII golden-orange. Legs black except distal parts of proand mesofemora, -tarsi, and -tibiae orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 5.7 punctures / 0.3 mm, punctures elongate longitudinally, margins of punctures confluent; in posterior half about 3.5 punctures / 0.3 mm, distance between punctures as wide as 0.5–1 punctures. Length of eyes ca. 0.8ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 25): 1st thickened only in distal half, becoming narrower at distal end, longer than 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–5th longer than greatest width, 6th quadrate, 7th– 10th wider than greatest length; 6th–11th densely covered by micro-trichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum slightly longer than wide, width : length = 0.91; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5

12

punctures); punctures flanking this area in 1–2 sparse longidutinal rows separated by distance as wide as 0.5–1 punctures, punctures of a row not contiguous, but distance varies; large, convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayishyellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, interspaces as wide as 0.5–1 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII well punctuated (V–VI less so medially); IV with yellowish-brown microsetae medially and posterolaterally, V–VI with yellow ones on posterolateral corners and laterally and with brown ones anteromedially, without setae posteromedially, VII with brown microsetae. Paratergites IV–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–V and VIII with brownish-yellow microsetae, VI–VII with brown ones, longer black ones on posterior margins of each segment. Segments VI–VII with black iridescent setae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 72) with evenly curved ventral surface, hind margin with two strong triangular lobes (tips rounded) and a shallow, broad U-shaped emargination medially; sternum IX (identical to that in Fig. 110) with two moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a slightly pointed extension (identical to that in Fig. 115). M a l e g e n i t a l i a . As in Figs. 134–137. Paramere in dorsal outline consistently converging to a rounded tip, apically slightly shorter than median lobe and narrower; completely straight in lateral outline. Sensory spinules arranged in two very short longitudinal rows restricted to tip. Median lobe with a broad subacute apex in dorsal outline, slightly upcurved apically in lateral outline, forming a small dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus antennatus, N. banghaasi, N. picticornis, N. aeneipennis and N. zischkai can all be distinguished from other species of Nordus by the following characters: punctures on anterolateral corners of sterna VI and VII less dense than on the rest of these sterna; punctures on anterior part of epicranium longitudinally elongate; and shape of sternum VIII in males (shape varies among these species, but overall is distinctive for this group of species). N. antennatus and N. banghaasi can be distinguished from N. picticornis, N. aeneipennis, and N. zischkai by antennomere 4 being longer than wide (subquadrate in other species). N. antennatus and N. banghaasi differ in the following characters: head, pronotum and elytra dark brown to black in N. antennatus and golden-orange in N. banghaasi; the small difference in the shape of sternum VIII in males (Figs. 72, 73); and the coloration of the antennae: in N. antennatus antennomeres 1–5 are dark yellow, 6–9 are whitish-yellow


and 10–11 are brown, while in N. banghaasi antennomeres 1–6 are yellow and 7–11 are whitish-yellow. Distribution. Known from elevations of 150–2000 m in Bolivia, Brazil, Colombia, Ecuador and Peru (Fig. 19). Habitat. Unknown. Parasites. FRANK (1982) mentions the occurrence of Mimeomyces brachyderi (Laboulbeniaceae) on N. antennatus as previously documented by THAXTER (1900, 1908, 1931). Remarks. Collected in feces traps, flight intercept traps, malaise traps and from fungusy logs. Material. Lectotype, ‚, here designated, with labels ‘Type’, ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905313’, ‘Brachydirus antennatus ‚ Type Amazons D. S.’, ‘Lectotype Brachydirus antennatus Sharp des. Chatzimanolis 2002’ (in BMNH). – Paralectotypes, 3 specimens: 1„ with labels ‘Type’, ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus antennatus „ Type D. S.’, [upside down] ‘No 1155 examined by Prof. Thatzer for Laboulbeniaceae’, ‘Paralectotype Brachydirus antennatus Sharp des. Chatzimanolis 2002’ (in BMNH). 1 specimen with labels ‘Ega’, ‘Bowring 6347*’, ‘antennatus Shr.’, ‘det. Bernhauer’, ‘Paralectotype Brachydirus antennatus Sharp des. Chatzimanolis 2002’ (in BMNH). 1‚ with labels ‘Amazons’, ‘S. America Brazil’, ‘Brachydirus antennatus 2nd Type Amazons D. S.’, ‘Sharp Coll. by exchange with the British Museum’, ‘Paralectotype Brachydirus antennatus Sharp des. Chatzimanolis 2002’ (in FMNH). – BOLIVIA: Coroico (1 in DEI, 3 in FMNH, 1 in NMW, 1 in AMNH). BRAZIL: Amazons (1 in FMNH). COLOMBIA: Amazones PNN, Amacayaco San Martin, 3o23´S 70o6´W 150 m, 8–16.ix.2000, B. Amado leg. malaise trap (1 in SEMC, 1 in UNCB); 3 km W Villavicencio Meta, 920 m, 11.iii.1955 (4 in CASC); F. W. Furry, 71 Villavicencio, 1928, L. Pena leg. (1 in FMNH); Muzo, 750 m (4 in FMNH); Sogamoso Ost-Cordillieren, i.1902, 900 m (2 in FMNH, 1 in AMNH); Sogamoso ost., 900 m, i.1902, ex. coll. Klima (10 in NMW); Ost-Villavieja, 450 m, ii.1911 (1 in NMW); Villavicencio, 450 m, v.1911 (1in FMNH); Fassl leg. (2 in NMW). ECUADOR: Napo, 12 km SW Tena, 8.vii.1976, S. & J. Peck, ex. suspended carrion trap (1 in CNCI); Napo, El Chaco, 2000 m, ii.1983, M. Sharkey & L. Masner leg. malaise trap (1 in CNCI); Napo, Misahualli, 350 m, ii.1983, M. Sharkey (1 in CNCI); Napo, Yasuni, N.P. Yasuni research station, 215 m, lowland forest, Ratcliffe, Jameson & Smith leg. FIT (1 in UNSM); Napo, 1200 m, 25.4 km Sarayacu-Loreto Rd., 12.vii.1994, F. Genier leg. feces trap (1 in SEMC); Napo, 950 m, 25.3 km Sarayacu-Loreto Rd., 14.vii.1994, F. Genier leg. feces trap (1 in SEMC); Pastaza, 1100 m, 1 km E Mera, 13–17.vii.1976, S. Peck leg. forest carrion trap 45 (1 in CNCI); Pastaza, 54.5 km S Puyo, PuyoMacas, R. David, 927 m, 20.ix.1993, D. Brzoska leg. (1 in SEMC); Santa Jnez, R. Haensch S. leg. (1 in FMNH); Sucumbios, Sacha Lodge, 0.5oS 76.5oW, 270 m, 1–31.xii.1994, Hibbs leg. malaise trap (1 in SEMC); Pompeya, v.1965 (1 in FMNH). PERU: Callanga (2 in DEI, 3 in NMW); Madre de Dios, Pentiacolla Lodge, 8 km NW El Mirador Trail, Alto Madre de Dios river, 800 m, 12o38´30´´S 71o16´41´´W, 23– 28.x.2000, R. Brooks leg. FIT (1 in SEMC); Pachitea (1 in FMNH); Pasco Dept., Villa Rica-Puerto Bermudas Rd., 910 m, 10o34´16´´S 75o5´30´´W, 17.x.1999, R. Brooks leg. on fungusy log (1 in SEMC); Pasco Dept., Villa Rica Rd., 1475 m, 10o47´6´´S 75o18´54´´W, 15–18.x.1999, R. Brooks & D. Brzoska leg. FIT (2 in SEMC); Ucayali Dept., Tingo Maria– Pucallpa Rd., Puente Chino km 205, 1300 m, 9o8´12´´S 75o47´20´´W, 11–14.x.2000, R. Brooks leg. FIT (3 in SEMC); Valle Chanchamayo, 800 m, viii.1957, Wevauch leg. (1 in NMW); Bang-Haas leg. (1 in FMNH).

4.7. Nordus banghaasi (Bernhauer, 1905) (Figs. 26, 73, 138–141) Brachydirus banghaasi Bernhauer, 1905: 184. Nordus banghaasi (Bernhauer); HERMAN (2001: 3579). Description. L e n g t h . 8.5–10.5 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres 1–6 yellow, 7–11 whitish-yellow.

13


Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII and posterior 1/3 of VII golden-orange. Legs golden-orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 4.9 punctures / 0.3 mm, punctures elongate longitudinally, margins of punctures confluent; in posterior half about 3.6 punctures / 0.3 mm, distance between punctures as wide as 0.5 puncture. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 26): 1st thickened only in distal half, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–4th longer than greatest width, 5th quadrate, 6th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 7th–11th and distal margin of 6th densely covered with microtrichae, 4th–10th with single circumferential row of prominent macrosetae near middle, 1st–3rd with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 0.94; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–2 punctures); punctures flanking this area in 1–2 sparse longitudinal rows separated by distance as wide as 0.5–1 punctures, punctures of a row not contiguous, but distance varies; large, convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayish-yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, and interspaces as wide as 0.5–1 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII well punctuated (V–VI less so medially); IV with yellow microsetae medially and posterolaterally, V with yellow ones throughout, VI with yellow ones laterally and brown ones medially, VII with brown ones. Paratergites IV–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: IV–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–VI and VIII with yellow microsetae, VII with brown ones. Segments VI–VII with black iridescent setae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 73) with evenly curved ventral surface, hind margin with two strong triangular lobes (tips rounded) and a shallow to moderately broad U-shaped emargination medially; sternum IX (identical to that in Fig. 110) with two moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures except margin of sternum VIII forming a slightly pointed extension (identical to that in Fig. 115).

M a l e g e n i t a l i a . As in Figs. 138–141. Paramere in dorsal outline consistently converging to a rounded tip, apically slightly shorter than median lobe and narrower; completely straight in lateral outline. Sensory spinules (many) arranged in a pair of bands and restricted to tip. Median lobe with subacute apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus antennatus, N. banghaasi, N. picticornis, N. aeneipennis and N. zischkai can all be distinguished from other species of Nordus by the following characters: punctures on anterolateral corners of sterna VI and VII less dense than on the rest of these sterna; punctures on anterior part of epicranium longitudinally elongate; and shape of sternum VIII in males (shape varies among these species, but overall is distinctive for this group of species). N. antennatus and N. banghaasi can be distinguished from N. picticornis, N. aeneipennis, and N. zischkai by antennomere 4 being longer than wide (subquadrate in other species). N. antennatus and N. banghaasi differ in the following characters: head, pronotum and elytra dark brown to black in N. antennatus and golden-orange in N. banghaasi; the small difference in the shape of sternum VIII in males (Figs. 72, 73); and the coloration of the antennae: in N. antennatus antennomeres 1–5 are dark yellow, 6–9 are whitish-yellow and 10–11 are brown, while in N. banghaasi antennomeres 1–6 are yellow and 7–11 are whitish-yellow. Distribution. Known from elevations of 730–1040 m in Bolivia (Fig. 19). Habitat. Unknown. Remarks. Collected in flight intercept traps. Material. Lectotype, ‚, here designated, with labels ‘Mapiri Bolivia’, ‘Bang-Haasi Bernh. Typus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Lectotype Brachydirus banghaasi Bernhauer des. Chatzimanolis 2001’ (in FMNH). – Paralectotypes: 1 specimen with labels ‘Bolivien’, ‘pendasoma für Bang Haas determ. Bernh.’, ‘Bang-Haasi Bernh. Typus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Paralectotype Brachydirus banghaasi Bernhauer des. Chatzimanolis 2001’ (in FMNH). – BOLIVIA: Cochabamba Prov., Cochabamba, 117 km E Yungas, Cochabamba-Villa Tunari Rd., 1040 m, 17o6´32´´S 65o41´12´´W, 1–12.ii.1999, R. Hanley leg. FIT (37 in SEMC); Cochabamba Prov., Cochabamba, 117 km E Yungas, Cochabamba–Villa Tunari Rd., 730 m, 17o3´54´´S 65o38´43´´W, 1–12.ii.1999, R. Hanley leg. FIT (5 in SEMC); Cochabamba Prov., Cochabamba, 117 km E Yungas, Lagunitas, 1000 m, 17o6´22´´S 65o40´57´´W, 1– 12.ii.1999, F. Genier leg. FIT (11 in SEMC); Coroico, BangHaas leg. (1 in AMNH, 1 in BMNH, 1 in DEI, 3 in FMNH, 1 in NMW); 1955, Karl Bransik leg. (1 in FMNH).

4.8. Nordus batesi (Sharp, 1876) (Figs. 27, 74, 113, 142–145) Brachydirus batesi Sharp, 1876: 114. Nordus batesi (Sharp); HERMAN (2001: 3579). Description. L e n g t h . 8.0–10 mm. C o l o r a t i o n . Head golden-orange with anterior 1/3 of epicranium shining black; mandibles and labrum dark brown; antennomeres 1–4 whitish-yellow, 7–11 black, 5–6 in some specimens whitish-yellow, in others black. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII and posterior 1/3 of VII golden-orange. Legs whitish-yellow. H e a d . Microsetae yellow on anterior 3/4 and black on posterior 1/4. Anteclypeus very small. Punctures in anterior half of epicranium moderately dense, about 5.3 punctures / 0.3 mm, not elongate longitudinally, margins of punctures

14

confluent, in females punctures less dense than in males; in posterior part about 3.1 punctures / 0.3 mm, but punctures almost confluent. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.06), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 27): 1st thickened only in distal half, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd longer than 3rd, 5th subequal in length to 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate, 6th quadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 0.94; anterolateral angles not prominent, subcariniform; lateral margins strongly concave in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in at least 1–2 dense, contiguous longitudinal rows, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayish-yellow setae and punctures covering entire surface. Elytra not depressed near scutellum, with large punctures, about 4.5 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae, except profemur with row of widely scattered long black macrosetae on anterior edge. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with yellow microsetae on medial disc; V–VII extremely punctuated (V somewhat less densely punctuated medially); V with yellow microsetae medially and along posterolateral angles, VI with yellow microsetae throughout, VII with yellow microsetae laterally and black ones medially. Paratergites III–VII covered with whitishyellow microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III– VII densely punctuated otherwise; III–IV and VIII with yellow microsetae. Segment V–VII with black and yellow iridescent microsetae. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 74) with evenly curved ventral surface, hind margin with single broad, triangular lobe rounded apically, no median cleft or emargination; sternum IX (identical to that in Fig. 105) with two small to moderate apical processes and a shallow U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a moderate subacute extension (Fig. 113). M a l e g e n i t a l i a . As in Figs. 142–145. Paramere in dorsal outline very wide, almost parallel-sided but a bit convex from middle to subapical region, then forming a broad subacute tip, apically slightly longer than median lobe and narrower; fairly straight in lateral outline but with a slight dorsal bend 2/3 from base. Sensory spinules arranged in a pair of


irregular bands and restricted to tip. Median lobe with broad emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming apically two widely separated, short dorsal lobes but no dorsomedial carina basal to these. Diagnosis. Among golden-orange species N. amazonicus and N. batesi can by identified by the combination of the following characters: antennomeres 8–10 and posterior half of 11 dark brown to black; epicranium with very dense punctuation anteriorly; punctuation on pronotum along medial impunctate area not dense; lateral margins of pronotum strongly concave; and in males sternum VIII forming a single elongate lobe at hind margin, without any median emargination or cleft. N. amazonicus can be distinguished from N. batesi by the following characters: antennomere 4– 7 longer than wide (4–6 subquadrate, 7 wider than long in N. batesi); antennomere 8 subquadrate (wider than long in N. batesi); antennomere 5 longer than 6 (subequal in N. batesi); and by the differences in sternum VIII (Figs. 70, 74) and the aedeagi (Figs. 126–129, 142–145). Distribution. Known from elevations of 200–400 m in Brazil, Colombia, French Guiana, and Peru (Fig. 58). There are no specimens known from Bolivia as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Habitat. Fungusy logs, Favolus sp. fungi, resupinate orange polypore fungi. Remarks. Also collected in flight intercept traps. Material. Lectotype, ‚, here designated, with labels ‘Type’, ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905313’, ‘Brachydirus batesi ‚ Type D. S.’, ‘Lectotype Brachydirus batesi Sharp des. Chatzimanolis 2002’ (in BMNH). – Paralectotypes, 3 specimens: 1„ with labels: on specimen paper card ‘Brachydirus batesi „ Type amazons D. S.’ and below ‘Type’, ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905313’, ‘Paralectotype Brachydirus batesi Sharp des. Chatzimanolis 2002’ (in BMNH). 2‚‚ with labels ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus batesi ‚ 2nd Type Amazon D. S.’, ‘Paralectotype Brachydirus batesi Sharp des. Chatzimanolis 2002’ (in BMNH). 1‚ with labels ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus batesi ‚ 2nd Type Amazon D. S.’, ‘Sharp Coll. by exchange with the British Museum’, ‘Paralectotype Brachydirus batesi Sharp des. Chatzimanolis 2002’ (in FMNH). – BRAZIL: Amazons, M. Cameron Bequest (1 in BMNH, 1 in DEI, 1 in FMNH); Ega (2 in BMNH); Sinop, M. Grosso, x.1975, M. Alvarenga (1 in CNCI). COLOMBIA: Putumayo PNN La Paya Mamansoya, Tierra Alta, 0o6´S 74o58´W, 350 m, 22–26.ix.2001, D. Campos leg. malaise trap (1 in SEMC, 1 in UNCB). FRENCH GUIANA: Saul, 8 km W, 280 m, 7.vi.1997, J.S. Ashe & R. Brooks leg. on Favolus sp. (1 in SEMC); Saul, 8 km NW, Boeuf Mort trail, 300 m, 7.vi.1997, J.S. Ashe & R. Brooks leg. on Favolus sp. (1 in SEMC); Saul, 7 km N, Les Eaux Claires, 220 m, 3o39´46´Ń 53o13´19´´W, 31.v.–3.vi.1997, J.S. Ashe & R. Brooks leg. FIT (1 in SEMC). PERU: Loreto Dept., 1.5 km N Teniente Lopez, 2o35´S 76o06´W, 240 m, 18.vii.1993, R. Leschen leg. on resupinate orange polypore (1 in SEMC); Loreto Dept., Teniente Lopez, 2o35´S 76o06´W, 210–240 m, 22.vii.1993, R. Leschen leg. FIT (1 in SEMC); Tambopata Prov., Madre de Dios Dpto., 15 km NE Puerto Cusco Amazonica, 12o33´S 69o03´W, 200 m, Maldonado Reserva, 9.vii.1989, J.S. Ashe & R.A. Leschen leg. FIT (2 in SEMC); Madre de Dios, Pentiacolla Lodge, Monk Saki Trail, Alto Madre de Dios river, 400 m, 12o39´22´´S 71o13´55´´W, 25.x.2000, R. Brooks leg. on fungus covered log (1 in SEMC).

4.9. Nordus championi (Sharp, 1884) (Figs. 28, 75, 104, 114, 146–149) Brachydirus championi Sharp, 1884: 350. Nordus championi (Sharp); HERMAN (2001: 3579). Description. L e n g t h . 10–12.0 mm. C o l o r a t i o n . Head golden-orange; mandibles and


labrum orange; antennomeres 1–5 golden-orange, 6–11 dark orange. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I–II, VIII and posterior 2/ of VII golden-orange. Legs golden-orange. 5 H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior 2/3 of epicranium dense, about 4.4 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior 1/3 about 2.7 punctures / 0.3 mm, punctures almost contiguous. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 28): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–5th longer than greatest width, 6th wider (basal margin) than greatest length, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.03; anterolateral angles extremely prominent and acutely delimited, forming angle of about 90°, cariniform; lateral margins slightly convex in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and shallow; medial impunctate area of equal width throughout (as wide as 3–4 punctures); punctures flanking this area in 2 rows separated by distance as wide as 1–1.5 punctures, punctures of a row not contiguous, but distance varies; large, raised, convex impunctate area in anterolateral quarters, and moderate impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, and interspaces as wide as 1 puncture, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV strongly punctuated on medial disc; V–VI strongly punctuated except medially; VII uniformly punctuated. IV–VI covered with yellow microsetae laterally, VII covered with brown ones. Paratergites III–VI covered with whitish-yellow microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise, with dark brown microsetae medially and yellow ones laterally. Segment VII with black iridescent setae in anterolateral quarters. Cerci slightly curved, basal half dark brown and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 63) with broad and shallow emargination medially; sternum VIII (Fig. 75) with evenly curved ventral surface, hind margin with broad and deep U-shaped emargination medially; sternum IX (Fig. 104) with two small to moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (Fig. 114).

15

M a l e g e n i t a l i a . As in Figs. 146–149. Paramere in dorsal outline consistently and strongly converging to a narrowly rounded tip, apically subequal in length to median lobe and narrower; slightly dorsally convex near apex in lateral outline. Sensory spinules arranged in a single cluster and restricted to tip. Median lobe with subtruncate apex in dorsal outline, abruptly bent upwards apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus facialis and N. championi can be distinguished from all other golden-orange species of Nordus by the combination of the following characters: epicranium convex; punctuation on anterior part of epicranium dense (but not very dense as in N. fungicola); and scutellum punctuated only in its posterior half. N. championi and N. facialis can be distinguished by the following characters: punctures on pronotum shallow in N. championi and deep in N. facialis; punctures on pronotum near medial impunctate area dense in N. facialis and less dense in N. championi; and antennomere 4 longer than wide in N. championi and subquadrate in N. facialis. Distribution. Known from elevations of 30–1371 m in Costa Rica, El Salvador, Guatemala, Mexico, and Nicaragua (Fig. 20). Habitat. Unknown. Remarks. Collected in flight intercept traps, fig fall-leaf litter and on leaves. Material. Lectotype, ‚, here designated, glued on a paper card, with labels: on paper card mount in D. Sharp’s handwriting ‘‚, Type, D. S. Brachydirus championi, Zapote, Guatemala, Champion’, below ‘Type’, ‘Zapote, Guatemala, Champion’, ‘B. C. A. Col. I. 2, Brachydirus championi Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Lectotype Brachydirus championi Sharp des. Chatzimanolis 2002’ (in BMNH). – Paralectotypes, 11 specimens: 3‚‚, on the same paper card in D. Sharp’s handwriting ‘Brachydirus championi, D. S. San Isidro, Guatemala 1600 ft Champion’, below ‘San Isidro, Guatemala, Champion’, ‘B. C. A. Col. I. 2, Brachydirus championi Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus championi Sharp des. Chatzimanolis 2002’ (in BMNH). 1‚ with labels: on paper card mount in D. Sharp’s handwriting ‘‚, D. S. Brachydirus championi, San Isidro, 1600 ft, Champion’, below ‘San Isidro, 1600 ft Champion’, ‘B. C. A. Col. I. 2, Brachydirus championi Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus championi Sharp des. Chatzimanolis 2002’ (in BMNH). 1„ with labels ‘Zapote, Guatemala, Champion’, ‘B. C. A. Col. I. 2, Brachydirus championi Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus championi Sharp des. Chatzimanolis 2002’ (in BMNH). 2„„ on the same card with labels ‘Zapote, Guatemala, Champion’, ‘B. C. A. Col. I. 2, Brachydirus championi Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus championi Sharp des. Chatzimanolis 2002’ (in BMNH). 2‚‚ on the same card with labels ‘Zapote, Guatemala, Champion’, ‘B. C. A. Col. I. 2, Brachydirus championi Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus championi Sharp des. Chatzimanolis 2002’ (in BMNH). 1„ with labels ‘Zapote, Guatemala, Champion’, ‘B. C. A. Col. I. 2, Brachydirus championi Sharp’, ‘Sharp Coll. by exchange with the British Museum’, ‘Paralectotype Brachydirus championi Sharp des. Chatzimanolis 2002’ (in FMNH). 1‚ with labels: on paper card mount in D. Sharp’s handwriting ‘‚, D. S. Brachydirus championi, Zapote, Guatemala, Champion’, below ‘Zapote, Guatemala, Champion’, ‘B. C. A. Col. I. 2, Brachydirus championi Sharp’, ‘Sharp Coll. by exchange with the British Museum’, ‘Paralectotype Brachydirus championi Sharp des. Chatzimanolis 2002’ (in FMNH). – COSTA RICA: Prov. Guanacaste, Cerro El Hacha, 300 m, 12 km SE de LaCruz,

16

malaise trap (3 in INBC); Prov. Guan., Est. Lomas Barbudal 30 m, vii.1991, A.C.T.U. Chavarria leg. malaise trap (2 in INBC); Prov. Guan, Est. Maritza 600 m Lado oeste del Volcan Orosi, II curso parataxonomos Ago 1990 (1 in INBC); Prov. Guan., Playas Nosara Asent, Financiera A.C. Guanacaste, 27– 30.x.1993, R. Vargas & D. Brenes leg. malaise trap (1 in INBC); Guanacaste Prov., Santa Rosa National Park, 13– 15.vii.1980, D.H. Jansen & W. Hallwachs leg. (1 in INBC); Guanacaste Prov., SSW side Cerro El Hacha, 300–400 m, viii– xi.1986, D.H. Jansen & W. Hallwachs leg. malaise trap (1 in INBC); Guanacaste Prov., Los Almenandros, P.N. Guanacasta, 1–22.vii.1992, E. Lopez leg. malaise trap (9 in INBC, 2 in SEMC); Guanacaste Prov., Maritza Biol. Sta., 650 m, 22.v.1993, J. & A. Ashe leg. FIT (1 in SEMC); Guanacaste Prov., Hacienda Comelco 24 km NW Canas Inter-Am. H’way, 50 m, 5.viii.1971, R. Heithaus leg. (4 in CASC); Barranca nr. Puntarenas, 25.viii.1954, E. Ross. leg. (1 in CASC); Las Canas Pac. Slope, 13.vi.1923, A. Smith leg. (1 in USNM). EL SALVADOR: Quezaltepeque, 500 m, 19.v.1963, D.Q. Cavagnaro & M.E. Irwin leg. (2 in CASC); San Salvador, 23.vii.1959 (2 in USNM); San Salvador, 13–14.vi.1958, O.L. Cartwright leg. (1 in USNM); Izalco Salv., Fred Knab leg. (1 in USNM). MEXICO: Chiapas, Municipio de Angel Albino Corzo (Jaltenango), above Finca Custepec, 1371 m, 11.xii.1981, D.E. & P.M. Breedlove leg. (2 in CASC); Chiapas, 96 km S Tuxtla Gutierrez on road to Nueva Concordia nr. junction with road to Mapastepec, 732 m, D.E. & J.A. Breedlove leg. (1 in CASC); Chis. 20–25 mi N Huixtla, xi.1978, E. Welling leg. malaise trap (1 in CNCI); Chis. 20–25 mi N. Huixtla, 3000 ft [1000 m], 2.vi.1969, B.V. Peterson leg. (1 in CNCI); Chis. 17 mi N. Huixtla, 2500 ft [833 m], 3.vi.1969, W.R.M. Mason leg. (1 in CNCI); Chis. 20 mi N Huixtla, 3000 ft [1000 m], 3.vi.1969, W.R.M. Mason leg. (4 in CNCI); Chiapas 1 mi SW Rizo de Oro, 2700 ft [900 m], 19–26.viii.1971, A. Newton leg. human dung trap (1 in FMNH); Chiapas 9 mi N Arriaga, 2000 ft [667 m], 19–26.viii.1971, A. Newton leg. human dung trap (1 in FMNH); Chiapas 25 mi SSW Vincente Guerrere, 9.vi.1987, B. Ratchliffe & M. Jameson leg. (1 in UNSM); Chiapas, La Victoria, 15.vii.1982, L. Delgado leg. (12 in CZUG); Chiapas, 19 km E Santa Isabel Rio Zoy Altenco, 9.vi.1991, 650 m, J.S. Ashe leg. fig fall-leaf litter (2 in SEMC). NICARAGUA: Granada Dept., Res. Nat. Volcan Mombacho, entrance Rd., 1060 m, 11o50.05Ń 85o58.83´W, 1–5.vi.2002, R. Brooks, S. Chatzimanolis & Z. Falin leg. FIT (1 in SEMC).

4.10. Nordus cherylae sp.n. (Figs. 29, 76, 150–153) Description. L e n g t h . 8.5–10.0 mm. C o l o r a t i o n . Head black; mandibles and labrum orange; antennomeres 1–6 brown, 7–11 brownish-yellow. Pronotum, elytra, and scutellum black. Abdomen black except: segments I–II light brown; narrow bands along posterior margins of sterna III–VI, posterior 1/3 of segment VII, and segment VIII golden-orange. Legs golden-orange except metatibiae and -tarsi black. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 5.3 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior half about 1.7 punctures / 0.3 mm, distance between punctures as wide as 0.5–1 puncture, middle of posterior half without punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1– 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 29): 1st thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th subequal in length to 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate, 6th–10th wider than greatest length, width of distal margin of 9th


and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum longer than wide, width : length = 0.85; anterolateral angles not prominent, subcariniform; lateral margins strongly concave in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures mediumsized and shallow; medial impunctate area broadest anteriorly (width 2–3 punctures); punctures flanking this area in 1–2 sparse longitudinal rows separated by distance as wide as 1–1.5 punctures, punctures of a row not contiguous, but distance varies; large, slightly convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming 2 rows, not covering entire width. Scutellum without arch-like carina, with grayish-yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, and interspaces as wide as 1–1.5 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V with subbasal carina; IV–VII well punctuated; IV–V with yellowish-brown microsetae medially and posterolaterally, VI with yellow ones laterally and brown ones medially, VII with brown ones. Paratergites IV–VII covered with whitishyellow and black microsetae on posterior half. Sterna IV– VII on anterolateral corners without punctures or with sparse punctures; sterna III–VII densely punctuated otherwise (IV–VI slightly less postero-medially); III–VI and VIII with brownish-yellow microsetae; VII with brown microsetae. Segments V–VII with black iridescent setae throughout. Cerci thin and straight, basal half light brown and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 76) with evenly curved ventral surface, hind margin with two moderate triangular lobes and a broad and deep U-shaped emargination; sternum IX (identical to that in Fig. 111) with no apparent apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII sinuate (identical to that in Fig. 118). M a l e g e n i t a l i a . As in Figs. 150–153. Paramere in dorsal outline very wide, widened apically, with broad subtruncate tip bearing a small, pointed medial projection, apically subequal in length to median lobe and slightly wider; dorsally slightly concave in lateral outline. Sensory spinules (many) arranged in a pair of loose clusters restricted to tip. Median lobe with subacute apex in dorsal outline, slightly upcurved apically in lateral outline, forming apically two widely separated, dorsal lobes (their bases at an angle of 90° to lateral margin of median lobe) but no dorsomedial carina basal to these. Diagnosis. Nordus cherylae, N. elegans, N. portokalis, and N. speciosus can be distinguished from all other species of Nordus by the presence of a subbasal carina (in addition to the basal one) on abdominal tergum V. N. portokalis can be distinguished from the other three species by the presence of the very distinctive orange abdomen (black in the other species) and the subquadrate pronotum (pronotum longer


than wide in other species). N. speciosus can be distinguished from N. cherylae and N. elegans by the metallic green head, pronotum and elytra. In N. cherylae head, pro-notum and elytra are all black, whereas in N. elegans the head is dark brown, the pronotum golden-orange, and the elytra dark orange to brown; sternum VIII in females is evenly rounded posteriorly in N. elegans (Fig. 114), and the aedeagus is as in Figs. 170–173, while in N. cherylae sternum VIII in females is sinuate (Fig. 118) and the aedeagus is as in Figs. 150–153. Distribution. Known from elevations of 1480–1800 m in Bolivia (Fig. 21). Habitat. Unknown. Derivatio nominis. The specific epithet is in honor of Cheryl A. Murphy. Remarks. Collected in flight intercept traps. Material. Holotype, ‚ with labels ‘Bolivia, Cochabamba,

Cochabamba, 105 km E Yungas nr. Rio Carmen Mayu (Cochabamba–Villa Tunari Rd.), 1800 m, 17o8´47´´S 65o43´55´´W, 8–12.ii.1999, F. Genier, BOL1G99 058 ex: FIT’, barcode label SM0173911, ‘HOLOTYPE Nordus cherylae Chatzimanolis des. Chatzimanolis 2003’ (in SEMC). – Paratypes, 24: same locality data and collector, barcode labels SM0173916, SM0173914, SM0173907, SM0173915, SM0173921, SM0173919, SM0173920, SM0173912 (7 in SEMC, 1 in FMNH); same locality data and collector but 1750 m, barcode labels SM0173551, SM0173556, SM0173554, SM0173557, SM0173552, SM0173549, SM0173550, SM0173555, SM0173548 (8 in SEMC, 1 in FMNH); same locality data but 1750 m, R.S. Hanley leg., barcode labels SM0161836, SM0162112, SM0161843, SM0161838 (4 in SEMC); ‘Cochabamba, Cochabamba, 109 km E Yungas (Cochabamba–Villa Tunari Rd.) 1480 m, 17o8´50´´S 65o42´29´´W, 1–6.ii.1999, F. Genier, BOL1G99 027 ex: FIT’, barcode labels SM0173974, SM0178973 (2 in SEMC); same locality data, R.S. Hanley leg., barcode label SM0162006 (1 in SEMC). All paratypes with label ‘Paratype Nordus cherylae Chatzimanolis des. Chatzimanolis 2003’.

4.11. Nordus cribricollis (Sharp, 1876) (Figs. 15, 30, 77, 154–157) Brachydirus cribricollis Sharp, 1876: 112. Nordus cribricollis (Sharp); HERMAN (2001: 3579). Description. L e n g t h . 8.0–10 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres 1–6 yellow, 7–11 darker yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/ of VII golden-orange. Legs golden-orange. 3 H e a d . Microsetae yellow on anterior 3/4 and black on posterior 1/4. Anteclypeus very small. Punctures in anterior 3/ of epicranium dense, about 6.2 punctures / 0.3 mm, not 4 elongate longitudinally, margins of punctures confluent; in posterior part about 3.1 punctures / 0.3 mm, punctures contiguous. Length of eyes ca. 0.8ⴛ length of head, distance between eyes less than 1.5ⴛ length of eye. Middle of epicranium convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.08), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 30): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–4th longer than greatest width, 6th much wider (basal margin) than greatest length and shorter than 5th, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with

17

single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum (Fig. 15) quadrate, width : length = 1.02; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 2–3 punctures); punctures flanking this area in at least 3 dense, contiguous longitudinal rows, punctures of a row conti-guous; large, convex, raised impunctate area in anterolateral quarter, and small impunctate area along posterior and posterolateral margins. Pronotum with dark brown microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayishyellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with large punctures, about 4.0 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; V densely punctuated on medial disc but less dense medially on posterior margin; VI densely punctuated except medially; IV with grayish-yellow microsetae on medial disc, V–VI with grayish-yellow ones except medially, VII with dark brown ones, VIII with sparsely distributed yellow ones. Paratergites III–VII covered with whitish-yellow setae on posterior half. Sterna: III–VII densely punctuated; III–VI and VIII with yellow microsetae. Segment VII with black iridescent setae throughout. Cerci slightly curved, basal half light brown and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 63) with broad and shallow emargination medially; sternum VIII (Fig. 77) with evenly curved ventral surface, hind margin with broad moderate U-shaped emargination medially; sternum IX (identical to that in Fig. 104) with two small to moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 154–157. Paramere in dorsal outline very slender, including base, almost parallel-sided from middle to subapical region, then forming a narrowly rounded tip, apically subequal in length to median lobe and narrower; slightly dorsally convex in lateral outline. Sensory spinules (many) arranged in a pair of bands and restricted to tip. Median lobe with broad subacute apex in dorsal outline, slightly upcurved apically in lateral outline, forming apically two well separated, small dorsal lobes but no dorsomedial carina basal to these. Diagnosis. Nordus cribricollis, N. elytisi and N. seferisi can be distinguished from all other golden-orange Nordus species by antennomere 6 being wider than long (subquadrate in all other species except N. portokalis, which by its orange abdomen differs from the three abovementioned species with black abdomen). N. cribricollis can be distinguished from N. seferisi and N. elytisi by antennomere 4 being longer than wide rather than subquadrate. In addition, N. cribricollis males have a paramere (Fig. 154) much narrower than in the other two species (Figs. 174, 218). Distribution. Known from elevations of 25–740 m in Bolivia, Brazil, Colombia, French Guiana, Guyana, Peru,

18

and Suriname (Fig. 58). There are no specimens known from Venezuela as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Habitat. Unknown. Remarks. Collected from fungusy logs, carrion traps, and flight intercept traps. Although SHARP (1876) mentioned both Ega and St. Paulo as the type localities, all specimens examined from the type series are from Ega. Material. Lectotype, ‚, here designated, with labels ‘Type’, ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905313’, ‘Brachydirus cribricollis ‚ Type D. S.’, ‘Lectotype Brachydirus cribricollis Sharp des. Chatzimanolis 2002’ (in BMNH). – Paralectotypes, 6 specimens: 1„ with labels: on specimen paper card ‘Brachydirus cribricollis amazons „ Type D. S.’ and below ‘Type’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Paralectotype Brachydirus cribricollis Sharp des. Chatzimanolis 2002’ (in BMNH). 3‚‚ with labels ‘Ega’, ‘S. America Brazil’, ‘Brachydirus cribricollis 2nd Type D. S. Amazon’, [upside down] ‘Sharp Coll. 1905313’, ‘Paralectotype Brachydirus cribricollis Sharp des. Chatzimanolis 2002’ (in BMNH). 1 specimen with labels ‘Ega’, ‘cribricollis Shr’, ‘Bowring 6347*’, ‘det. Bernhauer’, ‘Paralectotype Brachydirus cribricollis Sharp des. Chatzimanolis 2002’ (in BMNH). 1‚ with labels ‘Ega’, ‘S. America Brazil’, ‘Brachydirus cribricollis 2nd Type D. S. Amazon’, [upside down] ‘Sharp Coll. 1905-313’, ‘Sharp Coll. by exchange with the British Museum’, ‘Paralectotype Brachydirus cribricollis Sharp des. Chatzimanolis 2002’ (in FMNH). – BOLIVIA: Cochabamba Prov., Cochabamba, 124 km E Yungas, Cochabamba-Villa Tunari Rd., 700 m, 17o3´45´´S 65o38´38´´W, 1–6.ii.1999, F. Genier leg. carrion trap (1 in SEMC). BRAZIL: Amazons, Bernhauer det. (1 in DEI); Sinop, M. Grosso, x.1975, M. Alvarenga leg. (29 in CNCI); Sinop, M. Grosso, x.1976, M. Alvarenga leg. (6 in CNCI); Sinop, Mato Grosso, x.1976, M. Alvarenga leg. (1 in CNCI); Serra do Navio Terr. Amapa, 27.ix.1957, J. Lane leg. (1 in CNCI); Mato Grosso, Barra do Tapirare, 17.i.–2.ii.1964, B. Malkin leg. sweeping forest at night (1 in FMNH); Mato Grosso,Tapirape Indian village at confluence R. Tapirape and R. Araguata, 20–30.xi.196x, B. Malkin leg. night sweeping along forest trail (1 in FMNH); Para, Aldeia Aracu, Igarape Gurupumu Maranhao, 50 km E of Caninde, 1963, B. Malkin leg. howler’s feces (2 in FMNH); Para, Redencao vic. Keyapo Territory Pinkaiti R., xi.1998, tropical evergreen (1 in FMNH); Rondonia, Fazenda Rancho grande nr. Ariquemes, 16.x.1993, D.H & A.C. Kistner leg. fish pitfall trap (1 in FMNH); Para, Baker leg. (1 in FMNH); Para, Jacareacanga, xii.1968, M. Alvarenga leg. (4 in AMNH). COLOMBIA: Villavicencio, 450 m, v.1902, Fassl leg. (1 in FMNH). FRENCH GUIANA: Roura, 27.4 km SSE, 280 m, 4o44´20´Ń 52o13´25´´W, 23– 25.v.1997, J.S. Ashe & R. Brooks leg. FIT (9 in SEMC); Roura, 13 km SSE, 240 m, 4o38´38´Ń 52o17´56´´W, 13.vi.1997, J.S. Ashe & R. Brooks leg. FIT (1 in SEMC); Roura, 8.4 km SSE, 200 m, 4o40´41´Ń 52o13´25´´W, 29.v.– 10.vi.1997, J.S. Ashe & R. Brooks leg. FIT (1 in SEMC); Saul, 7 km N, Les Eaux Claires, Mt. La Fumee, 220–500 m, 3o39´46´Ń 53o13´19´´W, 1–9.vi.1997, J.S. Ashe & R. Brooks leg. FIT (5 in SEMC); Saul, Mt. Galbao summit, 740 m, 3o37´18´Ń 53o16´42´´W, 6.vi.1997, J.S. Ashe & R. Brooks leg. on fungusy logs (3 in SEMC); Cayenne, 33.5 km S and 8.4 km NW of H’way N2 on H’way D5, 30 m, 4o48´18´Ń 52o28´41´´W, 26–28.v.1997, J.S. Ashe & R. Brooks leg. FIT (2 in SEMC). GUYANA: Region 8, Iwokrama Forest, 26 km SW Kurupukari, Iwokrama Mt. base camp, 4o20´17´Ń 58o48´38´´W, 60–600 m, 23–25.v.2001, R. Brooks & Z. Falin leg. FIT (11 in SEMC); Region 8, Iwokrama Forest, Kabocalli Field Stn., 4o17´4´Ń 58o30´35´´W, 60 m, 3–5.vi.2001, R. Brooks & Z. Falin leg. FIT (1 in SEMC); Region 8, Iwokrama Forest, Pakatau hills, 4o44´54´Ń 59o1´36´´W, 70 m, 25– 29.v.2001, R. Brooks & Z. Falin leg. FIT (2 in SEMC); Region 8, Iwokrama Forest, 1 km W Kurupukari, Iwokrama field Stn., 4o40´19´Ń 58o41´43´´W, 60 m, 26–29.v.2001, R. Brooks & Z. Falin leg. FIT (1 in SEMC); Guiana [Guyana] (1 in FMNH). PERU: Madre de Dios, Rio Tambopata Res. 30 air km SW Pto. Maldonade, 290 m, 16–20.xi.1979, J.B. Heppner leg., subtropical moist forest (1 in USNM). SURINAME: Saramacca, West Suriname Rd., 128 km WSW Zanderij Airport, 25 m, 4o59´6´Ń 56o18´48´´W, 12–14.vi.1999, Z. Falin & B. DeDjin


leg. FIT (4 in SEMC); Marowijne, Nassau Mt., 480 m, 4o48´56´Ń 54o36´20´´W, 3–4.vi.1999, Z. Falin & B. DeDjin leg. FIT (1 in SEMC); Para, Carolina Creek, 11 km SE Zanderij Airport, 30 m, 5o23´36´Ń 59o9´29´´W, 18– 20.vi.1999, Z. Falin & B. DeDjin leg. FIT (1 in SEMC); Commewijne, Akintosoela, CELOS camp, 39 km SE Suriname River bridge road to Redi Doti, 40 m, 5o16´17´Ń 54o55´15´´W, 29.vi.–3.vii.1999, Z. Falin leg. FIT (2 in SEMC).

4.12. Nordus densiventris (Bernhauer, 1917) (Figs. 31, 62, 78, 105, 158–161) Brachydirus densiventris Bernhauer, 1917: 21. Nordus densiventris (Bernhauer); HERMAN (2001: 3579). Description. L e n g t h . 8.5–10 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres all yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/3 of VII golden-orange. Legs golden-orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 5.3 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior half about 4.4 punctures / 0.3 mm, punctures almost confluent. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 2ⴛ length of eye. Middle of epicranium flat; epicranium with microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.07), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 31): 1st thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate, 6th quadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th 1.5ⴛ length; 6th–11th densely covered by microtrichae; 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.03; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in at least 1–2 sparse longitudinal rows separated by a distance as wide as 1–1.5 punctures, punctures of a row not contiguous, but distance varies; large, slightly convex, raised impunctate area in anterolateral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming 2 rows not covering entire width. Scutellum without archlike carina, with yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with large punctures, about 3.6 punctures / 0.3 mm, and interspaces as wide as 0.5–1 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII well punctuated (V–VI less so medially); IV–V with yellow microsetae medially and posterolaterally, VI with yellow ones laterally and brown ones medially, VII with brown ones. Paratergites IV–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: IV–VII on anterolateral corners


without or with sparse punctures; III–VII densely punctuated otherwise; III–VI and VIII with yellow microsetae, VII with brown ones. Segment VII with black iridescent setae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 78) with evenly curved ventral surface, hind margin with two very small triangular lobes and a moderate U-shaped emargination; sternum IX (Fig. 105) with two small apical processes and a small U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 158–161. Paramere in dorsal outline consistently converging to a narrowly rounded tip, apically subequal in length to median lobe and narrower; slightly convex in lateral outline. Sensory spinules (many) arranged in a dense cluster restricted to tip. Median lobe with broadly rounded apex in dorsal outline, slightly upcurved apically in lateral outline, forming a small dorsomedial carina forked apically, these parts not forming discrete lobes. Diagnosis. Among species with golden-orange head, pronotum and elytra, N. densiventris, N. semelius, and N. periergos are unique in the presence of microsculpture of transverse microlines on the epicranium. N. densiventris is very similar to N. semelius and N. periergos but differs in the shape of sternum VIII of males (Figs. 78, 96, 100): two very small triangular lobes extending beyond the posterior sternal margin and a moderate U-shaped medial emargination in N. densiventris; two strong triangular lobes and a very deep V-shaped medial emargination in N. semelius and N. periergos. Distribution. Known from elevations of 650–2000 m in Colombia and Ecuador (Fig. 58). Habitat. Unknown. Remarks. Collected in flight intercept traps and malaise traps. Material. Lectotype, „, here designated, with labels ‘Sogamoso Col.[ombia] Ost. Co. W. 01/1911 900 m’, ‘densiventris Bern. Typus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Lectotype Brachydirus densiventris Bernhauer des. Chatzimanolis 2003’ (in FMNH). – Paralectotype, „ with labels ‘„’, ‘Sogamoso Col. Ost. Co. W. 01/1911 900 m’, ‘Brachydirus densiventris Bern, det. Bernhauer Cotypus ms.’, ‘Dr. M. Bernhauer 10.12.donavit.1938’, ‘ex. coll. Scheerpeltz’, ‘COTYPUS Brachydirus densiventris Dr. Bernhauer’, ‘Paralectotype Brachydirus densiventris Bernhauer des. Chatzimanolis 2003’ (in NMW). – COLOMBIA: Ost Cordillieren, 900 m, Sogamoso, Bernhauer det. (1 in FMNH); Valle de Cauca PNN, Farallones de Cali, Anchicaya, 3o26Ń 76o48´W, 650 m, 11.ix.–16.x.2001, S. Sarria leg. malaise trap (3 in SEMC); Hulla PNN, Guarcharos Alto, El Mirador, 1o38Ń 76o6´W, 1980 m, 7–21.xii.2001, R. Paramero leg. malaise trap (1 in SEMC, 2 in UNCB). ECUADOR: Provincia de Morona Santiago, 5 km S Macas, 1050 m, 18.xii.1992, E.S. Ross leg. (1 in CASC); Napo-Pastaza, 6–8 mi W Mera, 1500 m, 10.ii.1955, E.I. Schlinger & E.S. Ross leg. (1 in CASC); Napo Prov. El Chaco, 2000 m, ii.1983, M. Sharkey & L. Masner leg. malaise trap (3 in CNCI).

4.13. Nordus dichromos sp.n. (Figs. 1, 2, 32, 63, 79, 162–165) Description. L e n g t h . 8.5–10.0 mm. C o l o r a t i o n . (1) Golden-orange morph: Head goldenorange; mandibles and labrum golden-orange; antenno-

19

meres all orange. Pronotum, elytra, and scutellum goldenorange. Abdomen black except segments I–II, VIII, and posterior 1/4 of VII golden-orange. Legs golden-orange. (2) Black morph: Head black; mandibles and labrum goldenorange; antennomeres all orange. Pronotum, elytra, and scutellum black. Abdomen black except segment VIII and posterior 1/4 of VII golden-orange. Legs black. H e a d . Microsetae yellow on anterior 3/4 and brown on posterior 1/4. Anteclypeus very small. Punctures in anterior half of epicranium very dense, about 6.7 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 3.6 punctures / 0.3 mm, punctures almost confluent. Length of eyes ca. 0.6ⴛ length of head, distance between eyes less than 1.5ⴛ length of eye. Middle of epicranium slightly convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 32): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th slightly shorter than 4th, 1st–5th longer than greatest width, 6th quadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 1.5ⴛ length; 6th–11th densely covered by microtrichae; 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 0.94; anterolateral angles not prominent, subcariniform; lateral margins concave at dorsal aspect; disc laterally with distinct gaps in punctuation; punctures large and shallow; medial impunctate area broadest anteriorly (width 2–3 punctures); punctures flanking this area in at least 1–2 sparse longitudinal rows separated by distance as wide as 1–1.5 punctures, punctures of a row not contiguous, but distance varies; large, convex, raised impunctate area in anterolateral quarters, and small impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with whitish-yellow microsetae and punctures covering posterior half. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, punctures almost contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III–IV with 0–4 discal microsetae; V without subbasal carina; V densely punctuated on medial disc, less dense medially on posterior margin; VI densely punctuated except medially; V–VI with grayish-yellow microsetae except medially, VII with dark brown ones, VIII with sparsely distributed yellow ones. Paratergites III–VII covered with whitish-yellow setae on posterior half. Sterna: IV–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–VI and VIII with yellow microsetae. Segments VI–VII with black iridescent setae throughout. Cerci slightly curved, basal 1/3 brown and distal 2/3 yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (Fig. 63) with broad and shallow emargination medially; sternum VIII (Fig. 79) with evenly curved ventral surface, hind margin with very narrow and shallow V-shaped emargination medially; sternum IX (identical to that in Fig. 110) with two moderate apical processes and a

20

moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 162–165. Paramere in dorsal outline almost parallel-sided from middle to subapical region, then forming a narrowly rounded tip, apically subequal in length to median lobe and narrower; slightly convex near apex in lateral outline. Sensory spinules (many) arranged in a dense cluster and restricted to tip. Median lobe with narrowly subtruncate or slightly emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus terminalis, N. omios, N. solitarius, and N. dichromos can be distinguished from all other Nordus species by the lacking (or extremely reduced) microsetation on the medial disc of abdominal tergum IV. These setae are also absent in N. fungicola, which differs from the above species by its convex rather than concave lateral margins of the pronotum. N. terminalis and N. omios can be distinguished from the other two species by the depressed elytra near scutellum, the very sparse punctuation on the elytra (distance between punctures larger than width of 1 puncture versus less than 1 in the other two species). N. solitarius can be distinguished from N. dichromos by the following characters: antennomere 4 subquadrate (longer than wide in N. dichromos); antennomere 5 subequal in length to 6 (5 longer than 6 in N. dichromos); epicranium more ot less flat (convex in N. dichromos), shape of sternum VIII in males as in Fig. 90 (versus Fig. 79 in N. dichromos). Distribution. Known from Ecuador (Fig. 21). Considering specimens with elevation data available, all but two blackmorph specimens were found at elevations of 530–2300 m, while all but two golden-orange-morph specimens were collected at elevations of 25–300 m. Habitat. White polypore fungi. Derivatio nominis. The specific epithet is derived from the Greek word ‘␦␫␹␳␻␮␱␵’ (two-colored) and refers to the presence of two color morphs. Remarks. Also collected from dung traps, carrion traps, malaise traps and flight intercept traps. Material. Holotype, ‚ golden-orange morph, with labels ‘Ecuador: Pichincha Rio Palenque Science Center, 0o36´0´´S 79o21´0´´W, 200 m, 25.iv.–6.vi.1996, P. Hibbs, ECU1H96 002, ex: FIT’, barcode label SM0092684, ‘HOLOTYPE Nordus dichromos Chatzimanolis, des. Chatzimanolis 2003’ (in SEMC). – Paratypes, 105, with labels: (Golden-orange morph:) ‘Ecuador: Pichincha Rio Palenque Science Center, 0o36´0´´S 79o21´0´´W, 200 m, 25.iv.–6.vi.1996, P. Hibbs, ECU1H96 002, ex: FIT’, barcode labels SM0092680, SM0092682, SM0092683, SM0092679, SM0092681, SM0092678, SM0170379 (7 in SEMC); ‘Ecuador: Pichincha Rio Palenque Science Center, 0o36´0´´S 79o21´0´´W, 200 m, 31.iii.–25.iv.1996, P. Hibbs, ECU2H96 007C, ex: malaise’, barcode label SM0170324 (1 in SEMC); ‘Ecuador: Pichincha Pr. Rio Palenque, 47 km S Santo Domingo, 20.vii.1979, T. Ray, human dung trap’ (3 in MCZC); ‘Ecuador: 27 mi SW Quevedo, Los Rios, 50 m, 2.ii.1955’, ‘E.I. Schlinger & E.S. Ross collectors’ (3 in CASC); ‘Ecuador: Pichilingue, Los Rios, 40 m, 2.ii.1955’, ‘E.I. Schlinger & E.S. Ross collectors’ (1 in CASC); ‘Ecuador: Pichincha Pr. 47 km S. Sto. Domingo, Rio Palenque Station, leg: S. Peck, 27–31.v.1975’, ‘carrion trap’ (4 in FMNH); ‘Ecuador Esm. Pr., Zapallo Grande, 25–30.x.1987, Mike Huybenez’ (1 in FMNH); ‘Balzapamba, 5.vi.1938, ECUAD. elev. 700 m. Wm. Clarke-MacIntyre leg.’ (1 in FMNH); ‘ECU. Pich. 47 S. Sto. Domingo, Rio Palenque Station, 30.v.1975, 25 m, S. & J. Peck’ (3 in AMNH, 15 in


CNCI, 6 in SEMC); same locality data and collector but 25– 30.v.1975 (5 in CNCI); same locality data and colelctor but 250 m, 22–31.viii.[19]76 (2 in CNCI); same locality data but 250 m, 23–24.v.[19]75, S. Peck (2 in CNCI); same locality data but 300 m, 22–28.ii.1976, J.M. Campell (10 in CNCI); same locality data but 250 m, 17–25.ii., S.A. Marshall (2 in CNCI); same locality data but 700 m, 22–27.ii.1976, H. & A. Howden (1 in CNCI); same locality data, 18–21.ii.1979, L. Ling, dungtrap (1 in CNCI); ‘ECU, Rio Palenque, 49 km S. Santa Dom. Pich. Prov. malaise trap, J. Bellwood’ (1 in CNCI); ‘ECU, Los Rios Quevedo, vii.[19]76, Bolle (1 in CNCI). (Black Morph:) ‘Ecuador: Pichincha Tinalandia, Santo Domingo 16 km E, 750 m, 0o16´53´´S 79o3´39´´W, 27– 27.iii.1999, R. Brooks, D. Brzoska, ECU1B99 048 ex: FIT’, barcode labels SM0153309, SM0153323, SM0153321, SM0153317, SM0153326, SM0153324, SM0153322, SM0153318, SM0153296, SM0153305, SM0153308, SM0153312, SM0153304, SM0153311, SM0153310, SM0153306, SM0153315, SM0153319, SM0153307 (19 in SEMC); ‘Ecuador: Pichincha, Pedro Vicente Maldonado 3.5 km N, 530 m, 0o6´44´Ń 79o3´21´´W, 29.iii.1999, R. Brooks ECU1B99 066 ex: fungus, mushroom, polypore white’, barcode label SM0153175 (1 in SEMC); ‘Ecuador: Pichincha, 45 km NNW Quito Macquifucuna Station, 1600–1650 m, 18.v.– 5.vi.1996, ECU1H96 022, P. Hibbs, ex: FIT’, barcode labels SM0092766, SM0075101 (2 in SEMC); ‘Ecuador: Pichincha, Tinalandia, 15 km E S. Domingo, 650 m, D. Brzoska, 19.iii.1995’, barcode label SM0139615 (1 in SEMC); ‘Ecuador, Pich. E S. Domingo, 8–14.v.1988, Hanson & Bohart’, barcode labels SM0058158, SM0058157 (2 in SEMC); ‘Ecuador: Napo, 2300 m, Sierra Azul, Haciente Aragon, 0o40´0´´S 77o55´0´´W, 17.ii.–26.iii.1996, ECU1H96 009, P. Hibbs ex: FIT’, barcode label SM0092653 (1 in SEMC); ‘Ecuador: Pichincha, Mindo, 10.6 km W Mindo R., 0o4´23´´S 78o45´4´´W, 1445 m, 26–29.iii.1999, R. Brooks, D. Brzoska, ECU1B99 064 ex: FIT’, barcode label SM0153279 (1 in SEMC); ‘ECU, Pich. 16 km SE Sto. Domingo Tinalandia, 680 m, 20–21.vi.1975 S. & J. Peck’ (3 in CNCI); ‘ECU, Pich., 47 km S Sto. Domingo, Rio Palenque Sta. 30.v.[19]75, 25 m, S. & J. Peck’ (1 in CNCI); ‘ECU Guayas-Manabi borders, 10 m, Rio Ayampo, 26.vii.1976, S. & J. Peck’ (1 in CNCI); ‘ECU: Pich., Rio Palenque Sci. Center, 1–14.vii.1975, A. Forsyth’ (1 in CNCI); ‘ECU, Pich. Prov. 800 m, Tinalandia, ii.1983, L. Masner’ (2 in CNCI). All paratypes with label ‘Paratype Nordus dichromos Chatzimanolis des. Chatzimanolis 2003’.

4.14. Nordus diversiventris (Bernhauer, 1917) (Figs. 33, 80, 115, 166–169) Brachydirus diversiventris Bernhauer, 1917: 22. Nordus diversiventris (Bernhauer); HERMAN (2001: 3579). Description. L e n g t h . 8.5–10.5 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres 1–8 yellow, 9–11 darker orange to brown. Pronotum, elytra, and scutellum goldenorange. Abdomen black except segments I, II, VIII, and posterior 1/3 of VII golden-orange. Legs yellow. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium very dense, about 10.2 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 5.8 punctures / 0.3 mm, punctures almost confluent. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 33): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th slightly shorter than 4th, 1st–5th longer than greatest width, 6th quadrate, 7th–10th wider

21


than greatest length, width of distal margin of 9th and 10th at least 1.5ⴛ length; 6th–11th densely covered by microtrichae; 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 0.98; anterolateral angles not prominent, subcariniform; lateral margins almost parallel to each other in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and shallow; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in 1– 2 moderately dense longitudinal rows separated by a distance as wide as 0.5 punctures, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with whitishyellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with medium-sized punctures, about 3.6 punctures / 0.3 mm, and interspaces as wide as 0.5–1 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; V–VII extremely punctuated; IV with few microsetae on medial disc; IV–V and VIII covered with yellow microsetae, VI with yellow ones laterally and black ones medially, VII with black ones. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; III–IV and VIII with yellow microsetae. Segment VII with black iridescent microsetae in anterolateral quarters. Cerci slightly curved, basal 1/3 light brown and distal 2/3 yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 80) with evenly curved ventral surface, hind margin with two very small triangular lobes and a small V-shaped emargination medially; sternum IX (identical to that in Fig. 110) with two moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a slightly pointed extension (Fig. 115). M a l e g e n i t a l i a . As in Figs. 166–169. Paramere in dorsal outline weakly but consistently converging to a broadly rounded tip, apically subequal in length to median lobe and narrower; slightly convex near apex in lateral outline. Sensory spinules (many) arranged in a dense cluster and restricted to tip. Median lobe with broad subtruncate and slightly emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Among the golden-orange species N. diversiventris, N. testaceus, and N. styloceros can be identified by the almost parallel lateral margins of the pronotum. These three species are easily distinguished by the unique shapes of their aedeagi (Figs. 166, 237, 245) and sterna VIII in males and by the following characters in both sexes: N. diversiventris and N. styloceros differ from N. testaceus by the medial impunctate area on the pronotum being as wide as 1–1.5 punctures (0.5 punctures or less in N. testaceus). N. diversiventris differs from N. styloceros by antennomere 5 being longer than wide (subquadrate in N. styloceros) and

punctures on elytra being separated by a distance as wide as 0.5–1 punctures (confluent in N. styloceros). Distribution. Known from elevations of 500–1980 m in Colombia, Panama, and Peru (Fig. 60). There are no specimens known from Bolivia as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Habitat. Rotting palm fruits. Remarks. Collected in flight intercept and malaise traps, and human feces traps. Material. Holotype, ‚ with labels ‘Inner Peru Cordilleren Cerro de Pasca’, ‘diversiventris Bernh. Typus unic.’, ‘Chicago NHMuseum M. Bernhauer Collection’ (in FMNH). BERNHAUER (1917) states in the original description that he had available only a single specimen, which therefore is the holotype. – COLOMBIA: Anchicaya, 1000 m, 22.vii.1970, J.M. Campell leg. (3 in CNCI); Valle de Cauca PNN, Farallones de Cali, Anchicaya, 3o26Ń 76o48´W, 650 m, 2001, S. Sarria leg. malaise trap (10 in SEMC; 10 in UNCB); Hulla PNN, Guarcharos Alto, El Mirador, 1o38Ń 76o6´W, 1980 m, 22.xii.2001–5.i.2002, R. Paramero leg. malaise trap (1 in SEMC). PANAMA: Colon, 14 km N jct. Escobal & Pina Rd., 2–11.vi.1996, J.S. Ashe & R. Brooks leg. FIT (1 in SEMC); Darien, Cana Biological Station, 500–550 m, 7o45´18´Ń 77o41´6´´W, 9.vi.1996, J.S. Ashe & R. Brooks leg. human feces trap (1 in SEMC); Darien, Cana Biological Station, 500– 550 m, 7o45´18´Ń 77o41´6´´W, 6.vi.1996, J.S. Ashe & R. Brooks leg. rotting palm fruits (1 in SEMC); Darien, Estacion Ambiental Cana, 500 m, 7o45´20´Ń 77o41´6´´W, 7– 10.vi.1996, R. Anderson leg. human feces trap (1 in SEMC). PERU: Chanchamayo (1 in BMNH); Valle Chanchamayo, 800 m, viii.1957, Wervauchex leg. (1 in NMW); Klima leg., ex. coll. Scheerpeltz (1 in NMW).

4.15. Nordus elegans sp.n. (Figs. 5, 49, 94, 170–173) Description. L e n g t h . 9.0–10.5 mm. C o l o r a t i o n . Head dark brown; mandibles and labrum orange; antennomeres 1 and 7–11 brownish-yellow, 2–6 brown. Pronotum golden-orange, elytra dark orange to brown (one paratype with lighter colored elytra), scutellum black. Abdomen light brown except posterior margins of segments III–VI and entire segment VIII orange, and VII brown with posterior 1/3 orange. Legs golden-orange except metatibiae and -tarsi darker orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 5.8 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior half of epicranium about 2.2 punctures / 0.3 mm, distance between punctures as wide as 0.5–1 punctures; middle of posterior half without punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1–1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 49): 1st thickened clublike, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th subequal in length to 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate, 6th–10th wider than greatest length, width of distal margin of 9th and 10th at least twice length; 6th– 11th densely covered by microtrichae; 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum longer than wide, width : length = 0.87; anterolateral angles not prominent, subcariniform;

22

lateral margins strongly concave in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures medium-sized and shallow; medial impunctate area broadest anteriorly (width 2–3 punctures); punctures flanking this area in 1–2 sparse longitudinal rows separated by a distance as wide as 1–1.5 punctures, punctures of a row not contiguous, but distance varies; large, slightly convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming 2 rows, not covering entire width. Scutellum without arch-like carina, with yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, and interspaces as wide as 1–1.5 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V with subbasal carina; IV–VII well punctuated; IV–VIII with yellow microsetae medially and posterolaterally. Paratergites IV–VII covered with yellow microsetae on posterior half. Sterna: IV–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise, III–VIII with yellow microsetae. Segment VII with black iridescent setae throughout. Cerci thin and straight, basal half light brown and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 94) with evenly curved ventral surface, hind margin with a very deep and broad U-shaped emargination medially; sternum IX (identical to that in Fig. 111) with no apical processes and a deep U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 170–173. Paramere in dorsal outline very wide, widened apically, with broadly rounded and then narrow, acute tip (thus, distal margins slightly sinuate), apically subequal in length to median lobe and slightly narrower; dorsally slightly concave near base in lateral outline; with diamond-shaped depression dorsomedially below tip. Sensory spinules (many) arranged in a pair of clusters restricted to tip. Median lobe with broadly rounded apex in dorsal outline, upcurved apically in lateral outline, forming apically two widely separated, dorsal lobes (their bases at an angle of 90o to lateral margin of median lobe) but no dorsomedial carina basal to these. Diagnosis. Nordus cherylae, N. elegans, N. portokalis, and N. speciosus can be distinguished from all other species of Nordus by the presence of a subbasal carina (in addition to the basal one) on abdominal tergum V. N. portokalis can be distinguished from the other three species by the presence of the very distinctive orange abdomen (black in the other species) and the subquadrate pronotum (pronotum longer than wide in other species). N. speciosus can be distinguished from N. cherylae and N. elegans by the metallic green head, pronotum and elytra. In N. cherylae head, pronotum and elytra are all black, whereas in N. elegans the head is dark brown, the pronotum golden-orange, and the elytra dark orange to brown; sternum VIII in females is evenly rounded posteriorly in N. elegans (Fig. 114), and the aedeagus is as in Figs. 170–173, while in N. cherylae sternum VIII in females is sinuate (Fig. 118) and the aedeagus is as


in Figs. 150–153. Distribution. Known from Peru (Fig. 21). Habitat. Unknown. Derivatio nominis. The specific epithet is derived from the Latin word ‘elegans’ (elegant). The word ‘elegans’ was written on labels by Bernhauer, but he never published the name. Material. Holotype, ‚ with labels ‘Callanga, Peru’, ‘elegans Brh. Typ.’, ‘Chicago NHMus. M. Bernhauer Collection’, ‘HOLOTYPE Nordus elegans Chatzimanolis des. Chatzimanolis 2003’ (in FMNH). – Paratypes, 1‚, 1„ with labels ‘Callanga Peru’, ‘Brachydirus elegans Brh.’, ‘Bernhauer det.’, ‘coll. Kraatz’, ‘Coll. DEIC, Eberswalde’, ‘Paratype Nordus elegans Chatzimanolis des. Chatzimanolis 2003’ (in DEI).

4.16. Nordus elytisi sp.n. (Figs. 50, 95, 174–177) Description. L e n g t h . 8.5–10.0 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres all yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/3 of VII golden-orange. Legs yellow. H e a d . Microsetae yellow on anterior 3/4 and brown on posterior 1/4. Anteclypeus very small. Punctures in anterior half of epicranium very dense, about 6.7 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 3.6 punctures / 0.3 mm, punctures almost confluent. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 50): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th slightly shorter than 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate; 6th–10th wider than greatest length, width of distal margin of 9th and 10th at least 1.5ⴛ length; 6th–11th densely covered by microtrichae; 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.01; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in at least 2 dense, contiguous longitudinal rows, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and small impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with whitish-yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; V densely punctuated on medial disc but less dense medially on posterior margin; VI densely punctuated except medially; IV with grayish-yellow microsetae on medial disc, V–VI with grayish-yellow ones except


medially, VII with dark brown ones, VIII with sparsely distributed yellow ones. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; III–VI and VIII with yellow microsetae. Segment VII with black iridescent setae throughout. Cerci slightly curved, basal half light brown and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 95) with evenly curved ventral surface, hind margin with two very small triangular lobes and a small V-shaped emargination medially; sternum IX (identical to that in Fig. 110) with two moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 174–177. Paramere in dorsal outline very wide, almost parallel-sided from middle to subapical region, then strongly narrowed forming an acute tip, apically subequal in length to median lobe; dorsally concave basally and dorsally convex distally in lateral outline. Sensory spinules (few) arranged in a medial band and restricted to tip. Median lobe with broad emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus cribricollis, N. elytisi, and N. seferisi can be distinguished from all other golden-orange Nordus species by antennomere 6 being wider than long (subquadrate in all other species except N. portokalis, which by its orange abdomen differs from the three abovementioned species with black abdomen). N. cribricollis can be distinguished from N. seferisi and N. elytisi by antennomere 4 being longer than wide rather than subquadrate. In addition, N. cribricollis males have a paramere (Fig. 154) much narrower than in the other two species (Figs. 174, 218). N. elytisi differs from N. seferisi by the denser punctuation on the pronotum near the medial impunctate area, and males can be distinguished by the shape of the aedeagus (Figs. 174–177 and 222–225). Distribution. Known from elevations of 880–1100 m in Argentina and Bolivia (Fig. 21). Habitat. Unknown. Derivatio nominis. The specific epithet is in honor of Odysseas Elytis, a Greek Nobel laureate. Material. Holotype, ‚ with labels ‘Salta [Argentina], Dept Oran., Rio Saluga, P. Denier, 16.vi.1933’, ‘Brachydirus bruchi Brh. Typus unic.’, ‘Chicago NHMus. M. Bernhauer Collection’, ‘HOLOTYPE Nordus elytisi Chatzimanolis des. Chatzimanolis 2003’ (in FMNH). – Paratypes, 10 with labels: ‘Argent. Jujuy Padre Lozano, 15–17.xii.1967, C. C. Porter’ (1 in MCZC); ‘Argentina, Salta: Aguaray, i.1957, P. Wygodzinsky’ (2 in AMNH); ‘Arg. Salta Prov. El Rey Nat. Park, 880 m, 6–16.xii.1987, S. & J. Peck, Rio Los Puestos, malaise trap-FIT, Prosopis forest’ (1 in FMNH); ‘Arg. Salta, Pocitos, xii.1974, Bolle’ (1 in CNCI, 1 in SEMC); ‘Argentina, Salta, Oran, i.1975’ (2 in CNCI); ‘Argentina, Pcia Salta, Pocitos x.1961, A. Martinez leg.’ (1 in CNCI); ‘Bolivia, Yungas, La Paz, 6.i.1976, 1100 m, Bolle’ (1 in CNCI). All paratypes with label ‘Paratype Nordus elytisi Chatzimanolis des. Chatzimanolis 2003’.

4.17. Nordus facialis (Erichson, 1839) (Figs. 35, 81, 178–181) Staphylinus facialis Erichson, 1839: 354. Brachydirus facialis (Erichson); SCHEERPELTZ (1971: 98). Nordus facialis (Erichson); HERMAN (2001: 3580).

23

Description. L e n g t h . 8.5–10.0 mm. C o l o r a t i o n . Head golden-orange, in females anterior 1/ – 2/ shining black; mandibles and labrum orange; anten2 3 nomeres all golden-orange. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I–II, VIII, and posterior 1/8 of tergum VII golden-orange. Legs golden-orange, in females metafemora and -tibiae darker orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 5.8 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior half about 3.1 punctures / 0.3 mm, punctures almost contiguous. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium slightly convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 35): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate, 6th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th covered by dense dark microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 1.05; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in 2–3 moderately dense longitudinal rows separated by a distance as wide as 0.5 punctures, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and a moderate impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with large punctures, about 3.1 punctures / 0.3 mm, and interspaces as wide as 0.5 puncture, covered with yellow microsetae. Legs with yellow macrosetae, in females metafemora and -tibiae with brown macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV strongly punctuated on medial disc; V–VI strongly punctuated but without punctures anteriorly; VII uniformly punctuated; IV–VI covered with yellow microsetae laterally, VII covered with brown ones. Paratergites III–VI covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; with dark brown microsetae medially and yellow ones laterally. Segment VI–VII with black iridescent setae in anterolateral quarters. Cerci slightly curved, basal half dark brown to black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum (identical to that in Fig. 63) VII with broad and shallow emargination medially; sternum VIII (Fig. 81) with evenly curved ventral surface, hind margin with broad and moderately deep U-shaped emargination medially; sternum IX (identical to that in Fig. 104) with two small to moderate apical processes and a moderate U-shaped

24

emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 178–181. Paramere in dorsal outline slightly but consistently converging to a subacute tip, apically subequal in length to median lobe and narrower; dorsally convex near apex in lateral outline. Sensory spinules (few) arranged in two very short longitudinal rows close to midline and restricted to tip. Median lobe with subtruncate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus facialis and N. championi can be distinguished from all other golden-orange species of Nordus by the combination of the following characters: epicranium convex; punctuation on anterior part of epicranium dense (but not very dense as in N. fungicola); and scutellum punctuated only in its posterior half. N. championi and N. facialis can be distinguished by the following characters: punctures on pronotum are shallow in N. championi and deep in N. facialis; punctures on pronotum near medial impunctate area dense in N. facialis and less dense in N. championi; and antennomere 4 longer than wide in N. championi and subquadrate in N. facialis. Distribution. Known from elevations of 30–740 m in Brazil, French Guiana, Guyana, Suriname, and Venezuela (Fig. 60). Habitat. Unknown. Remarks. Collected in flight intercept traps and in banana fruit traps. Material. Holotype, with abdomen strongly damaged, and with labels ‘Type’, ‘5891’, ‘Hist. – Coll. (Coleoptera) Nr. 5891 Staphylinus fascialis [sic] Erichs. Cayenna, Buquet. Zool. Mus. Berlin’ (in ZMHB). The third label is not the original type label and the mistake in the species epithet was not made by Erichson. This is the only specimen in the type series and therefore is the holotype. – BRAZIL: Amapa, Rio Amapari, 10.vii.1969, J. Lane coll. (1 in CNCI). FRENCH GUIANA: Roura, 27.4 km SSE, 280 m, 4o44´20´Ń 52o13´25´´W, 23– 24.v.1997, J.S. Ashe & R. Brooks leg. FIT (4 in SEMC); Roura, 39.4 km SSE, 270 m, 4o32´43´Ń 52o8´26´´W, 29.v.– 10.vi.1997, J.S. Ashe & R. Brooks leg. FIT (2 in SEMC); Roura, 18.4 km SSE, 240 m, 4o36´38´Ń 52o13´25´´W, 29.v.– 10.vi.1997, J.S. Ashe & R. Brooks leg. FIT (5 in SEMC); Roura, 8.4 km SSE, 200 m, 4o40´41´Ń 52o13´25´´W, 25– 29.v.1997, J.S. Ashe & R. Brooks leg. FIT (3 in SEMC); Matoury, 41.5 km SSW on H’way N2, 50 m, 4o37´22´Ń 52o22´35´´W, 29.v.–9.vi.1997, J.S. Ashe & R. Brooks leg. FIT (2 in SEMC); Saul, 7 km N, Les Eaux Claires, Mt. La Fumee, 220–500 m, 3o39´46´Ń 53o13´19´´W, 1–9.vi.1997, J.S. Ashe & R. Brooks leg. FIT (11 in SEMC); Saul, Mt. Galbao summit, 740 m, 3o37´18´Ń 53o16´42´´W, 5–7.vi.1997, J.S. Ashe & R. Brooks leg. FIT (5 in SEMC); Cayenne, 33.5 km S and 8.4 km NW of H’way N2 on H’way D5, 30 m, 4o48´18´Ń 52o28´41´´W, 29.v.–9.vi.1997, J.S. Ashe & R. Brooks leg. FIT (1 in SEMC); Cayenne, Kraatz leg. (1 in DEI). GUYANA: Region 8, Iwokrama Forest, 26 km SW Kurupukari, Iwokrama Mt. base camp, 4o20´17´Ń 58o48´38´´W, 60–400 m, 23– 25.v.2001, R. Brooks & Z. Falin leg. FIT (7 in SEMC); Region 8, Iwokrama Forest, Kabocalli Field Stn., 4o17´4´Ń 58o30´35´´W, 60 m, 3–5.v. 2001, R. Brooks & Z. Falin leg. FIT (1 in SEMC); Region 8, Iwokrama Forest, 1 km W Kurupukari, Iwokrama Field Stn., 4o40´19´Ń 58o41´4´´W, 60 m, 20– 25.v.2001, R. Brooks & Z. Falin leg. FIT (1 in SEMC); Bartica, Br. Guiana [Guiana], C.E. Tottenham leg. (1 in BMNH). SURINAME: Saramacca, West Suriname Rd., 108 km WSW Zanderij Airport, 90 m, 5o13´37´Ń 55o52´54´´W, 8– 10.vi.1999, Z. Falin & B. DeDjin leg. FIT (1 in SEMC); Marowijne, Nassau Mt., 480–500 m, 4o48´56´Ń 54o36´20´´W, 1–4.vi.1999, Z. Falin & B. DeDjin leg. FIT (6 in SEMC); Marowijne, Palumeu, 160 m, 3o20´56´Ń 55o26´18´´W, 7– 8.vii.1999, Z. Falin leg. FIT (1 in SEMC); Brokopondo,


Brownsberg Nature Preserve, Witi Creek trail, 480 m, 4o56´55´Ń 55o10´53´´W, 23–25.vi.1999, Z. Falin, A. Gangadin & H. Hiwat leg. FIT (3 in SEMC); Commewijne, Akintosoela, 32 km SE Suriname River bridge road to Redi Doti, 40 m, 5o16´17´Ń 54o55´15´´W, 29.vi.–9.vii.1999, Z. Falin & B. DeDjin leg. FIT (5 in SEMC). VENEZUELA: Bolivar, La Gran Sabana Region, Hwy-170 89 km, 6o6´58´Ń 61o25´10´´W, 29–30.vii.1994, T. Philips leg. banana fruit trap (2 in SEMC).

4.18. Nordus fungicola (Sharp, 1884) (Figs. 18, 34, 82, 182–185) Brachydirus fungicola Sharp, 1884: 351. Nordus fungicola (Sharp); HERMAN (2001: 3580). Description. L e n g t h . 10–12.3 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres 1–5 golden-orange, 6–11 dark orange. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I–II, VIII, and posterior 1/3 of VII golden-orange. Legs golden-orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 3.5– 4.4 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 1.8–2.7 punctures / 0.3 mm, punctures almost contiguous. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 34): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st– 5th longer than greatest width, 6th wider (basal margin) than greatest length, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum (sub)quadrate, width : length = 1.02– 1.09; anterolateral angles extremely prominent and acutely delimited, forming angle of about 90o, cariniform; lateral margins moderately to strongly convex in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and shallow; medial impunctate area broa-dest anteriorly (width 3–4 punctures); punctures flanking this area in 2 longitudinal rows separated by distance as wide as 2–3 punctures, punctures of a row not contiguous, but distance varies; small and not raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow and black microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 4.4–4.9 punctures / 0.3 mm, and interspaces as wide as 1 puncture, covered with yellow and black microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III–IV with 0–4 discal microsetae; V without subbasal carina; V strongly punctuated except medially; VI–VII extremely punctuated; VI–VII covered laterally with yellow microsetae. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; with dark brown


microsetae medially and yellow ones laterally. Segment VII with black iridescent setae in anterolateral quarters. Cerci slightly curved, basal half dark brown to black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII with broad and shallow emargination medially (identical to that in Fig. 63); sternum VIII (Fig. 83) with evenly curved ventral surface, hind margin with broad and moderately deep U-shaped emargination medially; sternum IX (identical to that in Fig. 104) with two small to moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 182–185. Paramere in dorsal outline consistently converging to a rounded tip, apically subequal in length to median lobe and narrower; slightly convex near apex in lateral outline. Sensory spinules arranged in two very short longitudinal rows and restricted to tip. Median lobe with subtruncate apex in dorsal outline, strongly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus velitaris, N. fungicola, and N. major can be distinguished from other species of Nordus by the combination of a very convex epicranium and antennomere 4 being longer than wide. N. fungicola and N. major differ from N. velitaris by the convex rather than concave lateral margins of the pronotum. Furthermore, the temporal area of the head is strongly expanded in N. major but only slightly so in N. fungicola; a pair of raised anterolateral impunctate areas on the pronotum is present in N. fungicola but absent in N. major; the antyclepeus is reduced in N. fungicola but well developed in N. major. Distribution. Known from elevations of 0–800 m in Colombia, Costa Rica, Ecuador, Panama, and Venezuela (Fig. 20). Habitat. Rotting fruitfalls and rotting fungi. For a detailed account of the habitat and natural history see CHATZIMANOLIS (2003). Remarks. Collected also in flight intercept traps, malaise traps, carrion traps and banana traps. Specimens from Costa Rica appear to have a less concave pronotum than specimens from more southern localities. Material. Lectotype, ‚, here designated, glued on paper card, with labels: on paper card mount in D. Sharp’s handwriting ‘‚, Type, Brachydirus fungicola, Bugaba, Champion’, below ‘Bugaba, Panama, Champion’, ‘B. C. A. Col. I. 2, Brachydirus fungicola Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Lectotype Brachydirus fungicola Sharp des. Chatzimanolis 2002’, ‘Nordus fungicola (Sharp) det. S. Chatzimanolis 2002’ (in BMNH). – Paralectotypes, 11 specimens (9 in BMNH, 2 in FMNH): In BMNH: 1‚, 1„, on same paper card in D. Sharp’s handwriting ‘‚, „, Brachydirus fungicola, D. S. Bugaba, Champion’, below ‘Bugaba, Panama, Champion’, ‘B. C. A. Col. I. 2, Brachydirus fungicola Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus fungicola Sharp des. Chatzimanolis 2002’. 1‚, 1„, on same paper card in D. Sharp’s handwriting ‘‚, „, Brachydirus fungicola, D. S. Bugaba, Champion’, below ‘Bugaba 800–1500ft Champion’, ‘B. C. A. Col. I. 2, Brachydirus fungicola Sharp’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus fungicola Sharp des. Chatzimanolis 2002’. 1„, on paper card in D. Sharp’s handwriting ‘Brachydirus fungicola, D. S. V. de Chiriqui, Champion’, below, ‘V. de Chiriqui 3–4000ft, Champion’, ‘B. C. A. Col. I. 2, Brachydirus fungicola Sharp’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus fungicola Sharp des. Chatzimanolis 2002’. 1‚, on

25

paper card in D. Sharp’s handwriting ‘Brachydirus fungicola, Bugaba, Champion’, below, ‘Bugaba 800–1500 ft Champion’, ‘B. C. A. Col. I. 2, Brachydirus fungicola Sharp’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus fungicola Sharp des. Chatzimanolis 2002’. 1‚ with labels ‘Bugaba 800–1500 ft Champion’, ‘B. C. A. Col. I. 2, Brachydirus fungicola Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus fungicola Sharp des. Chatzimanolis 2002’. 1‚, 1„, on same paper card in D. Sharp’s handwriting ‘‚, „, Types, D. S. Brachydirus fungicola, D. S. Bugaba, Champion’, below [round red and white] ‘type’, ‘Bugaba, Panama, Champion’, ‘B. C. A. Col. I. 2, Brachydirus fungicola Sharp’, [upside down] ‘Sharp Coll. 1905-313’, [round light blue and white] ‘syntype’, ‘Paralectotype Brachydirus fungicola Sharp des. Chatzimanolis 2002’. In FMNH: 1„, 1‚, on same paper card in D. Sharp’s handwriting ‘‚, „, Brachydirus fungicola, D. S. Bugaba, Champion’, below ‘Bugaba, Panama, Champion’, ‘B. C. A. Col. I. 2, Brachydirus fungicola Sharp’, [upside down] ‘Sharp Coll. 1905-313’, ‘Sharp Colln. by exchange with the British Museum (N.H.)’, ‘Paralectotype Brachydirus fungicola Sharp des. Chatzimanolis 2002’. – COLOMBIA: Camp Curiche, 1967 (1 in FMNH); Choco Dept., Camp teresital, 1967 (4 in FMNH); Cali, Fassl leg. (1 in FMNH); Choco, Rio Tarapal, 100 m, 20–30.vii.1962, D.L. Hugh-Jones leg. (1 in BMNH); Huaso Ost-Colomb. (2 in FMNH); Le Bas, A. Breyer leg. (2 in ISNB); Lake Sepatoza region, Chiriguana region, 8.ix.1924, C. Allen leg. (1 in BMNH); Magdalena PNN, Tayrone Zaino, 11o20Ń 74o2´W, 50 m, 22.xi.–4.xii.2000, R. Herniquez leg. malaise trap (1 in SEMC); Magdalena, Carare Opon, Umg. Bogota, 4.viii.1968, H. Sturm leg. (1 in NMW); Rio Frio, Mgd., Darlington leg. (4 in MCZM); Rio Frio, 17.vii.1927, George Salt leg. (3 in BMNH); P’to Berrio Ant., 15.viii.1938, H. Dybas leg. (1 in FMNH); San Mante (1 in FMNH); San Esteban, iii.1888, E. Simon leg. (2 in BMNH); Valley of Magdalene, R. Carare-Opor, 4.viii.1968, Bet. Puerto Berrio of Barrancabermeja tropical rain forest, 150 m, H. Strum leg. (1 in AMNH). COSTA RICA: Prov. Guana., 3 km NO de Nacaome P.N. Barra Honda, 100 m, vii.1995 (1 in INBC); Prov. Guana., Est. Las Pailas, P.N. Rincon de la Vieja, 800 m, viii–ix.1993, G. Rogdiguez & J. Siezar leg. (1 in INBC); Puntarenas Prov., Rincon, 7km W, Osa Peninsula, Est. F.N. Aguas Buenas, 50 m, 21–25.vi.1997, S. & J. Peck leg. FIT (12 in SEMC); Puntarenas, Corconado National Park, Sirena, upper Rio Claro trail, 100 m, 8o28´29´Ń 83o35´8´´W, 28.vi.– 1.viii.2000, Z. Falin leg. FIT (65 in SEMC); Puntarenas, R.F. Golfo Dulce, 3 km SW Rincon, 10 m, v.–vi.1992, P. Hanson leg. FIT (16 in SEMC); Prov. Puntarenas, Est. Queb. Bonita, 50 m, Res. Biol. Carara, 18–29.x.1992, R. Guzman leg. (316 in INBC); Prov. Punta., R.N. Cabo Bianco, Est San Miguel, ix.1993, M. Ramirez leg. (1 in INBC); Prov. Puntarenas, Rancho Quemado, 200 m, Peninsula de Osa, vii.1992, M. Segura leg. (199 in INBC); Prov. Punt., Sector Laguna Meandrica, 100 m, R.B. Carara, vi.1990, R. Zuniga leg. (2 in INBC); Prov. Puntarenas, Sirena, Corcovado, N.P., 0–100 m, 1989, G. Fonseca leg. (62 in INBC); Prov. Punta., Est. Bijagual Res. Biol. Carara, 500 m, v.1993, J.C. Saborio leg. (4 in INBC); Prov. Puntarenas, Est. Carara, Res. Biol. Carara, 200 m, ii.1990, R. Zuniga leg. (1 in INBC); Prov. Punta., Bosque Esquinas Pen Osa, A.C. Osa, 200 m, vi.1994, M. Segura leg. (6 in INBC); Prov. Puntarenas, Est, Aguijas Sendero Hajo, 300 m, 22–30.viii.1997, M. Lobo leg. fruit trap (44 in INBC); Puntarenas Prov., Hincon de Osa, 14–26.vii.1969, T. Schuh & J. Crane leg. (3 in AMNH); Corcovado National Park, Osa Peninsula, 6.vii.1977, D.H. Jansen leg. (2 in INBC). ECUADOR: Granada (1 in BMNH). PANAMA: Bugaba, 800–1500 ft, Champion, B. C. A. Col. I. 2, (11 in BMNH, 1 in CNCI, 1 in FMNH); Canal Zone, Barro Colorado Island, 9o11Ń 79o51´W (3 in AMNH, 1 in CASC, 3 in CNCI, 8 in FMNH, 39 in MCZM, 171 in SEMC, 10 in USNM); Canal Zone, Skunk Hollow, G. NW Gatun locks, 13.vii.1976, B.C. Ratchliffe leg. (3 in UNSM); Canal Zone, Madden Forest pres., 20.vii.1976, A. Newton leg. around fermented palm fruit (2 in FMNH); Colon, Prq. Nat. Soberania, 9o07Ń 79o45´W, 40 m, J.S. Ashe & R. Brooks leg. (23 in SEMC); Darien, Cana Biological Station, 500–600 m, 7o45´18´Ń 77o41´6´´W, vi.1996, J.S. Ashe & R. Brooks leg. FIT (13 in SEMC); Darien Prov., Santa Fe, 1967, malaise trap (3 in FMNH); France Field, v.1930, Bierig leg. (2 in AMNH, 5 in FMNH); Gaboa, 9.x.1938 (5 in FMNH); Lino, 800 m, Fassl leg. (4 in FMNH, 3

26

in NMW); Panama Prov., Gamboa, Old Gamboa Rd. (30 in SEMC); Panama Prov., Pipeline Rd. (26 in SEMC); Panama Prov., Parq. Nat. Soberania, Pipeline Road km 2–4, 5– 15.vii.1994, B. & I. Ratchliffe & M. Jameson leg. (2 in UNSM); Pan. Prov., Altos de Maje, 6–15.x.1975 (1 in FMNH). VENEZUELA: no other labels (2 in BMNH).

4.19. Nordus gebieni (Bernhauer, 1917) (Figs. 36, 83, 106, 186–189) Brachydirus gebieni Bernhauer, 1917: 20. Nordus gebieni (Bernhauer); HERMAN (2001: 3580). Description. L e n g t h . 8.0–10 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres all yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/3 of VII golden-orange. Legs golden-orange. H e a d . Microsetae yellow on anterior 3/4, brownish-yellow on posterior 1/4. Anteclypeus very small. Punctures in anterior 3/ of epicranium dense, about 7.1 punctures / 0.3 mm, not 4 elongate longitudinally, margins of punctures confluent; in posterior part about 4.4 punctures / 0.3 mm, punctures almost contiguous. Length of eyes ca. 0.8ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 36): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–4th longer than greatest width, 5th subquadrate, 6th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 0.98; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (as wide as 2–3 punctures); punctures flanking this area in at least 3 dense, contiguous longitudinal rows, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayishyellow microsetae and punctures covering posterior 3/4. Elytra not depressed near scutellum, with large punctures, about 3.1 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with yellow microsetae on medial disc; V densely punctuated on medial disc but with fewer punctures medially on posterior margin; VI densely punctuated except medially; V with yellow microsetae throughout, VI with yellow ones laterally and brown ones medially, VII with dark brown ones, VIII with yellow ones. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely


punctuated otherwise; III–VI and VIII with yellow microsetae. Segment V–VII with black iridescent microsetae throughout. Cerci slightly curved, basal half light brown and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 63) with broad and shallow emargination medially; sternum VIII (Fig. 83) with evenly curved ventral surface, hind margin with very deep V-shaped notch medially; sternum IX (Fig. 106) with two moderate to large apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 186–189. Paramere in dorsal outline fairly wide but consistently converging to an acute tip, apically subequal in length to median lobe and narrower; slightly dorsally convex in lateral outline. Sensory spinules (very few) restricted to tip. Median lobe with broad truncate or slightly emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Among the golden-orange species N. nigroscutellatus and N. gebieni can by identified by the very dense and deep punctuation of the pronotum; this is also found in N. aeneiceps, which differs from the other two species by the metallic green anterior part of the head (golden-orange in N. nigroscutellatus and N. gebieni). N. gebieni can be distinguished from N. nigroscutellatus by the golden-orange scutellum (dark brown to black in N. nigroscutellatus) and by the shape of sternum VIII in males (Figs. 83, 86). Distribution. Known from elevations of 220–1480 m in Bolivia, Brazil, Ecuador, and Peru (Fig. 59). Habitat. Unknown. Remarks. Collected in flight intercept, malaise, and feces traps. Material. Lectotype, ‚, here designated, with labels ‘Jatahy (Brazil) A. Fauvel vend. 20.iv.1904’, ‘gebieni Bernh. Typus mus. Hamburg’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Lectotype Brachydirus gebieni Bernhauer des. Chatzimanolis 2001’ (in FMNH). – Paralectotype, „ with labels ‘Jatahy (Brazil) A. Fauvel vend. 20.iv.1904’, ‘gebieni Bernh. Cotypus mus. Hamburg’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Paralectotype Brachydirus gebieni Bernhauer des. Chatzimanolis 2001’ (in FMNH). – BOLIVIA: Cochabamba Prov., Cochabamba, 109 km E Yungas, Cochabamba-Villa Tunari Rd., 1480 m, 17o8´50´´S 65o42´29´´W, 1–6.ii.1999, R. Hanley leg. (1 in SEMC); Coroico (1 in BMNH); Santa Cruz, 9.5 km N Tatarenda, 25.xii.1993, D. Brzoska leg. (1 in SEMC). ECUADOR: Cachabe low, xii.1896, Rosenberg leg. (1 in FMNH); Napo, Yasuni Biological Station, 220 m, 0o40´12´´S 76o23´24´´W, 18–22.v.996, P. Hibbs leg. malaise trap (1 in SEMC); Napo, Yuturi Lodge, Rio Napo, 270 m, 0o32´54´´S 76o2´18´´W, 20– 22.iii.1999, R. Brooks & D. Brzoska leg. FIT (8 in SEMC); Napo Pr., Limoncocha, vii.1979, T. Ray leg. human dung trap (1 in MCZC); Napo, Tiputini Research station, 0o38´0´´S 76o9´0´´W, 220 m, 5–25.ix.2000, D.J. Inward & K.A. Jackson leg. (1 in BMNH); Napo Prov., Pto. Misahualli, 350 m, ii.1983, M. Sharkey leg. (1 in CNCI); Pastaza Liandia, 17 km N Puyo, 1000 m, 19.vii.1994, F. Genier leg. feces trap (1 in SEMC); Sucumbios, Sacha Lodge, 270 m, 0o28´14´´S 76o27´35´´W, 21–24.iii.1999, R. Brooks leg. FIT (16 in SEMC); Sucumbios, Sacha Lodge, 270 m. 0.5oS 76oW, iii–vi.1994, Hibbs leg. malaise trap (6 in SEMC). PERU: Loreto Dept., 1.5 km N Teniente Lopez, 2o35´66´´S 76o06´92´´W, 26.vii.1993, 210– 240 m, R. Leschen leg. FIT (1 in SEMC); Madre de Dios, Cocha Cashu Biol. Stn., Manu National Park, 350 m, 11o53´45´´S 71o24´24´´W, 17–19.x.2000, R. Brooks leg. FIT (8 in SEMC); Madre de Dios, Pentiacolla Lodge, Alto Madra

27


de Dios River, 400 m, 12o39´22´´S 71o13´55´´W, 23–28.x.2000, R. Brooks leg. FIT (3 in SEMC); 50 mi S Tingo Maria, Carpish Crest, 28.xii.1954, E.I. Schlinger & E.S. Ross leg. (1 in CASC).

4.20. Nordus longipes (Sharp, 1876) (Fig. 51) Brachydirus longipes Sharp, 1876: 115. Nordus longipes (Sharp); HERMAN (2001: 3580). Description. L e n g t h . 8.5–10 mm. C o l o r a t i o n . Head golden-orange with anterior 2/3 of epicranium shining black; mandibles and labrum light brown; antennomeres light brown to black. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/3 of VII goldenorange. Legs brownish-orange (yellow in one specimen). H e a d . Microsetae brown. Anteclypeus very small. Punctures in anterior half of epicranium moderately dense, about 5.3 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 3.6 punctures / 0.3 mm, punctures almost confluent. Length of eyes ca. 0.8ⴛ length of head, distance between eyes ca. 1ⴛ length of eye. Middle of epicranium strongly convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci are widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 51): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–4th longer than greatest width, 5th and 6th quadrate, 7th subquadrate, 8th–10th wider than greatest length, width of distal margin of 9th and 10th at least 1.5ⴛ length; 7th–11th densely covered by microtrichae, 6th with very few microtrichae, 4th–10th with single circumferential row of prominent macrosetae near middle, 1st–3rd with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.00; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1.5–2 punctures); punctures flanking this area in at least 3 dense, contiguous longitudinal rows, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and narrow impunctate area along posterior and posterolateral margin. Pronotum with brown microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with medium-sized punctures, about 4.5 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with brownishyellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with microsetae on medial disc; V– VII extremely punctuated, slightly less so medially; V–VII with yellow microsetae laterally and brown iridescent ones medially. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; III–IV and VIII with yellow microsetae. Segment VII without black iridescent setae. Cerci slightly curved, basal half brown and distal half yellow.

S e c o n d a r y s e x u a l s t r u c t u r e s . Male: unknown. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . Males unknown. Diagnosis. N. longipes is very similar to N. amphivolos and N. stomachoponos. Only these species are golden-orange with the anterior 3/4 of the head shining black and have antennomeres 2–10 light brown to black. N. stomachoponos can be distinguished from the other two by the very shallow and sparse punctuation on the pronotum, the very dense punctuation on the anterior part of the head, and antennomere 7 being wider than long (subquadrate in the other two species). In N. longipes antennomeres 5 and 6 are quadrate, while in N. amphivolos they are longer than wide; in N. longipes the last antennomere is brown-black while in N. amphivolos it is pale yellow. Distribution. Known from lowland forest in Brazil (Fig. 59). There are no specimens known from Bolivia as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Habitat. Unknown. Remarks. Collected in flight intercept and/or dung pitfall traps. Material. Holotype, „ with labels: on specimen card box ‘Brachydirus longipes Type Amazon D. S. „’, below ‘Type’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Pará’ (in BMNH). SHARP (1876) specifically states in the original description that he had available only a single female specimen, which is therefore the holotype. – BRAZIL: Para, Baker leg. (1 in FMNH); Para, Redencao vic. Keyapo Territory Pinkaiti R., xi.1998, tropical evergreen, seasonal lowland forest on clay soil, P.Y. Schaffer leg. dung pitfall or FIT (5 in FMNH).

4.21. Nordus maculiceps (Sharp, 1876) (Figs. 37, 84, 107, 190–193) Brachydirus maculiceps Sharp, 1876: 109. Brachydirus schuberti Wendeler, 1926: 74; new synonym. Nordus maculiceps (Sharp); HERMAN (2001: 3580). Nordus schuberti (Wendeler); HERMAN (2001: 3581). Description. L e n g t h . 9.5–12 mm. C o l o r a t i o n . Head black with two large red dots at middle of epicranium; mandibles and labrum orange; antennomeres (Fig. 37) all golden-orange. Pronotum, elytra, and scutellum black. Abdomen black except segment VIII and posterior 1/3 of VII golden-orange; in some specimens segment VII completely orange to light brown. Front and middle legs orange except coxae dark brown to black; hind legs dark brown to black except femora light brown; in some specimens (both sexes) all legs completely black except protarsi, protibiae, anterior part of profemora, and anterior parts of mesofemora dark orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior 3/4 of epicranium dense, about 4.5 punctures / 0.3 mm, punctures elongate longitudinally, margins of punctures confluent; in posterior 1/4 about 3.1 punctures / 0.3 mm, punctures almost contiguous. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micro-punctures. Gula narrow near midlength (least width : length = 0.09), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres: 1st gradually thickened club-like, but narrower near distal

28

end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–5th longer than greatest width, 6th almost quadrate, 9th and 10th almost quadrate; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.04; anterolateral angles not prominent, subcariniform; lateral margins almost parallel to each other in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures large and shallow; medial impunctate area of equal width throughout (width 2–3 punctures); punctures flanking this area in 1–2 sparse longitudinal rows separated by a distance as wide as 1 puncture, punctures of a row not contiguous, but distance varies; large, slightly convex, not raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayish-yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with mediumsized punctures, about 3.6 punctures / 0.3 mm, and interspaces as wide as 1 puncture, covered with grayish-yellow microsetae. Legs: fore and middle legs with yellow macrosetae except mesotibiae and -tarsi with black ones; hind legs with dark brown macrosetae except metacoxae, metatrochanters and anterior parts of metafemora with orange ones. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII extremely punctuated; IV–V and VIII covered with yellow microsetae, VI and VII with yellow and black ones. Paratergites III–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: III–VII densely punctuated; III–VI and VIII with yellow microsetae, VI and VII with yellow and black ones. Segment VII with black iridescent microsetae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 84) with evenly curved ventral surface, hind margin with very deep V-shaped notch medially; sternum IX (Fig. 107) with two long apical processes, hind margin without emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 190–193. Paramere in dorsal outline almost parallel-sided from middle to subapical region, then narrowed to a subacute tip, apically subequal in length to median lobe and narrower; completely straight in lateral outline. Sensory spinules (few) arranged in two irregular longitudinal bands close to midline and restricted to tip. Median lobe with narrowed subacute apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Among species with black head, pronotum and elytra, Nordus maculiceps and N. xanthocerus can be recognized by the shape of sternum VIII: in females it is evenly rounded and not strongly produced at its hind margin (as in Fig. 114), while in all other black species it is either sinuate (as in Fig. 118) or more strongly produced into


a lobe (as in Fig. 121); in males sternum VIII has a very deep V-shaped notch (as in Figs. 84, 103), while in all other black species it is as in Figs. 87, 88, 92 or 93. N. maculiceps is very similar to N. xanthocerus but can be distinguished from it by the presence of two bright red spots on the middle of the epicranium and the almost parallel lateral margins of the pronotum (margins concave in N. xanthocerus); also, in N. xanthocerus the pronotum is more densely punctuated than in N. maculiceps. Synonymy. Brachydirus schuberti, known from only one female specimen, is identical to B. maculiceps in all respects. Distribution. Known from elevations of 5–1800 m in Bolivia, Brazil, Colombia, Ecuador, Guyana, Peru, and Suriname (Fig. 58). Habitat. Unknown. Remarks. Collected in carrion traps, dung traps, flight intercept traps, and malaise traps. Material. Lectotype of B. maculiceps, ‚, here designated, with labels ‘Type’, ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus maculiceps ‚ Type D. S.’, ‘Lectotype Brachydirus maculiceps Sharp des. Chatzimanolis 2001’ (in BMNH). – Paralectotypes of B. maculiceps, 7 specimens (in BMNH): 1„ with labels ‘Amazon’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus maculiceps „ 2nd Type D. S. Amazon’, ‘Paralectotype Brachydirus maculiceps Sharp des. Chatzimanolis 2001’. 1 specimen with labels ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus maculiceps „ 2nd Type D. S. Amazon’, ‘Paralectotype Brachydirus maculiceps Sharp des. Chatzimanolis 2001’. 1 specimen with labels ‘ Peru’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus maculiceps var. Peru’, ‘Peru’, ‘Paralectotype Brachydirus maculiceps Sharp des. Chatzimanolis 2001’. 1 specimen with labels ‘S. Paulo Amazon’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus maculiceps var. amazons’, ‘Paralectotype Brachydirus maculiceps Sharp des. Chatzimanolis 2001’. 1 specimen with labels ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus maculiceps var. Amazon’, ‘Paralectotype Brachydirus maculiceps Sharp des. Chatzimanolis 2001’. 1 specimen with labels ‘Amazon’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘„’, ‘Brachydirus maculiceps 2nd Type D. S. Amazon’, ‘Paralectotype Brachydirus maculiceps Sharp des. Chatzimanolis 2001’. 1 specimen with labels ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus maculiceps’, ‘Paralectotype Brachydirus maculiceps Sharp des. Chatzimanolis 2001’. – Holotype of B. schuberti, „ with labels ‘Brachydirus n. sp’, ‘Oberer Madre de Dios, Peru 500 m’, ‘Brachydirus schuberti n. sp. Wend. Typus!’ (in ZMHB). WENDELER (1926) states in the original description that he had available only a single specimen, which therefore is the holotype. – BOLIVIA: Chapare, 400 m, 27.viii.1947, R. Zischka leg. (2 in NMW); Cochabamba, Cristalmayu, 17.xi.1949, L. Pena leg. (1 in NMW); Cochabamba, Cristalmayu, 30.ix.1949, L. Pena leg. (2 in NMW); Cochabamba, Chapare Villa Tunari, xii.1976, Bolle leg. (1 in CNCI); Cochabamba Prov., Cochabamba, 67.5 km NE Est. Biol. Valle de Sajita of Univ. de San Simon, 300 m, 17o6´33´´S 64o47´52´´W, 9–13.ii.1999, R. Hanley leg. FIT (13 in SEMC); Cochabamba Prov., Cochabamba, 67.5 km NE Est. Biol. Valle de Sajita of Univ. de San Simon, 300 m, 17o6´33´´S 64o47´52´´W, 9–13.ii.1999, F. Genier leg. FIT (29 in SEMC); Coroico (1 in AMNH, 1 in BMNH, 1 in DEI, 8 in FMNH, 1 in NMW); Mapiri (2 in FMNH, 2 in NMW); La Paz, Atto Bani, Palos Blancos, 600 m, 11.i.1976, L.E. Pena leg. (3 in FMNH); Coroico, 1800 m, 10.xii.1906, Gerloff & Fassl leg. (1 in FMNH); Farinas (1 in BMNH); Rio Juntas (1 in BMNH); Rio Longo, 750 m, Fassl leg. (1 in FMNH); Yungas de la Paz, 1000 m, H. Rolle leg. (1 in ZMHB); Bang Haas leg. (1 in AMNH); Karl Brancik leg. (1 in AMNH); (no labels) (5 in FMNH). BRAZIL: Amazon, Kraatz leg. (4 in DEI); Amazon, Bates leg. (3 in BMNH); Amazons (2 in BMNH); Amazons Nanta (1 in BMNH); Amazonias, xi.1923, Bierig leg. (1 in


FMNH); Ega (1 in FMNH); Guanabara, Rio de Janeiro iii.1969, M. Alvarenga leg. (1 in AMNH); Repress de Rio Grando, iii.1970, M. Alvarenga leg. (2 in AMNH); Para, Aldeia Aracu, Igarape Gurupumu Maranhao, 50 km E of Caninde, v.1963, B. Malkin leg. (4 in FMNH); Para, Aldeia Coraci, 12 km W of Caninde, R. Gurupi, v.1963, B. Malkin leg. (2 in FMNH); Para, Aldeia Maracacume Maranhao 80 km E of Caninde, 22–23.v.1963, B. Malkin leg. (1 in FMNH); Para, Caninde, Rio Gurupi 7–15.iv.1963, B. Malkin leg. forest sweep (1 in FMNH); Para, Caninde, Rio Gurupi, 30.iii.1963, B. Malkin leg. beating leaf forest debris (1 in FMNH); Para, Redencao vic. Keyapo Terittory Pinkaiti R., xi.1998, tropical evergreen seasonal lowland forest on clay soil, P.Y. Schaffer leg. (1 in FMNH); Para, Baker leg. (1 in FMNH); Rondonia, Fazenda Rancho grande, xi.1991, S. Passos leg. (2 in FMNH); Sinop, Mato Grosso, x.1975, M. Alvarenga leg. (3 in CNCI); Sinop, Mato Grosso, x.1976, M. Alvarenga leg. (1 in CNCI); Umarituba, Rio Negro (1 in FMNH). COLOMBIA: Meta PNN, Tinigua Vda. Bajo Raudal, 2o16Ń 73o48´W, 390 m, 21.xii. 2001, D. Campos leg. (1 in SEMC); Putomayo, Santa Rosa de Sucumbios, Rio San Miguel, 400 m, 1–7.ix.1971, B. Malkin leg. forest trail beating rotten foliage (1 in FMNH); Villavicencio ost Colombia, 450 m, iv.1910 (2 in NMW). ECUADOR: Coca, v.1965, L. Pena leg. (2 in FMNH); Napo Prov., Anaconda Lodge, Napo River, 15.ii.1991, D. Brzoska leg. (1 in SEMC); Napo Prov., Napo R. 1 mi E Anvano, Anaconda Is. 14.ii.1991, J. Stamatov leg. (1 in AMNH); Napo Prov., Limoncocha, 250 m, 15–28.vi.1976, S. & J. Peck leg. (1 in FMNH); Napo, 25 km E of Puerto Napo Selva Alinahui, 450 m, xii.1992, E.S. Ross leg. (1 in CASC); Napo Prov., Pto. Misahualli, 350 m, ii.1983, M. Sharkey leg. (1 in CNCI); Sucumbios, Sacha Lodge, 0.5oS 76.5oW, 270 m, 3–13.iv.1994, P. Hibbs leg. malaise trap (2 in SEMC). FRENCH GUIANA: 3.5 mi N Saul, Les Eaux Claires, 3.xi.1995, A. Berkov leg. (1 in AMNH); 3.5 mi N Saul, Les Eaux Claires, 14.iv.1996, C. Chaboo leg. by hand (1 in AMNH). GUYANA: British Guiana [Guyana], Mazaruni High Forest, 21.viii.1937, Richards & Smart leg. (1 in BMNH, 1 in FMNH); (no labels) (1 in FMNH). PERU: Callanga (3 in NMW); Pachitea (2 in FMNH); Junia, San Ramon de Pangoa, 40 km S Satipo, 750 m, 26.i.1974, R.T. Schuh leg. (1 in AMNH); Loreto Dept. 1.5 km N Teniente Lopez, 2o35´66´´S 76o06´92´´W, 210–240 m, 18.vii.1993, R. Leschen leg. FIT (13 in SEMC); Madre de Dios, Coche Salvador Reserved Zone, Manu national park, 310 m, 12o0´13´´S 71o31´36´´W, 20–21.x.2001, R. Brooks leg. FIT (2 in SEMC); Tambopata Prov., 15 km NE Pto. Maldonaldo, 200 m, 26.vi.1989, R. Leschen leg. malaise trap (3 in SEMC); Tingo Maria Huan., 20.v.1947, 2200 ft [733 m], J.C. Pallister leg. (3 in AMNH); Monsou Valley Tingo Maria, 2.xii.1954, E.I. Schlinger & E.S. Ross leg. (9 in CASC); Vilcanota, Kraatz leg. (2 in DEI). SURINAME: Marowijne Perica, 70 km E Paramaribo on E-W Rd., 5 m, 5o40´28´Ń 54o36´31´´W, 29–31.v.1999, Z. Falin & B. DeDijin leg. FIT (1 in SEMC); Kraatz leg. (1 in DEI).

4.22. Nordus major (Bernhauer, 1905) (Figs. 16, 38, 85, 194–197) Brachydirus major Bernhauer, 1905: 186. Nordus major (Bernhauer); HERMAN (2001: 3580). Description. L e n g t h . 11.5–13.5 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres all golden-orange. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/6 of VII golden-orange. Legs golden-orange. Head. Microsetae yellow on anterior 3/4 of epicranium and black on posterior 1/4. Anteclypeus well developed. Punctures in anterior 3/4 of epicranium dense, about 4.5 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior 1/4 about 3.6 punctures / 0.3 mm, punctures almost contiguous. Middle of epicranium convex; epicranium without microsculpture of transverse microlines and micropunctures. Length of eyes ca. 0.6ⴛ

29

length of head, distance between eyes less than 1.5ⴛ length of eye. Eyes positioned far anteriorly on head (rather than around middle), temporal area posterior to eyes greatly expanded, strongly convex. Gula very narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and posteriorly and almost contiguous near midlength. Antennomeres (Fig. 38): 1st gradually thickened club-like, but narrower near distal end, shorter than 2nd plus 3rd, 2nd shorter to 3rd, 5th subequal in length to 4th, 1st–5th longer than greatest width, 6th almost quadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. A particular point of intraspecific variation only found in N. major is that the size of the head in relation to body size varies dramatically among specimens. T h o r a x . Pronotum somewhat wider than long, width : length = 1.14; anterolateral angles extremely prominent and acutely delimited, forming angle of about 90°, cariniform; lateral margins of pronotum strongly convex in dorsal aspect; punctures large and shallow; medial impunctate area present, of uniform width throughout (width 2–3 punctures); otherwise punctuation uniform, hence disc laterally without distinct gaps in punctuation, no discrete rows of punctures flanking medial impunctate area, and no impunctate areas in anterolateral quarters and along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with 1–2 rows of punctures anteriorly. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with large punctures, about 4.5 punctures / 0.3 mm, punctures nearly contiguous, covered with yellow microsetae, longer black macrosetae along margins. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII extremely punctuated; IV–VII covered with yellow microsetae laterally and few black ones medially. Paratergites III–VII with whitish-yellow microsetae throughout. Sterna: III–VII densely punctuated; III–VII with yellow microsetae. Segment VII with black iridescent setae throughout. Cerci slightly curved, basal half light brown and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 85) with evenly curved ventral surface, hind margin with a broad and moderately deep U-shaped emargination medially; sternum IX (identical to that in Fig. 110) with two moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 194–197. Paramere in dorsal outline very wide but with a very narrow base, with slightly convex lateral margins hardly converging distally, tip broadly rounded, apically subequal in length to median lobe and wider; almost completely straight in lateral outline. Sensory spinules (many) arranged in a pair of far lateral rows extending from tip 1/3 of paramere length towards paramere base, some additional spinules ventrally between rows. Median lobe strongly narrowed distally and

30

with acute apex in dorsal outline, hardly upcurved apically in lateral outline, without dorsomedian carina and without pair of dorsal lobes. Diagnosis. Nordus velitaris, N. fungicola, and N. major can be distinguished from other species of Nordus by the combination of a very convex epicranium and antennomere 4 being longer than wide. N. fungicola and N. major differ from N. velitaris by the convex rather than concave lateral margins of the pronotum. Furthermore, the temporal area of the head is strongly expanded in N. major but only slightly so in N. fungicola; a pair of raised anterolateral impunctate areas on the pronotum is present in N. fungicola but absent in N. major; the antyclepeus is reduced in N. fungicola but well developed in N. major. Distribution. Known from elevations of 210–700 m in Bolivia, Brazil, Ecuador, and Peru (Fig. 59). Habitat. Unknown. Remarks. Collected in flight intercept and malaise traps. Material. Lectotype, ‚, here designated, with labels ‘Pachitea Peru’, [white] ‘major Bernhauer Typus’, [yellow] ‘major Bernhauer Typus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Lectotype Brachydirus major Bernhauer des. Chatzimanolis 2003’ (in FMNH). – Paralectotype, „ with labels ‘Mapiri Bolivia’, [yellow] ‘major Bernhauer cotypus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Paralectotype Brachydirus major Bernhauer des. Chatzimanolis 2003’ (in FMNH). – BOLIVIA: Mapiri (1 in BMNH). BRAZIL: Itatiaya, Est. do. Bio., 700 m, 16.ii.1924, J.F. Zikan leg. (1 in FMNH); S. Catharina, Rio Capivary, 1888, Fruhstorfer leg. (8 in BMNH, 8 in DEI, 3 in FMNH, 4 in NMW). ECUADOR: Limoncocha, 11.iii.1976, J.M. Campell leg. (1 in CNCI); Limoncocha, 31.iii.1974, 300 m, H.P. Stockwell leg. (1 in FMNH); Napo, Yasuni Biological Station, 220 m, 0o40´12´´S 76o23´24´´W, 18–22.v.1996, P. Hibbs leg. malaise trap (1 in SEMC); Sucumbios, Sacha Lodge, 270 m, 0o28´14´´S 76o27´35´´W, 21–24.iii.1999, R. Brooks leg. FIT (1 in SEMC). PERU: Loreto Dept., 1.5 km N Teniente Lopez, 2o35´66´´S 76o06´92´´W, 210–240 m, 18.vii.1993, R. Leschen leg. FIT (6 in SEMC); Marcapata, Bierig leg. (1 in FMNH); Marcapata (3 in NMW).

4.23. Nordus nigroscutellatus (Bernhauer, 1905) (Figs. 39, 86, 198–201) Brachydirus nigroscutellatus Bernhauer, 1905: 185. Brachydirus chapareanus Scheerpeltz, 1971: 103; new synonym. Nordus nigroscutellatus (Bernhauer); HERMAN (2001: 3580). Nordus chapareanus (Scheerpeltz); HERMAN (2001: 3579). Description. L e n g t h . 8.0–10 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres all yellow. Pronotum and elytra golden-orange, scutellum dark brown to black. Abdomen black except segments I, II, VIII, and posterior 1/3 of VII goldenorange. Legs golden-orange; in some specimens middle legs light brown, distal halves of metafemora and -tibiae light brown to brown. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 5.3 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 4.1 punctures / 0.3 mm, punctures almost contiguous. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 39): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal


in length to 3rd, 5th shorter than 4th, 1st–4th longer than greatest width, 5th subquadrate, 6th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th– 10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 0.96; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in at least 2–3 dense, contiguous longitudinal rows, punctures of a row conti-guous; large, convex, raised impunctate area in anterolateral quarters, and small impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with whitishyellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with large punctures, about 3.5 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae, except metatibia with brown ones. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with yellow microsetae medially and posterolaterally; V densely punctuated on medial disc but with fewer punctures medially on posterior margin,; VI densely punctuated except medially; V with yellow microsetae throughout, VI with yellow ones laterally and brown ones medially, VII with dark brown ones, VIII with yellow ones. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–VI and VIII with yellow microsetae. Segment VII with black iridescent setae throughout. Cerci slightly curved, basal half light brown and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 63) with broad and shallow emargination medially; sternum VIII (Fig. 86) with evenly curved ventral surface, hind margin with deep Vshaped notch medially; sternum IX (identical to that in Fig. 111) with two moderate apical processes and a moderate Ushaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig. 114). M a l e g e n i t a l i a . As in Figs. 198–201. Paramere in dorsal outline fairly wide but consistently converging to an acute tip, apically subequal in length to median lobe and narrower; slightly dorsally convex in lateral outline. Sensory spinules (very few) restricted to tip. Median lobe with broad truncate or slightly emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, for-ming a dorsomedial carina forked apically into two short lobes. Diagnosis. Among the golden-orange species N. nigroscutellatus and N. gebieni can by identified by the very dense and deep punctuation of the pronotum; this is also found in N. aeneiceps, which differs from the other two species by the metallic green anterior part of the head (golden-orange in N. nigroscutellatus and N. gebieni). N. gebieni can be distinguished from N. nigroscutellatus by the golden-orange scutellum (dark brown to black in N. nigroscutellatus) and

31


by the shape of sternum VIII in males (Figs. 83, 86); male genitalia are very similar in these two species (Figs. 186– 189, 198–201). Synonymy. Brachydirus nigroscutellatus and B. chapareanus are identical in every respect. Distribution. Known from elevations of 300–1475 m in Bolivia and Peru (Fig. 59). Habitat. Unknown. Remarks. Collected in flight intercept, carrion and feces traps. Material. Holotype of B. nigroscutellatus, „ with labels ‘Mapiri Bolivia’, ‘nigroscutellatus – Typus Bang-Haas’, ‘Chicago NHMuseum M. Bernhauer Collection’ (in FMNH). BERNHAUER (1905) states in the original description that he had available only a single specimen, which therefore is the holotype. – Holotype of B. chapareanus, ‚ with labels ‘‚’, ‘Bolivia Chapara 400 m leg. H. Zischka’, ‘Bolivia Chapara 400 m Zischka’, ‘8.ii.1948’, ‘Brachydirus chapareanus n.sp.m.’, ‘ex. coll. Scheerpeltz’, ‘Typus Brachydirus chapareanus O. Scheerpeltz’ (in NMW). SCHEERPELTZ (1971) explicitely mentioned that there was only one specimen for his description. However, in his collection there are two specimens, one labeled ‘Typus’ (the specimen above) and another, female one labeled ‘Cotypus’. The latter specimen was not mentioned in the original description and therefore is not a valid paratype for this species. – BOLIVIA: Cochabamba, 67.5 km NE Est. Biol. Valle de Sajita of Univ. de San Simon, 300 m, 17o6´33´´S 64o47´52´´W, 7–9.ii.1999, F. Genier leg. FIT (7 in SEMC); same collecting data, R. Hanley leg. FIT (1 in SEMC); Cochabamba, 124 km E Yungas, Cochabamba–Villa Tunari Rd., 730 m, 17o3´54´´S 65o38´43´´W, 1–6.ii.1999, R. Hanley leg. FIT (9 in SEMC); same collecting data, F. Genier leg. carrion/feces trap (4 in SEMC); Cochabamba, 117 km E Yungas, Lagunitas, 1000 m, 17o6´22´´S 65o40´57´´W, 6.ii.–8.vi.1999, F. Genier leg. FIT (2 in SEMC); Mapiri (2 in FMNH). PERU: Pasco Dept., Villa Rica Rd., 1475 m, 10o47´5´´S 75o18´54´´W, 15–18.x.1999, R. Brooks & D. Brzoska leg. FIT (2 in SEMC); Ucayali Dept., Tingo Maria-Pucallpa Rd., Puente Chino, km 205, 1300 m, 9o8´12´´S 75o47´20´´W, 11–14.x.1999, R. Brooks leg. FIT (2 in SEMC).

4.24. Nordus omios sp.n. (Figs. 40, 64, 87, 202–205) Description. Exactly as for the black morph of N. terminalis with the following differences: H e a d . Distance between punctures in posterior 1/4 of epicranium as wide as 1.5 punctures. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 64) with broad and very deep emargination medially; sternum VIII (Fig. 88) with evenly curved ventral surface, hind margin with single broad, long lobe and deep, narrow, parallel-sided median cleft, with slightly expanded brush-like patch of prominent setae on basal part of each side of cleft; sternum IX (Fig. 108) with two long apical processes and a deep U-shaped emargination medially. M a l e g e n i t a l i a . As in Figs. 202–205. Paramere in dorsal outline very slender but with distinctly widened, disclike part around middle, distally slightly narrowed to a subacute tip, apically subequal in length to median lobe and narrower; altogether slightly dorsally convex in lateral outline, but with abrupt dorsal bend near middle (at disc) and with hook-like apex. Sensory spinules (many) arranged in a distal medial cluster and two regular longitudinal rows more basally, all restricted to apical 1/10 of paramere. Median lobe with broadly rounded apex in dorsal outline, slightly upcurved apically in lateral outline, forming apically two widely separated, short dorsal lobes but no dorsomedial carina basal to these.

Diagnosis. Nordus terminalis, N. omios, N. solitarius, and N. dichromos can be distinguished from all other Nordus species by the lacking (or extremely reduced) microsetation on the medial disc of abdominal tergum IV. These setae are also absent in N. fungicola, which differs from the above species by its convex rather than concave lateral margins of the pronotum. N. terminalis and N. omios can be distinguished from the other two species by the depressed elytra near scutellum, the very sparse punctuation on the elytra (distance between punctures larger than width of 1 puncture versus less than 1 in the other two species) and, in the males, by their very distinctive shape of the paramere (disc-like middle surrounded by very slender parts; Figs. 202, 241) and sternum VIII (one large lobe on hind margin, with deep, narrow medial cleft, and brush-like patch of prominent setae at anterior end of cleft; Figs. 87, 88). N. terminalis and N. omios differ only in the shapes of the aedeagus and male sternum VIII: lobe of sternum VIII slightly shorter in N. omios; paramere (lateral view, Figs. 203, 242) hook-like apically in N. omios, but almost straight apically in N. terminalis; sensory spinules extending further towards paramere base in N. terminalis than in N. omios; different shape of apex of median lobe. Distribution. Known from elevations of 500–1450 m in Darien, Panama (Fig. 68). Habitat. Unknown. Derivatio nominis. The specific epithet is derived from the Greek word ‘ó␮␱␫␱␵’ (same) and refers to the similarity between N. terminalis and N. omios. Remarks. Collected in flight intercept traps and human dung traps. In SEMC, there are two females from Coclé, Panama, and four females from Cerro Campana (Capira), Panama, that are probably N. omios but they cannot be reliably distinguished from females of N. terminalis. Material. Holotype, ‚ with labels ‘Panama: Darien, Cana Biological Station, Serrania de Pirre, 1450 m, 7o48´18´Ń 77o41´6´´W, 4–7.vi.1996, J.S. Ashe, R. Brooks, PAN1A96 108 ex: FIT’, barcode label SM0034293, ‘HOLOTYPE Nordus omios Chatzimanolis des. Chatzimanolis 2003’ (in SEMC). – Paratypes, 13 specimens: Same locality and collector data, barcode label SM0034287 (1 in SEMC). Same locality name and collector but 1200 m, 7o45´18´Ń 77o41´6´´W, barcode labels SM0013834, SM0013833, SM0021494, SM0021490, SM0013835, SM0013837, SM0021498, SM0034288 (8 in SEMC). Same locality name and collector, 1250–1390 m, 7o45´18´Ń 77o41´6´´W, 5.vi.1996, barcode labels SM0013866, SM0013867 (2 in SEMC). Same locality name and collector but 1380 m, 7–9.vi.1996, 7o45´18´Ń 77o41´6´´W, barcode label SM0034301 (1 in SEMC). ‘Panama: Darien, 500 m, Estacion Ambiental Cana, 7o45´20´Ń 77o41´6´´W, 6–7.vi.1996, R.S. Anderson, PAN2A96 96-109, ex: human dung trap’, barcode label SM0034536 (1 in SEMC). All paratypes with label ‘Paratype Nordus omios Chatzimanolis des. Chatzimanolis 2003’.

4.25. Nordus periergos sp.n. (Figs. 52, 96, 206–209) Description. L e n g t h . 8.0–9.0 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres all yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior half of VII golden-orange. Legs golden-orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 5.3 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior half about 3.1 punctures /

32

0.3 mm, punctures almost confluent. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium with microsculpture of transverse microlines but without micropunctures. Gula narrow near midlength (least width : length = 0.06), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 52): 1st thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate, 6th–10th wider than greatest length, width of distal margin of 9th and 10th 1.5ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 0.94; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in at least 2 dense, contiguous longitudinal rows, punctures of a row contiguous; large, slightly convex, raised impunctate area in anterolate-ral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering the posterior 4/5. Elytra not depressed near scutellum, with large punctures, about 3.6 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII well punctuated (V–VI less so medially); IV–VI with yellow microsetae medially and posterolaterally, VII with brown ones. Paratergites IV–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–VI and VIII with yellow microsetae, VII with brown ones. Segment VII with black iridescent microsetae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 96) with evenly curved ventral surface, hind margin with two very large triangular lobes and a deep V-shaped emargination medially; sternum IX (identical to that in Fig. 104) with two small to moderate apical processes and a moderate U-shaped emargination medially. Female: unknown. M a l e g e n i t a l i a . As in Figs. 206–209. Paramere in dorsal outline abruptly narrowed near middle, almost parallelsided basal and distal to middle, distally forming a subacute tip, apically subequal in length to median lobe and narrower; almost completely straight in lateral outline. Sensory spinules (many) arranged in two longitudinal rows placed near midline and restricted to the apical and subapical areas. Median lobe slightly widened distally, with broadly subacute apex in dorsal outline, quite strongly upcurved in distal half in lateral outline, forming a small dorsomedial carina forked apically into two short lobes. Diagnosis. Among species with golden-orange head, pronotum and elytra, N. densiventris, N. semelius, and N. periergos are unique in the presence of microsculpture of


transverse microlines on the epicranium. N. densiventris is very similar to N. semelius and N. periergos but differs in the shape of sternum VIII of males (Figs. 78, 96, 100): two very small triangular lobes extending beyond the posterior sternal margin and a moderate U-shaped medial emargination in N. densiventris; two strong triangular lobes and a very deep V-shaped medial emargination in N. semelius and N. periergos. N. semelius and N. periergos differ in the aedeagi (Figs. 206, 222): e.g., in N. semelius the median lobe in dorsal view is distally emarginate, while in N. periergos it is distally narrowed to a broad subacute apex. Distribution. Known from elevations of 1000 m in Peru (Fig. 21). Habitat. Unknown. Derivatio nominis. The specific epithet is derived from the Greek word ‘␲⑀␳␫⑀␳␥␱␵’ (bizarre) and refers to the shape of the aedeagus and the 8th sternum. Material. Holotype, ‚ with labels ‘Chauchamayo, Peru, 8.viii.[19]48, J.M. Schunke’, ‘HOLOTYPE Nordus periergos Chatzimanolis des. Chatzimanolis 2003’ (in CASC). – Paratypes, 4‚‚: Same locality data and collector (2 in CASC). ‘1000 m, 11o8´S 75o17´W, Greenw. N. Iconnicoff, 3.ii.1906’, ‘densiventris det. Bernhauer’, ‘testaceus Peru, Mus. Moskov, det. Bernhauer’, ‘Chicago NHMus. M. Bernhauer Collection’ (2 in FMNH). All paratypes with label ‘Paratype Nordus periergos Chatzimanolis des. Chatzimanolis 2003’.

4.26. Nordus picticornis (Solsky, 1872) (Figs. 53, 97, 210–213) Brachydirus picticornis Solsky, 1872: 307. Nordus picticornis (Solsky); HERMAN (2001: 3581). Description. L e n g t h . 10.0 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres 1–4 yellow, 5–9 and distal part of 11 pale yellow, 10 and basal part of 11 dark brown to black. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/4 of VII golden-orange. Legs golden-orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 4.9 punctures / 0.3 mm, punctures elongate longitudinally, margins of punctures confluent; in posterior half about 3.6 punctures / 0.3 mm, distance between punctures as wide as 0.5 punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 53): 1st thickened only at distal half, but narrower near distal end, longer than 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate, 6th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 0.96; anterolateral angles not prominent, subcariniform; lateral margins strongly concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1– 1.5 punctures); punctures flanking this area in 1–2 sparse

33


longitudinal rows separated by a distance as wide as 0.5–1 punctures, punctures of a row not contiguous, but distance varies; large, convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayishyellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.6 punctures / 0.3 mm, and interspaces as wide as 0.5–1 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII well punctuated (V–VI less so medially); IV with yellow microsetae medially and posterolaterally, V with yellow ones throughout, VI with yellow ones laterally and brown ones medially, VII with brown ones. Paratergites IV–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: IV–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–VI and VIII with yellow microsetae, VII with brown ones. Segment VI–VII with black iridescent microsetae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 97) with evenly curved ventral surface, hind margin with two strong triangular lobes (tips pointed) and a moderate U-shaped emargination medially; sternum IX (identical to that in Fig.110) with two moderate apical processes and a moderate U-shaped emargination medially. Female: unknown. M a l e g e n i t a l i a . As in Figs. 210–213. Paramere in dorsal outline slightly but consistently converging to a narrowly rounded tip, apically slightly shorter than median lobe and narrower; completely straight in lateral outline. Sensory spinules (few) arranged in a medial distal cluster and some isolated spinules a bit more basally, all restricted to tip. Median lobe with narrowly rounded apex in dorsal outline, not upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two tiny lobes. Diagnosis. Nordus antennatus, N. banghaasi, N. picticornis, N. aeneipennis, and N. zischkai can all be distinguished from other species of Nordus by the following characters: punctures on anterolateral corners of sterna VI and VII less dense than on the rest of these sterna; punctures on anterior part of epicranium longitudinally elongate; and shape of sternum VIII in males (shape varies among these species, but overall is distinctive for this group of species). N. antennatus and N. banghaasi can be distinguished from N. picticornis, N. aeneipennis, and N. zischkai by antennomere 4 being longer than wide (subquadrate in other species). In N. picticornis head, pronotum and elytra are golden-orange, whereas in N. aeneipennis and N. zischkai head and pronotum are reddish-brown to black and elytra are dark brown to black. The very similar N. picticornis and N. aeneipennis are well distinguished by the shapes of the aedeagus (Figs. 210–213, 134–137) and sternum VIII (Figs. 97, 69). Distribution. Known from an elevation of 1300 m in Peru (Fig. 60). Habitat. Unknown. Remarks. Collected in flight intercept traps. The original type by Solsky appears to be lost. V. Gusarov and A. Solodovnikof have both checked the collections in St.

Petersburg and Moscow and suggest that the type should be considered lost. SOLSKY (1872) did not specify any type locality in his original description. The specimen designated here as neotype agrees in every respect with Solsky’s description. SOLSKY (1875) mentioned that he had seen some specimens of N. picticornis from Peru that exhibit dramatic color variation, however, based on his description of color variation, these were specimens of N. aeneipennis. These two species look very similar if someone ignores the color but they have a few differences in addition to those in coloration (distribution of punctures on the pronotum, shape of aedeagus, shape of sternum VIII; see diagnosis of N. aeneipennis). Material. Neotype, ‚, here designated, with label ‘Peru: Ucayali dept. Tingo Maria-Pucallpa Rd. Puente Chino, km 205, 1300 m, 9o8´12´´S 75o47´20´´W, 11–14.x.1999; R. Brooks PERU1B99 007A ex. flight intercept trap’, barcode label SM0135073, ‘NEOTYPE Brachydirus picticornis Solsky des. Chatzimanolis 2003’ (in SEMC).

4.27. Nordus portokalis sp.n. (Figs. 54, 98, 112, 214–217) Description. L e n g t h . 10.5–11.5 mm. C o l o r a t i o n . Head dark brown to black with two bright red dots at middle of epicranium; mandibles and labrum light brown; antennomeres 1–5 brown, 6–11 yellowish-brown. Pronotum dark brown to black, elytra brown, scutellum black. Abdomen dark orange to light brown. Legs orange except meso- and metatibiae and -tarsi black and protibiae brownish-orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 5.8 punctures / 0.3 mm, punctures elongate longitudinally, margins of punctures confluent; in posterior half about 2.3 punctures / 0.3 mm, distance between punctures as wide as 0.5–1 punctures, middle of posterior half without punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.03), gular sulci widely separated anteriorly and near midlength and almost contiguous posteriorly. Antennomeres (Fig. 54): 1st thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th subequal in length to 4th, 1st– 3rd longer than greatest width, 4th and 5th subquadrate, 6th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum longer than wide, width : length = 0.88; anterolateral angles not prominent, subcariniform; lateral margins strongly concave in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures medium-sized and shallow; medial impunctate area broadest anteriorly (width 2–3 punctures); punctures flanking this area in 1–2 sparse longitudinal rows separated by a distance as wide as 1.5–2.5 punctures, punctures of a row not contiguous, but distance varies; large, slightly convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming 2 rows, not covering entire width. Scutellum without

34

arch-like carina, with yellow microsetae and punctures covering posterior half. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, and interspaces as wide as 0.5–1 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V with subbasal carina; IV–VII well punctuated; IV–VIII with yellow microsetae medially and posterolaterally. Paratergites IV–VII covered with yellow microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–VII with yellow microsetae. Segment VII with black iridescent setae throughout. Cerci thin and slightly curved, basal 2/3 light brown and distal 1/3 yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: distal margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 98) with evenly curved ventral surface, hind margin with two moderate triangular lobes and a narrow and deep U-shaped emargination medially; sternum IX (Fig. 112) with very long apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII sinuate (identical to that in Fig. 118). M a l e g e n i t a l i a . As in Figs. 214–217. Paramere in dorsal outline slightly but abruptly narrowed near middle, almost parallel-sided basal and distal to middle (or even slightly expanding distally in basal part), distally forming an acute tip, apically subequal in length to median lobe and narrower; almost completely straight in lateral outline; with depression dorsomedially below tip. Sensory spinules (only 6) arranged in two longitudinal, basally diverging doublerows and restricted to tip. Median lobe strongly narrowed distally to subacute apex in dorsal outline, with tiny mediodistal projection, slightly upcurved in distal half in lateral outline, forming apically two long, widely separated dorsal lobes but no dorsomedial carina basal to these. Diagnosis. Nordus cherylae, N. elegans, N. portokalis, and N. speciosus can be distinguished from all other species of Nordus by the presence of a subbasal carina (in addition to the basal one) on abdominal tergum V. N. portokalis can be distinguished from the other three species by the presence of the very distinctive orange abdomen (black in the other species) and the subquadrate pronotum (pronotum longer than wide in other species). Distribution. Known from an elevation of 1700 m in Peru (Fig. 21). Habitat. Collected in flight intercept traps. Derivatio nominis. The specific epithet is derived from the Greek word ‘␲␱␳␶␱␬␣␭␫’ (orange) and refers to the color of the abdomen. Material. Holotype, ‚ with labels ‘PERU: Huánuco Dept. Cordillera Azul, 39 km NE Tingo María, 1700 m, trap site 672, 11–14.i.1983 montane rainforest, A. Newton & M. Thayer’, ‘window trap’, ‘Holotype Nordus portokalis Chatzimanolis des. Chatzimanolis 2003’ (in FMNH). – Paratypes, 1‚, 2„„ with same labels except paratype label ‘Paratype Nordus portokalis Chatzimanolis des. Chatzimanolis 2003’ (‚ in SEMC, „„ in FMNH).

4.28. Nordus seferisi sp.n. (Figs. 55, 99, 218–221) Description. L e n g t h . 8.5–9.0 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres yellow, 7–11 darker yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black


except segments I, II, VIII, and posterior 1/3 of VII goldenorange. Legs golden-orange. H e a d . Microsetae yellow on anterior 3/4 and black on posterior 1/4. Anteclypeus very small. Punctures on anterior 3/4 of epicranium dense, about 5.8 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior 1/4 about 3.1 punctures / 0.3 mm, punctures almost contiguous. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near mid-length (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 55): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate, 6th wider (basal margin) than greatest length and shorter than 5th, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 1.07; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc of pronotum laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1– 1.5 punctures); punctures flanking this area in at least 3 dense, contiguous longitudinal rows, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and small impunctate area along posterior and posterolateral margins. Pronotum with dark brown microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayish-yellow microsetae and punctures covering posterior 2/ . Elytra not depressed near scutellum, with large punctu3 res, about 4.0 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with grayish-yellow microsetae on medial disc; V densely punctuated on medial disc but less so medially on posterior margin; VI densely covered with punctures except medially; V–VI with grayish-yellow microsetae except medially, VII with dark brown ones, VIII with sparsely distributed yellow ones. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated, III–VI and VIII with yellow microsetae. Segment VII with black iridescent setae throughout. Cerci slightly curved, basal half light brown and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 63) with broad and shallow emargination medially; sternum VIII (Fig. 99) with evenly curved ventral surface, hind margin with broad and moderately deep U-shaped emargination medially; sternum IX (identical to that in Fig. 104) with two small to moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (identical to that in Fig.114).


M a l e g e n i t a l i a . As in Figs. 218–221. Paramere in dorsal outline very wide, almost parallel-sided from middle to subapical region, then strongly narrowed forming a subacute tip, apically subequal in length to median lobe; slightly dorsally concave basally and slighly dorsally convex distally in lateral outline. Sensory spinules (few) arranged in an irregular medial band and restricted to tip. Median lobe with broad emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus cribricollis, N. elytisi, and N. seferisi can be distinguished from all other golden-orange Nordus species by antennomere 6 being wider than long (subquadrate in all other species except N. portokalis, which by its orange abdomen differs from the three abovementioned species with black abdomen). N. cribricollis can be distinguished from N. seferisi and N. elytisi by antennomere 4 being longer than wide rather than subquadrate. In addition, N. cribricollis males have a paramere (Fig. 154) much narrower than in the other two species (Figs. 174, 218). N. elytisi differs from N. seferisi by the denser punctuation on the pronotum near the medial impunctate area, and males can be distinguished by the shape of the aedeagus (Figs. 174–177 and 222–225). Distribution. Known from elevations of 210–240 m in Peru (Fig. 21). Habitat. Unknown. Derivatio nominis. The specific epithet is in honor of Georgios Seferis, a Greek Nobel laureate. Remarks. Collected in flight intercept traps. Material. Holotype, ‚ with labels ‘Peru, Dept. Loreto Teniente Lopez, 2o35´66´´S, 76o06´92´´W, 18.vii.1993, 210– 240 m Richard Leschen #117 ex: flt. inter. Trap, Qd11’, barcode label SM0058051, ‘Holotype Nordus seferisi Chatzimanolis, des. Chatzimanolis 2003’ (in SEMC). – Paratype, „ with labels ‘Peru, Dept. Loreto Teniente Lopez, 2o35´66´´S, 76o06´92´´W, 20.vii.1993, 210–240 m Richard Leschen #133 ex: flt. inter. Trap, Qd11’, barcode label SM0058049, ‘Paratype Nordus seferisi Chatzimanolis, des. Chatzimanolis 2003’ (in SEMC).

4.29. Nordus semelius sp.n. (Figs. 7, 56, 100, 222–225) Description. L e n g t h . 9.0–10.5 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres (Fig. 56) all yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior half of VII golden-orange. Legs golden-orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 4.9 punctures / 0.3 mm, punctures elongate longitudinally, margins of punctures confluent; in posterior half about 3.6 punctures / 0.3 mm, distance between punctures as wide as 0.5 puncture. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium convex; epicranium with microsculpture of transverse microlines but without micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres: 1st thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–3rd longer than greatest width, 4th and 5th subquadrate, 6th– 10th wider than greatest length, width of distal margin of

35

9th and 10th 1.5ⴛ length; 6th–11th densely covered by microtrichae, 6th–10th with single circumferential row of prominent macrosetae near middle, 1st–5th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 0.95; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in at least 3 dense, contiguous longitudinal rows, punctures of a row contiguous; large, slightly convex, raised impunctate area in anterola-teral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering posterior 4/5. Elytra not depressed near scutellum, with large punctures, about 4.4 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII well punctuated (V–VI less so medially); IV–VI with yellow microsetae medially and posterolaterally, VII with brown ones. Paratergites IV–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–VI and VIII with yellow microsetae, VII with brown ones. Segment VII with black iridescent setae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 100) with evenly curved ventral surface, hind margin with two very large triangular lobes and a deep V-shaped emargination medially; sternum IX (identical to that in Fig. 104) with two small to moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a rounded extension (identical to that in Fig. 117). M a l e g e n i t a l i a . As in Figs. 222–225. Paramere in dorsal outline weakly converging in basal part and more strongly in subapical region, then forming an acute tip, apically subequal in length to median lobe and narrower; slightly dorsally convex in lateral outline. Sensory spinules (few) arranged in a loose medial cluster (alternatively seen as two vague rows near midline) and restricted to tip. Median lobe with broad subtruncate, emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Among species with golden-orange head, pronotum, and elytra, N. densiventris, N. semelius, and N. periergos are unique in the presence of microsculpture of transverse microlines on the epicranium. N. densiventris is very similar to N. semelius and N. periergos but differs in the shape of sternum VIII of males (Figs. 78, 96, 100): two very small triangular lobes extending beyond the posterior sternal margin and a moderate U-shaped medial emargination in N. densiventris; two strong triangular lobes and a very deep V-shaped medial emargination in N. semelius and N. periergos. N. semelius and N. periergos differ in the aedeagi (Figs. 206, 222): e.g., in N. semelius the median lobe in dorsal view is distally emarginate, while in N.

36

periergos it is distally narrowed to a broad subacute apex. Distribution. Unknown from elevations of 1000–1480 m in Bolivia (Fig. 21). Habitat. Not known. Derivatio nominis. The specific epithet is derived from the Greek word ‘⌺⑀␮⑀␭␩’ (Semeli), which was an ancient Greek minor goddess and mother of the ancient Greek god Dionysos. Remarks. Collected in flight intercept traps. Material. Holotype, ‚ with labels ‘Bolivia, Cochabamba, Cochabamba, 117 km E Yungas, Lagunitas, 1000 m 17o6´22´´S 65o40´57´´W, 8–12.ii.1999, F. Genier, BOL1G99 062 ex: FIT’, barcode label SM0174215, ‘HOLOTYPE Nordus semelius Chatzimanolis des. Chatzimanolis 2003’ (in SEMC). – Paratypes, 5 specimens: Same locality data and collector, barcode label SM0174207 (1 in SEMC); ‘Bolivia, Cochabamba, Cochabamba, 109 km E Yungas (Cochabamba– Villa Tunari Rd.), 1480 m, 17o8´50´´S 65o42´29´´W, 1– 6.ii.1999, R. Hanley, BOL1H99 026 ex: FIT’, barcode labels SM0162060, SM0161890, SM0161891 (3 in SEMC). Same locality data, F. Genier leg., barcode label SM0173971 (1 in SEMC). All paratypes with label ‘Paratype Nordus semelius Chatzimanolis des. Chatzimanolis 2003’.

4.30. Nordus simplex (Sharp, 1876) (Figs. 41, 89, 116, 226–229) Brachydirus simplex Sharp, 1876: 112. Nordus simplex (Sharp); HERMAN (2001: 3581). Description. L e n g t h . 9.5–11 mm. C o l o r a t i o n . Head golden-orange with anterior 3/4 of epicranium metallic green except a narrow golden-orange area medial to each eye and around insertion of antenna; mandibles and labrum dark brown; antennomeres all yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/ of VII golden-orange. Legs golden-orange. 3 H e a d . Microsetae yellow on metallic green part and black on golden-orange part. Anteclypeus very small. Punctures in anterior 3/4 of epicranium dense, about 4.0 punctures / 0.3 mm, punctures elongate longitudinally, margins of punctures confluent; in posterior part about 3.1 punctures / 0.3 mm, distance between punctures as wide as 0.5–1 punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 41): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–5th longer than greatest width, 6th wider (posterior margin) than greatest length, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 0.96; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 2–3 punctures); punctures flanking this area in at least 3 dense, contiguous longitudinal rows, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters,


and narrow impunctate area along posterior and posterolateral margins. Pronotum with black and few yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayish-yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with large punctures, about 4.5 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with few microsetae medially; V– VII extremely punctuated; IV–V and VIII covered with yellow microsetae (V with slightly fewer microsetae medially), VI and VII with yellow ones laterally and black ones medially. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; III–IV and VIII with yellow microsetae. Segment V–VII with black and yellow iridescent microsetae, throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 89) with evenly curved ventral surface, hind margin with broad and deep V-shaped emargination medially; sternum IX (identical to that in Fig. 110) with two moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a moderate rounded extension (Fig. 116). M a l e g e n i t a l i a . As in Figs. 226–229. Paramere in dorsal outline consistently converging to a narrowly rounded tip, apically subequal in length to median lobe and narrower; dorsally convex in lateral outline. Sensory spinules (many) arranged in a pair of narrowly separated bands and restricted to tip. Median lobe with broadly rounded apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Nordus aeneiceps and N. simplex can be distinguished from other golden-orange species by the metallic green anterior part of the epicranium. They differ from each other as follows: punctures on anterior part of epicranium dense and not longitudinally elongate in N. aeneiceps but very dense and longitudinally elongate in N. simplex; cerci straight in N. aeneiceps but slightly curved in N. simplex; structure of aedeagus as shown in Figs. 122–125, 226–229; paramere tip in dorsal outline pointed in N. aeneiceps but rounded in N. simplex. Distribution. Known from elevations of 150–450 m in Brazil, Colombia, Ecuador, and Peru (Fig. 60). There are no specimens known from Bolivia as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Habitat. Collected in flight intercept traps and on fleshy polypore fungi. Parasites. FRANK (1982) mentions the occurrence of Dichomyces peruvianus (Laboulbeniaceae) on N. simplex as documented by THAXTER (1900, 1908). Material. Lectotype, ‚, here designated, in cardboard box with dissected genitalia, with labels: on cardboard box mount in D. Sharp’s handwriting ‘Brachydirus simplex Type D. S. Amazon’, below ‘Type’, ‘Pará’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Lectotype Brachydirus simplex Sharp des. Chatzimanolis 2002’ (in BMNH). – Paralectotypes, 2 specimens: 1„ with labels ‘Type’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus simplex „ Amazon Type D. S.’, ‘Paralectotype


Brachydirus simplex Sharp des. Chatzimanolis 2002’ (in BMNH). 1‚ with labels ‘S. America Brazil’, ‘Brachydirus simplex 2nd Type Amazon D. S.’, [upside down] ‘Sharp Coll. 1905-313’, ‘Sharp Coll. by exchange with the British Museum’, ‘Paralectotype Brachydirus simplex Sharp des. Chatzimanolis 2002’ (in FMNH). – BRAZIL: Amazon (3 in BMNH); Amazon, Nanta (1 in BMNH); Brasilien (1 in DEI); Para (1 in BMNH); Para, 1904, M.H.D. Mathan leg. (1 in FMNH); Para, Jacareacanga, i.1969, F.R. Barbosa leg. (1 in AMNH). COLOMBIA: Amazonas PNN, Amacayaco Matanata, 3o41´S 70o15´W, 150 m, 19.ix.2001, D. Chota leg. malaise trap (1 in SEMC; in UNCB); Buenar[v]a (1 in BMNH); Villavicencio, 1938, F.W. Furry (3 in FMNH); Villavicencio, Ost-Columbia, 450 m, iv.1910, Fassl leg. (1 in FMNH, 1 in NMW); Villaviceneio, 21.vii.1938, C.H. Seevers leg. (1 in FMNH); Putomayo, Santa Rosa, Kofan Indian village, 2–24.x.1970, B. Malkin leg. (3 in FMNH); Putumayo PNN La Paya, Mamansoya, Tierra Baja, 0o6´S 74o58´W, 330 m, 22–26.ix.2001, R. Cobete leg. (1 in SEMC; 3 in UNCB). ECUADOR: Napo Prov., Napo R. 1 mi E Anvano, Anaconda Is., 14.ii.1991, J. Stamatov leg. (1 in AMNH); Pompeyo, v.1965, L. Pena leg. (1 in FMNH); Sucumbios, Sacha Lodge, 270 m, 0o28´14´´S 76o27´35´´W, 21–24.iii.1999, R. Brooks leg. FIT (5 in SEMC). PERU: Loreto Prov., Rio Ucayali, Yucuruche, 4o30´8´´S 73o26´1´´W, 1.i.1999, D. Brzoska leg. (3 in SEMC); Loreto, Explorama Lodge, Yanamono R., 80 km NE lquitos, 19.iv.1995, D. Bzroska leg. (2 in SEMC); Pachitea (1 in FMNH); Dept. Madre de Dios, Manu Prov., Parque Nac. Manu Zone Res. Rio Manu, Cocha Juarez, trail nr. Manu, Lodge, 18–24.ix.1991, A. Hartman leg. FIT (1 in FMNH); Madre de Dios, Cocha Cashu Biol. Stn., Manu National Park, 350 m, 11o53´45´´S 71o24´24´´W, 17–19.x.2000, R. Brooks leg. FIT (44 in SEMC); Tambopata Prov., Madre de Dios Dpto., 15 km NE Puerto Cuzco Amazonica, 12o33´S 69o03´W, 200 m, Maldonado Reserva, 20.vi.1989, J.S. Ashe & R. Leschen leg. FIT (2 in SEMC); Tambopata Prov., 15 km NE Puerto, Maldonado, 200 m, 15.vi.1989, R. Leschen leg. FIT (1 in SEMC).

4.31. Nordus solitarius (Sharp, 1884) (Figs. 42, 90, 117, 230–233) Brachydirus solitarius Sharp, 1884: 350. Brachydirus occipitalis Bernhauer, 1905: 185; new synonym. Brachydirus ecuadorensis Scheerpeltz, 1971: 104; new synonym. Nordus occipitalis (Bernhauer); HERMAN (2001: 3580). Nordus solitarius (Sharp); HERMAN (2001: 3581). Nordus ecuadorensis (Scheerpeltz); HERMAN (2001: 3580). Description. L e n g t h . 10–12 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres golden-orange except tip of 11 darker. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I–II, VIII, and posterior 1/ of VII golden-orange. Legs golden-orange except meta3 tibiae and -femora black. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior part of epicranium dense, about 4–4.5 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 3 punctures / 0.3 mm, distance between punctures as wide as 0.5–1 punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat in males, slightly convex in females; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 42): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–3rd longer

37

than greatest width, 5th and 6th almost quadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by micro-trichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 0.98; anterolateral angles not prominent, subcariniform; lateral margins slightly concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and shallow; medial impunctate area broadest anteriorly (width 0.5 punctures); punctures flanking this area in 2 longitudinal rows separated by a distance as wide as 1–2 punctures, punctures of a row not contiguous, but distance varies; large, slightly convex, raised impunctate area in anterola-teral quarters, and moderate impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with medium-sized punctures, about 5–5.5 punctures / 0.3 mm, and interspaces as wide as 1 puncture, covered with yellow microsetae. Legs with yellow macrosetae, except metafemora and -tibiae with black and yellow ones. A b d o m e n . Terga: III–IV with 0–4 discal microsetae; V without subbasal carina; V–VII strongly punctuated; V covered with yellow microsetae except for few darker ones medially, VI with tiny patch of yellowish-white microsetae near lateral margins. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III– VII densely punctuated otherwise; III–VII with yellow microsetae. Segment VII with black iridescent setae in anterolateral quarters. Cerci slightly curved, basal half dark brown to black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 90) with almost evenly curved ventral surface, hind margin with single broad lobe, apex of lobe with moderate U-shaped emargination medially, left and right edges of emargination continuing basally into subcarinate ridges that define a small triangular impunctate and depressed area at base of emargination; with brush-like patch of prominent setae on each side of emargination; sternum IX (identical to that in Fig. 107), with two long apical processes and shallow Ushaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a well rounded extension (Fig. 117). M a l e g e n i t a l i a . As in Figs. 230–233. Paramere in dorsal outline increasingly narrowed from base to middle, then widened again to a very broad subapical part, then converging to a short, narrow, parallel-sided apical part with a rounded tip, apically subequal in length to median lobe and narrower; almost completely straight in lateral outline, tip pointed and curved downwards. Distal part of paramere continuing basally into an elevated part on the dorsomedial face of the basal part, thus paramere with irregular dorsal surface around middle; paramere with a dorsomedian projection apically that in lateral view appears fin-like. Sensory spinules (many) arranged in a pair of regular far lateral rows and a regular medial band, both extending from tip almost 1/2 of paramere length towards paramere base.

38

Median lobe with broad subtruncate apex in dorsal outline, hardly upcurved apically in lateral outline, forming apically two widely separated, short, subtruncate dorsal lobes but no dorsomedial carina basal to these; slight emargination between lobes, with which the apex of the paramere interdigitates. Diagnosis. Nordus terminalis, N. omios, N. solitarius, and N. dichromos can be distinguished from all other Nordus species by the lacking (or extremely reduced) microsetation on the medial disc of abdominal tergum IV. These setae are also absent in N. fungicola, which differs from the above species by its convex rather than concave lateral margins of the pronotum. N. terminalis and N. omios can be distinguished from the other two species by the depressed elytra near scutellum, the very sparse punctuation on the elytra (distance between punctures larger than width of 1 puncture versus less than 1 in the other two species). N. solitarius can be distinguished from N. dichromos by the following characters: antennomere 4 subquadrate (longer than wide in N. dichromos); antennomere 5 subequal in length to 6 (5 longer than 6 in N. dichromos); epicranium more ot less flat (convex in N. dichromos); shape of sternum VIII in males as in Fig. 90 (versus Fig. 79 in N. dichromos). Synonymy. Brachydirus solitarius Sharp, B. occipitalis Bernhauer, and B. ecuadorensis Scheerpeltz are identical in all respects, including male sexual structures and male and female morphology and coloration. Distribution. Known from elevations of 0–1000 m in Colombia, Costa Rica, Ecuador, Nicaragua, Panama, and Venezuela (Fig. 66). Habitat. Rotting fruits. Remarks. Collected in flight intercept and malaise traps, and in Heliconia leaves. Material. Holotype of B. solitarius, ‚ glued on a paper card, with labels: on the paper card mount in D. Sharp’s handwriting ‘‚, Type, Brachydirus solitarius, Nicaragua, Sallé’, below ‘Type’, ‘Nicaragua’, ‘Sallé coll.’, ‘B. C. A. Col. I. 2, Brachydirus solitarius Sharp’, [upside down] ‘1096’, ‘Nordus solitarius (Sharp) det. S. Chatzimanolis 2002’ (in BMNH). SHARP (1884) states in the original description that he had available only a single specimen, which therefore is the holotype. The type locality for this species is ‘Fort San Carlos’ at the borders of Nicaragua and Costa Rica. Although the type locality is not written on the labels of the holotype, it is specified in the original description of the species by Sharp. – Lectotype of B. occipitalis, ‚, here designated, with labels ‘Collection Schild-Bargdorf, Costa Rica, San Carlos’, ‘occipitalis für Bang Haas detern. [sic] Bernh.’, ‘occipitalis Bernh. Typus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Lectotype Brachydirus occipitalis Bernhauer des. Chatzimanolis 2003’ (in FMNH). – Holotype of Brachydirus ecuadorensis, ‚ with labels ‘‚’, ‘Rep. Ecuador Esmeralda Parr. San Marcos, 5.x.1956’, ‘Rep. Ecuador Esmeralda Parr. San Marcos, 5.x.1956’, ‘Brachydirus ecuadorensis n. sp. ms.’, ‘ex. coll. Scheerpeltz’, ‘TYPUS, Brachydirus ecuadorensis O. Scheerpeltz’, ‘ecuadorensis Scheerp.’ (in NMW). SCHEERPELTZ (1971) explicitely mentioned that there is only one specimen for his description. However, in his collection there are two specimens, one labeled ‘Typus’ (the specimen above) and another, female one labeled ‘Cotypus’. The latter specimen was not mentioned in the original description and therefore is not a valid paratype for this species. – COLOMBIA: (no labels) (2 in BMNH); Anchicaya, 1000 m, 22–27.vii.1970, J.M. Campell leg. malaise trap (9 in CNCI); Anchicaya, 1000 m, 22.vii.1970, J.M. Campell leg. (3 in CNCI); Granial Muzo, ost Cordill., 750 m, xi.1910 (1 in BMNH, 2 in FMNH); Huaso Ost-Columb. (1 in FMNH); Muzo (1 in BMNH, 1 in NMW); Muzo, 400 m, vii.1910, Fassl leg. (1 in FMNH, 8 in NMW); Muzo, Dept. Boyaca, 900 m, 20–30.vi.1936, J. Banquaert leg. (1 in MCZC); Muzo, Rio Villanuzar, 600 m, viii.1910 (1 in NMW); Sogamoso ost Col., iii.1902 (1 in FMNH); Meta, Rio Guayriba, tributary of Rio Meta, xii.1946, L. Richter leg. (2 in


AMNH); Rio Negro, 500 m (1 in NMW); Meta, 3 mi W Villavicencio, 920 m, 11.iii.1955, E.I. Schlinger & E.S. Ross leg. (1 in CASC); Villavicencio, 450 m, ii.1902 (3 in FMNH, 3 in NMW); Valle del Cauca PNN Farallones de Cali, Anchicaya, 3o26Ń 76o48´W, 650–900 m, S. Sarria leg. malaise trap (23 in SEMC, 41 in UNCB). COSTA RICA: Alajuela, Sector Colonial Palmarena, 9 km SO de Bajo Rodriguez, 700 m, 10.vi.– 2.vii.1997, G. Carballo leg. malaise trap (2 in INBC); Alaj. Prov., 7 km N Fortuna, 10.viii.1989, F.D. Parker leg. (1 in SEMC); Alaj. Prov., 5.5 mi NW Florencia, 23.vii.1966, J.B. Karren leg. in Heliconia leaf (1 in SEMC); Prov. Alaju., Est. San Ramon, 620 m, 26.iv.1994, Fam. Hurtado Garcia leg. (1 in INBC); Carillo, Plason leg. (2 in NMW); Cartago, Turrialba, 20.viii.1989, F.D. Parker leg. (1 in SEMC); Cartago, Turrialba, 646 m, ix.1964, M.G. Naumann leg. (1 in SEMC); Ebene von Limon bei Las Mercedes, Farm Hamburg am Reventazon 15.x.1923, F. Nevermann leg. (3 in FMNH); Farm Cardiella, vii.1938, Bierig leg. (6 in FMNH); Farm Cabriella, vii.1938 (2 in BMNH); Guanac. Prov., Estac. Pitillia, 700 m, 9 km S Santa Cecilia, iii.1989 (20 in INBC); Guanac., Estac. Pitilla, Santa Cecilia, 700 m, v.1989, Gould leg. (1 in INBC); Guanacaste, Patilla Biological Station, 610 m, 10o59´22´Ń 85o25´33´´W, 13-15.vii.2000, J.S. Ashe, R. Brooks & Z. Falin leg. FIT (1 in SEMC); Prov. Heredia, Est. Magsasay, 200 m, P.N. Braulio Carillo, v.1991, R. Aguilar leg. (4 in INBC); Prov. Here., Sendero R. Sardinalito a Raravis, 850–700 m, A. Fernandez leg. (2 in INBC); Prov. Here., Est. El Celbo, P.N. Braulio Carillo, 400–600 m, ii.1990, C. Chaves leg. (27 in INBC); Heredia, La Selva, 80 m, 19.v.1993, J.S Ashe leg. FIT (2 in INBC, 5 in SEMC); Heredia, La Selva, 3.3 km S Puerto Viejo, 100 m, 10o25´0´Ń 84o0´0´´W (26 in SEMC); Heredia, Chilamate, 75 m, v.1989, Hanson & Godoy leg. (6 in INBC); Heredia, Finca La Selva, 30.v.1979, Andrews leg. abandonded cocoa plantantion (1 in MCZC); Heredia, Puerto Viejo, Finca La Selva, vi.1979, C. Andrews leg. on Rudga leaf in forest (1 in MCZC); Heredia Prov., Sardinalito, Braulio Carrillo, 360–500 m, x.1998, R. Aguilar & M. Zumbado leg. (1 in INBC); Limon, Farm Castilla, vii.1938, Bierig leg. (1 in AMNH); Prov. Limon, Valle la Estrella, R.B. Hitoy Cerere, A.C. Amistad, 100–200 m, 1994, G. Carballo leg. (1 in INBC); Prov. Limon, Sector Cerro Cocori Fcs de E. Rojas, 150 m, 26.iii.–24.iv.1992, F.A. Quesada leg. (1 in INBC); Prov. Limon, Est. Hitoy Cerere, 100 m, Res. Biol. Hitoy Cerere, ix.1992, G. Garballo leg. (17 in INBC); Limon Prov., Pandora, 200 ft [67 m], 23.viii.1963, Wille, Kerfoot, C.D. & D.R. Michener leg. (1 in SEMC); Prov. Limon, Puerto Viejo, 0 m, xii.1990, M. Zumbaco leg. (1 in INBC); Limon, 16 km W Guapiles, 400 m, x–xi.1989, P. Hanson leg. (1 in INBC); Prov. Limon, Amubri, 70 m, 12–31.x.1993, G. Gallardo leg. (1 in INBC); Prov. Punta., Es. Q. Bonita, 50 m, ix.1993, R. Guzman leg. (1 in INBC); Prov. Slsj., Falda Este Vol. Tenorio Colonia Rio Celeste Fca Magli, 400–500 m, xi.1988, A. Solis leg. (1 in INBC); San Carlos, Schlid & Burgdorf leg. (4 in USNM); Turrialba, 7.viii.1965, under Ficus bark, A. Raske leg. (1 in CNCI). ECUADOR: Cochabe low, i.1897, Rosenberg leg. (1 in FMNH, 1 in ISNB); Esmeraldas, Bilsa, 0o20´0´´S 79o43´0´´W, iv–vi.1996, Hibbs leg. malaise trap (6 in SEMC); Manabi, Cerro pala de Pajaro, 300–500 m, 0o0´0´Ń 75o57´0´´W, 19–21.vi.1996, Hibbs leg. malaise trap (1 in SEMC); Pich. 16 km SE Sto. Domingo Tinalandia, 680 m, 20– 21.vi.1975, S. Peck leg. (2 in CNCI); Pich. 4 km SE Sto Domingo, 500 m, 8–11.vi.1976, S. Peck leg. (1 in CNCI); St. Domingo, Rio Palenque 47 km S, 700 m, 22–27.ii.1976, H. & A. Howden leg. malaise trap (2 in CNCI); Pich. E Sto. Domingo, 8–14.v.1988, Hanson & Bohart leg. (1 in SEMC); Pich. Prov., 800 m, Tinalandia, ii.1983, M. Sharkey & Masner leg. malaise trap (1 in CNCI); Pichincha, Tinalandia, Santo Domingo, 16 km E, 750 m, 0o16´53´´S 79o3´39´´W, 26– 27.iii.1999, R. Brooks & D. Brzoska leg. FIT (17 in SEMC); Pich. 47 km SE Sto Domingo, Rio Palenque Sta., 22– 28.ii.1976, 300 m, J.M. Campell leg. (1 in CNCI); Pichincha, 6 mi E Santo Domingo de los Colorados, 24.ii.1955, E.I. Schlinger & E.S. Ross leg. (3 in CASC). NICARAGUA: Rio San Juan Dept., 60 km SE San Carlos, Refugio Bartola, 100 m, 10o58´40´Ń 84o20´30´´W, 25–28.v.2002, R. Brooks, S. Chatzimanolis & Z. Falin leg. FIT (2 in SEMC). PANAMA: Cocle, 7.2 km N El Cope, 730 m, 8o37Ń 80o35´W, 20.v.– 7.vi.1995, J.S. Ashe & R. Brooks leg. FIT (4 in SEMC); Darien, Cana, Biological Station, 500–600 m, 7o45´18´Ń


77o41´6´´W, vi.1996, J.S. Ashe & R. Brooks leg. FIT, rotting palm fruits (48 in SEMC); Darien, Cana El., 450 m, 24.ix.1982, Stockwell leg. (1 in FMNH); Panama Pr., Cerro Campana, 850 m, 8o40Ń 79o56´W, 12.iii.1972, Stockwell leg. (1 in BMNH); Panama Prov., Cerro Campana, 550 m, 8.xii.1972, Bivia, 8o40Ń 79o56´W (1 in USNM); Panama Pr., Cerro Campana, 820 m, 24.iii.1982, Stockwell leg. (1 in FMNH). VENEZUELA: Merida, 35 km S El Vigia, 2.vi.1976, A.S. Menke & D. Vincent leg. (1 in USNM).

4.32. Nordus speciosus (Bernhauer, 1905) (Figs. 4, 14, 43, 101, 111, 118, 234–236) Brachydirus speciosus Bernhauer, 1905: 187. Nordus speciosus (Bernhauer); HERMAN (2001: 3581). Description. L e n g t h . 11.0–13.0 mm. C o l o r a t i o n . Head metallic green with two bright red spots at middle of epicranium; mandibles and labrum orange; antennomeres all yellow. Pronotum, elytra, and scutellum metallic green except elytra orange along posterior margin. Abdomen black except segments I–II brown, VII–VIII and posterior margin of VI golden-orange. Legs golden-orange except around tibiofemoral articulations dark brown; in some females from La Paz (Bolivia) golden-orange area on elytra larger and lighter; abdomen light brown except segment VII dark brown with posterior 1/3 golden-orange (no males of such coloration known). H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 6.2 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior half about 3.1 punctures / 0.3 mm, distance between punctures as wide as 0.5 punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1–1.5ⴛ length of eye. Middle of epicra-nium slightly convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 43): 1st thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–5th longer than greatest width, 6th subquadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 1.5ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum longer than wide, width : length = 0.80; anterolateral angles not prominent, subcariniform; lateral margins strongly concave in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures medium-sized and shallow; medial impunctate area broadest anteriorly (width 3–4 punctures); punctures flanking this area in 1–2 sparse longitudinal rows separated by a distance as wide as 1–1.5 punctures, punctures of a row not contiguous, but distance varies; moderately sized, slightly convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming 2 rows, not covering entire width. Scutellum without arch-like carina, with yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, and interspaces as wide as 1–1.5 punctures, covered with yellow microsetae. Legs with yellow

39

macrosetae; in males legs modified (see under secondary sexual structures); in female mesotarsus 1st tarsomere elongated, 2ⴛ length of 2nd; 5th elongated similar to 1st. A b d o m e n . Terga: III with 0–4 discal microsetae; V with subbasal carina; IV–VII well punctuated (V less so medially); IV–VI and VIII with brownish-yellow microsetae throughout, VII with yellow ones throughout. Paratergites IV–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise, IV–VI slightly less so posteromedially; III–VI and VIII with brownish-yellow microsetae, VII with brown ones. Segment VII with black iridescent setae throughout. Cerci thin and straight, basal 2/3 light brown and distal 1/3 yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: profemur very thick and robust; protibia short, robust, expanded from base to broad apex; mesofemur very robust, dorsoventrally flattened distally; mesotibia (Fig. 14) on dorsal face adjacent to femorotibial articulation with flattened triangular area, from which a basally directed domed process arises that articulates with large U-shaped socket in distal dorsal margin of femur; articular region sharply curved in posterior aspect, mesotibia greatly expanded posteriorly towards apex, ventral surface of apical half of expansion strongly excavated and concave; 1st mesotarsomere greatly expanded, 2.2ⴛ length of 2nd, rest of mesotarsus also expanded, 5th mesotarsomere shorter than 1st; metafemur with basal third strongly expanded and concave ventrally; the margins of the concavity (two lines along the anteroventral and posteroventral faces of the femur) are prolonged as large blade-like carinae, each of which is expanded distally (near knee) into a prominent tooth. Hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 101) with evenly curved ventral surface, hind margin with two long trapezoid setose lobes and a very broad and deep U-shaped emargination medially; sternum IX (Fig. 111) without any apical processes and with moderately large U-shaped emargination. Female: with no obvious secondary sexual structures except hind margin of sternum VIII sinuate (Fig. 118). M a l e g e n i t a l i a . As in Figs. 234–236. Paramere in dorsal outline very wide, widened in subapical part, then abruptly and strongly narrowing to narrow, acute apical part (thus, distal margins sinuate), apically shorter than median lobe; dorsally slightly concave in lateral outline, very distinctive through basal dorsal bulge followed by deep constriction; with diamond-shaped depression dorsomedially below tip. Sensory spinules (many) arranged in a pair of clusters shortly basal to tip. Median lobe with strong lateral constriction in distal third and with broadly subacute apex in dorsal outline, strongly upcurved in distal half in lateral outline, bearing a dorsomedian process near midlength, otherwise without a dorsomedian carina, and without pair of distal dorsal lobes. Diagnosis. Nordus cherylae, N. elegans, N. portokalis, and N. speciosus can be distinguished from all other species of Nordus by the presence of a subbasal carina (in addition to the basal one) on abdominal tergum V. N. speciosus can be distinguished from the other three species by the metallic green head, pronotum and elytra. Distribution. Known from elevations of 750–2300 m in Bolivia and Peru (Fig. 67). No specimens are known from Ecuador as erroneously mentioned in SCHEERPELTZ (1971). Habitat. Unknown.

40

Remarks. Collected in flight intercept traps. Material. Holotype, ‚ with labels ‘Bolivien’, ‘Bolivien Bang Haas’, ‘prasinus für .i.l.’, ‘speciosus Bern. Typus’, ‘Chicago NHMuseum M. Bernhauer Collection’ (in FMNH). BERNHAUER (1905) states in the original description that he had available only a single specimen, which therefore is the holotype. There are 6 specimens in Scheerpeltz’ collection labeled as ‘Cotypus’. These specimens do not belong to the type series since Bernhauer mentioned that he had only 1 specimen. – BOLIVIA: Cochabamba Prov., Cochabamba, 109 km E Yungas, Cochabamba–Villa Tunari Rd., 1400 m, 17o8´52´´S 65o42´54´´W, 8–12.ii.1999, F. Genier leg. FIT (1 in SEMC); Coroico (1 in BMNH, 1 in FMNH); La Paz Prov., Chulumani, 9.2 km E, 2100 m, 16o20´58´´S 67o30´58´´W, 19–21.i.2001, J.S. Ashe & R.S. Hanley leg. FIT (6 in SEMC); La Paz Prov., Chulumani, 9.2 km E, 2200 m, 16o20´59´´S 67o30´18´´W, 19– 21.i.2001, J.S. Ashe & R.S. Hanley leg. FIT (1 in SEMC); La Paz Prov., Chulumani, 9.2 km E, 2100–2300 m, 16o20´59´´S 67o30´18´´W, 21.i.2001, J.S. Ashe & R.S. Hanley leg. (1 in SEMC); Rio Longo, 750 m, Fassl leg. (3 in FMNH, 5 in NMW); Rio Longo (4 in NMW). PERU: Ocobambe, Kraatz leg. (1 in DEI).

4.33. Nordus stomachoponos sp.n. (Figs. 44, 119) Description. L e n g t h . 8.5–10 mm. C o l o r a t i o n . Head golden-orange with anterior 3/4 of epicranium shining black; mandibles and labrum dark brown; antennomeres light brown to black. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/3 of VII goldenorange. Legs yellow. H e a d . Microsetae brown to black. Anteclypeus very small. Punctures in anterior half of epicranium very dense, about 9.3 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior part about 4.5 punctures / 0.3 mm, punctures almost confluent. Length of eyes ca. 0.8ⴛ length of head, distance between eyes ca. 1ⴛ length of eye. Middle of epicranium convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 44): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th slightly shorter than 4th, 1st–5th longer than greatest width, 6th quadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 1.5ⴛ length; 6th–11th densely covered by microtrichae, 4th–10th with single circumferential row of prominent macrosetae near middle, 1st–3rd with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.02; anterolateral angles not prominent, subcariniform; lateral margins slightly concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and shallow; medial impunctate area broadest anteriorly (width 1 puncture); punctures flanking this area in 1–2 moderately dense longitudinal rows separated by a distance as wide as 1 puncture, punctures of a row not contiguous, but distance varies; large, convex, raised impunctate area in anterolateral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with brown microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with whitishyellow microsetae and punctures covering entire surface.


Elytra not depressed near scutellum, with medium-sized punctures, about 3.6 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with few microsetae on medial disc; V–VII extremely punctuated, slightly less so medially; V–VI with yellow microsetae laterally and brown ones medially, VII completely covered with brown ones. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise; III–IV and VIII with yellow microsetae. Segment VII with black iridescent setae in anterolateral quarters. Cerci slightly curved, basal 1/3 brown and distal 2/ yellow. 3 S e c o n d a r y s e x u a l s t r u c t u r e s . Male: unknown. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a rounded extension (Fig. 119). M a l e g e n i t a l i a . Males unknown. Diagnosis. N. stomachoponos is very similar to N. amphivolos and N. longipes. Only these species are golden-orange with the anterior 3/4 of the head shining black and have antennomeres 2–10 light brown to black. N. stomachoponos can be distinguished from the other two by the very shallow and sparse punctuation on the pronotum, the very dense punctuation on the anterior part of the head, and antennomere 7 being wider than long (subquadrate in the other two species). Distribution. Known from elevations of 500–1000 m in Colombia and in Darien, Panama (Fig. 20). Habitat. Rotting fruits. Derivatio nominis. The specific epithet is derived from the Greek word ‘␴␶␱␮␣␹ó␲␱␯␱␵’ (stomach ache). This was the last new species discovered during the present study. Remarks. Collected from human dung traps and flight intercept traps. The absence of male specimens is problematic for this new species, however the overall morphology, especially of the head and the pronotum, make it easily recognizable. In addition, this species is known only from regions that are not easily accessible to scientists and thus additional specimens of this species may not be collected in the near future. Material. Holotype, „ with labels ‘Panama, Darien, Cana, Biological Station, 530 m, 7o45´18´Ń 77o41´6´´W, 7.vi.1996, J.S. Ashe & R. Brooks PAN1AB96 069 ex: rotting fruits’, barcode label SM0034206, ‘HOLOTYPE Nordus stomachoponos Chatzimanolis des. Chatzimanolis 2003’ (in SEMC). – Paratypes, 12„„: Same locality data and collector but ex: FIT, barcode labels SM0011060, SM0048077, SM0048100, SM0039812, SM0034207 (5 in SEMC). Same locality and collector but 550 m, ex: FIT, barcode labels SM0049043, SM0049010 (2 in SEMC). Same locality data but 500 m, R.S. Anderson, ex: human dung trap, barcode labels SM0034445, SM0034493, SM0034492 (3 in SEMC). ‘Colom. 1000 m Anchicaya, 20.vii.1970’, ‘H.F. Howden’ (1 in CNCI). ‘Colom. 1000 m Anchicaya, 20.vii.1970’, ‘H.F. Howden’, ‘Brachydirus aeneiceps Sharp, det. J.M. Campell 1976’, ‘Nordus facialis (Er.) det. Smetana’ (1 in CNCI). All paratypes with label ‘Paratype Nordus stomachoponos Chatzimanolis des. Chatzimanolis 2003’.

4.34. Nordus styloceros (Sharp, 1876) (Figs. 45, 65, 91, 120, 237–240) Brachydirus styloceros Sharp, 1876: 111. Nordus styloceros (Sharp); HERMAN (2001: 3581).


Description. L e n g t h . 9.5–11 mm. C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres all yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I, II, VIII, and posterior 1/4 of VII golden-orange. Legs golden-orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior 2/3 of epicranium dense, about 6.2 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior 1/3 about 4.4 punctures / 0.3 mm, punctures almost confluent. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 2ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 45): 1st thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–5th longer than greatest width, 6th quadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th 1.5ⴛ length; 6th–11th densely covered by microtrichae, 6th–10th with single circumferential row of prominent macrosetae near middle, 1st–5th with irregularly scattered macrosetae. T h o r a x . Pronotum slightly wider than long, width : length = 1.06; anterolateral angles not prominent, subcariniform; lateral margins almost parallel to each other in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–1.5 punctures); punctures flanking this area in at least 1–2 moderately dense longitudinal rows separated by a distance as wide as 1–1.5 punctures, punctures of a row not contiguous, but distance varies; large, convex, raised impunctate area in anterolateral quarters, and narrow impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming 2–3 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with large punctures, about 4.9 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with microsetae on medial disc; V– VII extremely punctuated; V, VI and VIII covered with yellow microsetae, VII with brown ones. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; III–VII with yellow microsetae. Segment VII with black and yellow iridescent microsetae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (Fig. 65) with very strong notch medially and prominent setae around margin of notch; sternum VIII (Fig. 91) not with evenly curved ventral surface but rather elevated along middle, forming a carina, distal end of carina continuing into a single, very long and narrow, discrete median lobe upon the hind margin, lobe distally forked into two branches that diverge and then converge apically; sternum IX (identical to that in Fig. 107) with two long apical processes and a shallow emargination medially. Female: with no obvious secondary sexual structures except hind

41

margin of sternum VIII forming a pointed extension (Fig. 120). M a l e g e n i t a l i a . As in Figs. 237–240. Paramere in dorsal outline consistently converging to a rounded tip, apically subequal in length to median lobe and narrower; dorsally convex in lateral outline. Sensory spinules (few) arranged in a median band and restricted to tip. Median lobe with broad and emarginate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Among the golden-orange species N. diversiventris, N. testaceus, and N. styloceros can be identified by the almost parallel lateral margins of the pronotum. These three species are easily distinguished by the unique shapes of their aedeagi (Figs. 166, 237, 245) and sterna VIII in males and by the following characters in both sexes: N. diversiventris and N. styloceros differ from N. testaceus by the medial impunctate area on the pronotum being as wide as 1–1.5 punctures (0.5 punctures or less in N. testaceus). N. diversiventris differs from N. styloceros by antennomere 5 being longer than wide (subquadrate in N. styloceros) and punctures on elytra being separated by a distance as wide as 0.5–1 punctures (confluent in N. styloceros). Distribution. Known from elevations of 180–800 m in Brazil, Colombia, French Guiana, Guyana, and Peru (Fig. 67). There are no specimens known from Bolivia as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Habitat. Unknown. Remarks. Collected in flight intercept traps, howler monkeys’ feces, and on a dead spider monkey. SHARP (1876) mentioned that he had 11 specimens, however only 8 specimens were found that belong to the type series. Material. Lectotype, ‚, here designated, with labels ‘Type’, ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905313’, ‘Brachydirus styloceros ‚ Type D.S.’, ‘Lectotype Brachydirus styloceros Sharp des. Chatzimanolis 2002’ (in BMNH). – Paralectotypes, 8 specimens: 1„ with labels ‘Type’, ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905313’, ‘Brachydirus styloceros „ Type amazons D.S’, ‘Paralectotype Brachydirus styloceros Sharp des. Chatzimanolis 2002’ (in BMNH). 2„„ with labels ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus styloceros „ 2nd Type amazons D.S’, ‘Paralectotype Brachydirus styloceros Sharp des. Chatzimanolis 2002’ (in BMNH). 3‚‚ with labels ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Brachydirus styloceros ‚ 2nd Type amazons D.S’, ‘Paralectotype Brachydirus styloceros Sharp des. Chatzimanolis 2002’ (2 in BMNH, 1 in FMNH). 1‚ with labels: on specimen card in D. Sharp’s handwriting ‘Brachydirus styloceros ‚ 2nd Type D.S.’, below ‘Ega’, ‘S. America Brazil’, [upside down] ‘Sharp Coll. 1905-313’, ‘Paralectotype Brachydirus styloceros Sharp des. Chatzimanolis 2002’ (in BMNH). – BRAZIL: Amazon, Bates leg. (1 in BMNH); Amazon, Kraatz leg. (1 in DEI); Amazons (1 in BMNH); Ega (3 in BMNH, 1 in FMNH); Castanhal PA, 28.x.1962, J. Bechyne leg. (1 in CNCI); Para, Aldeia Aracu, Igarape Gurupumu Maranhao, 50 km E of Caninde, 10– 11.v.1963, B. Malkin leg. howler’s feces (5 in FMNH); Para, Caninde, Rio Gurupi, 7–15.iv.1963, B. Malkin leg. dead spider monkey (1 in FMNH); Para, Caninde, Rio Gurupi, 7– 15.iv.1963, B. Malkin leg. forest sweep (1 in FMNH); Para, Jacareacanga, xii.1968, M. Alvarenga leg. (6 in AMNH); Para, Redencao vic. Keyapo Territory Pinkaiti R., xi.1998, tropical evergreen seasonal lowland forest on clay soil, P.Y. Schaffer leg. (2 in FMNH); Para, Baker leg. (1 in FMNH); Rondonia, 62 km SE Ariquemes, 17–24.iii.1989, 180 m, W.J. Hanson leg. (2 in SEMC); Sinop, M. Grosso, x.1975, M. Alvarenga leg. (10 in CNCI); Sinop, M. Grosso, x.1976, M. Alvarenga leg. (44 in CNCI). COLOMBIA: Putumayo PNN La Paya Mamansoya, Tierra Alta, 0o6´S 74o58´W, 350 m, 22–26.ix.2001, D. Campos

42

leg. malaise trap (1 in SEMC; 1 in UNCB). FRENCH GUIANA: Mai, Bas Moroni leg. Le Moul (1 in FMNH). GUYANA: Brit. Guiana [Guyana], upper Kutari R., 10–12.xii. 1935, G.A. Hudson leg. (1 in BMNH). PERU: (no labels) (3 in NMW); 1955, Karl Brancsik leg. (1 in AMNH); Callanga (1 in DEI, 1 in NMW); Callanga, Bierig leg. (1 in FMNH); Loreto Dept., 1.5 km N Teniente Lopez, 2o35´66´S 76o06´92´´W, 22.vii.1993, 210–240 m, R. Leschen leg. FIT (2 in SEMC); Madre de Dios Dept., Tambopata, 25.x.1982, L.E. Watrous & G. Mazurek leg. (1 in FMNH); Madre de Dios, Pentiacolla Lodge, 5.5 km NW El Mirador trail, Alto Madra de Dios River, 500 m, 12o39´10´´S 71o15´28´´W, 23–26.x.2000, R. Brooks leg. FIT (3 in SEMC); Madre de Dios, Pentiacolla Lodge, 8 km El Mirador trail, Alto Madra de Dios River, 800 m, 12o38´30´´S 71o16´41´´W, 23–26.x.2000, R. Brooks leg. FIT (1 in SEMC); Madre de Dios, Pentiacolla Lodge, Alto Madra de Dios River, 400 m, 12o39´22´´S 71o13´55´´W, 23– 26.x.2000, R. Brooks leg. FIT (1 in SEMC); Madre de Dios, Rio Tambopata Res., 30 km (air) SW Pto. Maldonato, 200 m, 12o50´S 69o20´W, 1982, N.E. Stork leg. (2 in BMNH, 1 in SEMC); Marcapata, Bierig leg. (1 in FMNH); Pachitea (2 in FMNH); 50 mi E Tingo Maria, 12.xii.1954, E.I. Schlinger & E.S. Ross leg. (1 in CASC).

4.35. Nordus terminalis (Sharp, 1884) (Figs. 46, 88, 108, 121, 241–244) Brachydirus terminalis Sharp, 1884: 348. Brachydirus divisus Sharp, 1884: 349; new synonym. Brachydirus gracilis Sharp, 1884: 349; new synonym. Nordus divisus (Sharp); HERMAN (2001: 3580). Nordus gracilis (Sharp); HERMAN (2001: 3580). Nordus terminalis (Sharp); HERMAN (2001: 3581). Description. L e n g t h . 9.0–10.5 mm. C o l o r a t i o n . (1) Black morph: Head black; mandibles and labrum brown; antennomere 1 dark orange, 2–7 light brown, 8–11 yellow. Pronotum, elytra, and scutellum black. Abdomen black except segments I–II, VIII, and posterior 1/ of VII golden-orange. Legs black except all tarsi and 3 pro- and mesotibiae dark orange, distal parts of pro- and mesofemora orange. (2) Golden-orange/black morph (previously identified as B. divisus Sharp): Head golden-orange, anterior 3/4 slightly darker than posterior; mandibles and labrum orange; antennomere 1 dark orange, 2–7 light brown, 8–11 yellow. Pronotum and scutellum goldenorange, elytra dark brown to black. Abdomen black except segments I–II, VIII, and posterior 1/3 of VII golden-orange. Legs golden-orange. (3) Golden-orange morph (previously identified as B. gracilis Sharp): Head golden-orange, anterior 3/4 slightly darker than posterior; mandibles and labrum orange; antennomere 1 dark orange, 2–7 light brown, 8–11 yellow. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segments I–II, VIII, and posterior 1/ of VII golden-orange. Legs golden-orange. 3 H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior 3/4 of epicranium dense, about 5.8 punctures / 0.3 mm, punctures elongate longitudinally, margins of punctures confluent; in posterior 1/4 about 3.1 punctures / 0.3 mm, distance between punctures as wide as 1 puncture. Length of eyes ca. 0.8ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.04), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 46): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–5th longer than greatest width, 6th


quadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.02; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures medium-sized and shallow; medial impunctate area broadest anteriorly (width 2–3 punctures); punctures flanking this area in 1–2 sparse rows, separated by distance as wide as 1 puncture, punctures of a row not contiguous, but distance varies; large, convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron without any punctures anteriorly. Metepisternum with punctures forming 2 rows. Scutellum without arch-like carina, with whitish-yellow microsetae and punctures covering posterior half. Elytra depressed near scutellum, with medium-sized punctures, about 4 punctures / 0.3 mm, and interspaces as wide as 1–2 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III–IV with 0–4 discal microsetae; V without subbasal carina; V–VI moderately punctuated with fewer setae medially; IV–V and VIII covered with yellow microsetae, VI and VII with black ones. Paratergites III– VII covered with whitish-yellow setae on posterior half. Sterna: VI–VII on anterolateral corners without punctures or with sparse punctures; III–VII densely punctuated otherwise; III–VII with yellow microsetae. Segment VII with black iridescent setae in anterolateral quarters. Cerci slightly curved, basal half dark brown to black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 64) with broad and very deep emargination medially; sternum VIII (Fig. 88) with evenly curved ventral surface, hind margin with single broad, long lobe and deep, narrow, parallel-sided median cleft, with brush-like patch of prominent setae on basal part of each side of cleft; sternum IX (Fig. 108) with two long apical processes and a deep U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a well rounded extension (Fig. 121). M a l e g e n i t a l i a . As in Figs. 241–244. Paramere in dorsal outline very slender but with distinctly widened, disclike part around middle, distally slightly narrowed to a narrowly rounded tip, apically subequal in length to median lobe and narrower; altogether slightly dorsally convex in lateral outline, but with dorsally arched portion immediately beyond middle (at disc), apex not hook-like. Sensory spinules (many) arranged in a distal medial cluster and two irregular longitudinal rows more basally, all restricted to apical 1/6 of paramere. Median lobe with fairly narrowly rounded apex in dorsal outline, slightly upcurved apically in lateral outline, forming apically two widely separated, short dorsal lobes but no dorsomedial carina basal to these. Diagnosis. Nordus terminalis, N. omios, N. solitarius, and N. dichromos can be distinguished from all other Nordus species by the lacking (or extremely reduced) microsetation on the medial disc of abdominal tergum IV. These setae are also absent in N. fungicola, which differs from the above species by its convex rather than concave lateral margins of

43


the pronotum. N. terminalis and N. omios can be distinguished from the other two species by the depressed elytra near scutellum, the very sparse punctuation on the elytra (distance between punctures larger than width of 1 puncture versus less than 1 in the other two species) and, in the males, by their very distinctive shape of the paramere (disclike middle surrounded by very slender parts; Figs. 202, 241) and sternum VIII (one large lobe on hind margin, with deep, narrow medial cleft, and brush-like patch of prominent setae at anterior end of cleft; Figs. 87, 88). N. terminalis and N. omios differ only in the shapes of the aedeagus and male sternum VIII: lobe of sternum VIII slightly shorter in N. omios; paramere (lateral view, Figs. 203, 242) hooklike apically in N. omios, but almost straight apically in N. terminalis; sensory spinules extending further towards paramere base in N. terminalis than in N. omios; different shape of apex of median lobe. Synonymy. Brachydirus terminalis Sharp, B. gracilis Sharp, and B. divisus Sharp are identical in all respects, including male sexual structures and male and female morphology except the coloration as noted above. Distribution. Golden-orange morph: known from elevation of 100 m in Nicaragua (Fig. 68); no specimens are known from El Salvador as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Golden-orange/ black morph: known from elevations of 200–1200 m in Costa Rica and Panama (Fig. 68); no specimens are known from Nicaragua as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Black morph: known from elevations of 0–1400 m in Costa Rica, Guatemala, and Panama (Fig. 68). The color morphs of N. terminalis are not known to be sympatric at any locality. Habitat. Unknown. Remarks. Collected in flight intercept traps and on rotting palm trunk. For entomologists familiar with D. Sharp’s work, it comes with surprise that he described in the same publication three species that are proposed here to be synonymous. An explanation for this can be the fact that he had only a female of B. terminalis (and thus was not able to compare the unique male sternum VIII), a single male for B. gracilis, and only one male in poor condition for B. divisus. The differences in the color pattern and maybe the fact that he described an enormous number of species in the ‘Biologia Centrali Americana’ may also have contributed to the above mistakes. Material. Holotype of B. terminalis, „ glued on paper card, with labels: on paper card mount in D. Sharp’s handwriting ‘Brachydirus terminalis „ Type, D.S. Panima Vera Pag. Champion’, below, ‘Type’, ‘Panima, Guatemala, Champion’, ‘B. C. A. Col. I. 2, Brachydirus terminalis Sharp’, [upside down] ‘Sharp Coll. 1905-313’, ‘Nordus terminalis (Sharp) det. S. Chatzimanolis 2002’ (in BMNH). SHARP (1884) states in the original description that he had available only a single specimen, which therefore is the holotype. – Holotype of B. gracilis, ‚, glued on a paper card, with labels: on paper card mount in D. Sharp’s handwriting ‘‚ Brachydirus gracilis Type, D.S. Chontales, Nicaragua, Belt’, below, ‘Type’, ‘Chontales, Nicaragua, T. Belt’, ‘B. C. A. Col. I. 2, Brachydirus gracilis Sharp’, [upside down] ‘Sharp Coll. 1905-313’, ‘Nordus terminalis (Sharp) det. S. Chatzimanolis 2002’ (in BMNH). SHARP (1884) states in the original description that he had available only a single specimen, which therefore is the holotype. – Lectotype of B. divisus, ‚, here designated, glued on paper card, with labels: on cardboard box mount in D. Sharp’s handwriting ‘‚ Brachydirus divisus ‚ Type D. S. Bugaba, Champion’, below ‘Type’, ‘Bugaba, Panama, Champion’, ‘B. C. A. Col. I. 2, Brachydirus divisus Sharp’, [upside down] ‘Sharp Coll. 1905-313’, ‘Syn-type’, ‘Lectotype Brachydirus divisus Sharp des. Chatzimanolis 2002’ (in BMNH). –

Paralectotypes of B. divisus, 2 specimens: 1„ glued on paper card, with labels: on cardboard box mount in D. Sharp’s handwriting ‘„ Brachydirus divisus Type D.S. Bugaba, Champion’, below ‘Type’, ‘Bugaba, Panama, Champion’, ‘B. C. A. Col. I. 2, Brachydirus divisus Sharp’, [upside down] ‘Sharp Coll. 1905-313’, ‘Syn-type’, ‘Paralectotype Brachydirus divisus Sharp des. Chatzimanolis 2002’ (in BMNH). 1„ glued on paper card, with labels: on cardboard box mount in D. Sharp’s handwriting ‘„ Brachydirus divisus Type D.S. Bugaba, Champion’, below ‘Bugaba, Panama, Champion’, ‘B. C. A. Col. I. 2, Brachydirus divisus Sharp’, ‘Sharp Coll. by exchange with the British Museum’, ‘Paralectotype Brachydirus divisus Sharp des. Chatzimanolis 2002’ (in FMNH). – (Goldenorange morph:) NICARAGUA: Rio San Juan Dept. 60 km SE San Carlos, Refugio Bartola, 100 m, 10o58.40Ń 84o20.30´W, 25–28.v.2002, R. Brooks, S. Chatzimanolis & Z. Falin leg. FIT (1 in SEMC). (Golden-orange/Black morph:) COSTA RICA: San Vito de C.B., Las Cruces, 1200 m, 9.vii.–7.viii.1982, B. Gill leg. malaise trap (5 in CNCI); Puntarenas, Las Cruces botanical garden, nr. San Vito 3500 ft [1167 m], 27–28.ii.1985, L. Herman leg. (1 in AMNH); S.J. Nara, NE Quepos, 16.vii.1975, W.J. Hanson leg. (1 in SEMC). PANAMA: Bugabila (1 in NMW); Bugabila, 200 m, Fassl leg. (3 in FMNH, 2 in NMW); Lino (1 in BMNH); Lino, Bierig leg. (1 in FMNH); Lino, 800 m, Fassl leg. (3 in AMNH, 12 in FMNH, 29 in NMW). (Black morph:) COSTA RICA: Alajuela, Estac. Biol. San Ramon, 900 m, 1.viii.–30.ix.1995, P. Hanson leg. malaise trap (11 in SEMC); Alajuela, Sector Colonia Palmarina, 9 km SO. de Bajo Rodriguez, 700 m, 10.vi.– 2.vii.1997, G. Carballo leg. Malaise (4 in INBC); Prov. Alaju., R.B. San Ramon, 800 m, 21.x.–3.xi.1994, G. Carballo leg. (10 in INBC); Prov. Alaju., R. San Lorencito, 900 m, R.F. San Ramon, 5 km N de Colonia, Palmarena, 13–18.vi.1993 (3 in INBC); Alajuela Prov., Fca. San Gabriel, 2 km SW Dos Rios, 600 m, v.1989 (5 in INBC); Cartago, Cachi, 2 km NE, 1200 m, 1.x.–30.xii.1995, P. Hanson leg. malaise trap (7 in SEMC); Puis Cartago, 1000 m, Fassl leg. (3 in FMNH); Prov. Cartago, Monumento Nacional Guayabo, Turrialba, 1100 m, vii.1994, G. Fonseca leg. (3 in INBC); Farm Cardiella, vii.1938, Bierig leg. (6 in FMNH); Guanacaste, Patilla Biological Station, 610 m, 10o59´22´Ń 85o25´33´´W, 13–15.vii.2000, J.S. Ashe, R. Brooks & Z. Falin leg. FIT (3 in SEMC); Prov. Guanacaste, Rio San Lorenzo, 1050 m, Tierras Morenas, Z.P. Tenorio, iv.1992, F. Quesada leg. (7 in INBC); Guanac. Prov., Estac. Pitillia, 700 m, 9 km S. Santa Cecilia, iii.1989 (76 in INBC); Prov. Guanacaste, Est Cacao, 1000–1400 m, Lado Vol. Cacao Guan., 21–29.v.1992, F. Araya leg. (1 in INBC); Prov. Here, Est. El Celbo, P.N. Braulio Carillo, 400–600 m, ii.1990, C. Chaves leg. (1 in INBC); Heredia, La Selva, 80 m, 19.v.1993, J.S. Ashe leg. FIT (1 in INBC); Heredia, La Selva, 3.2 km SE Puerto Viejo, 100 m, ii–iii.1992, W. Bell leg. FIT (48 in SEMC); Heredia Prov., Puerto Viejo, La Selva, vi–vii.1979, T. Ray leg. plantain-bated trap (1 in MCZC); Limon, 4 km NE Bribri, 50 m, iv.–vi.1990, P. Hanson Coll (1 in SEMC); Prov. Limon, Est. Cuatro Esquinas P.N. Tortuguero, 0 m, v.1990, J. Solano leg. (29 in INBC); Prov. Limon, Sector Cerro Cocori Fcs de E. Rojas, 150 m, 26.iii.–24.iv.1992, F.A. Quesada leg. (2 in INBC); Prov. Limon, Cerro Tortuguero P.N. Tortuguero, 100 m, iv.1989, R. Aguilar & J. Solano leg. (1 in INBC); Orosi (1 in FMNH); Turrialba, 22.vi.1951, O.L. Cartwright leg. (1 in USNM). PANAMA: Chiriqui Prov., La Fortuna ‘Cont. Div. trail’, 8o42Ń 82o14´W, 1150 m, 9–12.vi.1995, J.S. Ashe & R. Brooks leg. FIT (1 in SEMC).

4.36. Nordus testaceus (Fabricius, 1801) (Figs. 6, 102, 245–247) Staphylinus testaceus Fabricius, 1801: 59. Preoccupied (non Staphylinus testaceus Paykull, 1789). Petition to conserve S. testaceus Fabricius, 1801 proposed by HERMAN (2002); approved by ICZN Opionion 2053 (ICZN 2003). Brachydirus testaceus (Fabricius); GEMMINGER & HAROLD (1868: 598). Nordus testaceus (Fabricius); HERMAN (2001: 3581). Description. L e n g t h . 9.5–10 mm (Fig. 6).

44

C o l o r a t i o n . Head golden-orange; mandibles and labrum orange; antennomeres golden-orange, but only 1–5 present. Pronotum, elytra, and scutellum golden-orange. Abdomen black except segment I–II, VIII, and posterior 1/8 of VII golden-orange. Legs golden-orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior 3/4 of epicranium dense, about 5.8 punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior 1/4 about 4.5 punctures / 0.3 mm, punctures almost contiguous. Length of eyes ca. 0.4ⴛ length of head, distance between eyes more than 2ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.02), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres: 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st–5th longer than greatest width; 5th with with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.07; anterolateral angles not prominent, subcariniform; lateral margins almost parallel to each other in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and shallow; medial impunctate area broadest anteriorly (width 1–2 punctures); punctures flanking this area in 2 longitudinal rows separated by width equal to 0.5 or less punctures, punctures of a row not contiguous, but distance varies; large, slightly convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with medium-sized punctures, about 3.6 punctures / 0.3 mm, and interspaces as wide as 1 puncture, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with only few yellow setae on medial disc; V covered with yellow microsetae except for few darker ones medially, VI with tiny patch of yellowish-white microsetae near lateral margins and with darker microsetae medially. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; III–VII with black microsetae. Segment VII with black iridescent setae in anterolateral quarters. Cerci slightly curved, basal half dark brown to black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 102) with evenly curved ventral surface, hind margin divided into two broad lobes and deep, V-shaped emargination medially, each lobe with brush-like patch of prominent, very dense and long setae; sternum IX (identical to that in Fig. 107) with two long apical processes and a shallow emargination medially. Female: unknown. M a l e g e n i t a l i a . As in Figs. 245–247. Paramere in dorsal outline fairly wide, almost parallel-sided though somewhat convex from near base to subapical region, then narrowing to a subacute tip, apically subequal in length to


median lobe and narrower; slightly and irregularly dorsally convex in lateral outline, tip curved downwards and (in contrast to all other species) a bit anteriorly. Sensory spinules (many) arranged in a medial cluster and restricted to tip. Median lobe with broad subtruncate apex in dorsal outline, slightly upcurved apically in lateral outline, forming a small dorsomedial carina, apically without two lobes. Diagnosis. Among the golden-orange species N. diversiventris, N. testaceus, and N. styloceros can be identified by the almost parallel lateral margins of the pronotum. N. testaceus is easily distinguished from the other two species by the unique shape of its aedeagus (Figs. 245 versus 166, 237) and sternum VIII in males and by the medial impunctate area on the pronotum being as wide as 0.5 punctures or less (1–1.5 punctures in the two other species). Distribution. ‘Middle America’. ERICHSON (1839) mentioned the occurrence of N. testaceus in Colombia, but this is likely to be incorrect since he had misidentified specimens in his collection as N. testaceus. Habitat. Unknown. Remarks. Despite the extensive sampling of Central and South America by collectors in the Snow Entomological Collection and the study of almost 3,000 specimens from all over the world, this species is known only from the lectotype and the paralectotype. All previous staphylinid specialists that described species of Nordus and mentioned N. testaceus had misidentified specimens of N. fungicola, N. solitarius, or N. styloceros as N. testaceus. Thus, previous descriptions that compared various species of Nordus with N. testaceus should be treated with caution. Material. Lectotype, ‚, here designated, with labels ‘Type’, ‘S. testaceus, Am. Mer.’, ‘Lectotype Staphylinus testaceus Fabricius des. Chatzimanolis 2003’ (in ZMUC). – Paralectotype, ‚ with labels ‘Type’, ‘Paralectotype Staphylinus testaceus Fabricius des. Chatzimanolis 2003’ (in ZMUC).

4.37. Nordus velitaris (Erichson, 1839) (Figs. 3, 47, 92, 109, 248–251) Staphylinus velitaris Erichson, 1839: 354. Brachydirus velitaris (Erichson); BERNHAUER & SCHUBERT (1914: 405). Brachydirus erosus Bernhauer, 1917: 20; new synonym. Brachydirus schadei Scheerpeltz, 1971: 97; new synonym. Nordus velitaris (Erichson); HERMAN (2001: 3581). Nordus erosus (Bernhauer); HERMAN (2001: 3580). Nordus schadei (Scheerpeltz); HERMAN (2001: 3581). Description. L e n g t h . 10.0–10.5 mm. C o l o r a t i o n . (1) Black morph (previously identified as B. erosus or B. schadei): Head black; mandibles and labrum orange; antennomeres yellow but 6–11 darker yellow. Pronotum black, scutellum dark brown, elytra dark brown to black. Abdomen dark brown to black except segments I– II, VIII, and posterior 1/3 of VII golden-orange. Fore and middle legs light brown; hind legs dark brown except around coxotrochanteral and femorotibial articulations orange. (2) Golden-orange/black morph: Head golden-orange; mandibles and labrum orange; antennomeres yellow but 6– 11 darker yellow. Pronotum and scutellum golden-orange, elytra dark brown to black. Abdomen dark brown to black except segments I–II, VIII, and posterior 1/3 of VII goldenorange. Fore and middle legs golden-orange (Fig. 3); hind legs dark brown except tarsus orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior 3/4 of epicranium dense, about 4.4

45


punctures / 0.3 mm, not elongate longitudinally, margins of punctures confluent; in posterior 1/4 about 3.1 punctures / 0.3 mm, distance between punctures as wide as 0.5 puncture. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium convex; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.06), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 47): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd shorter than 3rd, 5th shorter than 4th, 1st–5th longer than greatest width, 6th wider (posterior margin) than greatest length, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 6th–11th densely covered by microtrichae, 6th–10th with single circumferential row of prominent macrosetae near middle, 1st–5th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.02; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally with distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 1–2 punctures); punctures flanking this area in 3 dense, contiguous longitudinal rows, punctures of a row contiguous; large, strongly convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with light brown microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with whitish-yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with large punctures, about 3.1 punctures / 0.3 mm, punctures contiguous, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV with microsetae on medial disc; V– VI strongly punctuated; IV–VIII covered with yellow microsetae (V–VI with fewer microsetae medially). Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; III–VII with yellow microsetae. Segment VII with black iridescent setae in anterolateral quarters. Cerci slightly curved, basal 1/3 light brown to black and distal 2/3 yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 92) with evenly curved ventral surface, hind margin with broad, moderately deep U-shaped emargination medially; sternum IX (Fig. 109) with two small and wide apical processes and a shallow U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a slightly pointed extension (Fig. 115). M a l e g e n i t a l i a . As in Figs. 248–251. Paramere in dorsal outline very slender, consistently converging to a rounded tip (though distal third fairly parallel-sided), apically subequal in length to median lobe and narrower; dorsally slightly convex in lateral outline. Sensory spinules (many) arranged in a cluster and restricted to tip. Median lobe strongly narrowed to an acute apex in dorsal outline, strongly upcurved in distal half in lateral outline, apex shaped as a single median carina. Diagnosis. Nordus velitaris, N. fungicola, and N. major can be distinguished from other species of Nordus by the combination of a very convex epicranium and antennomere

4 being longer than wide. N. fungicola and N. major differ from N. velitaris by the convex rather than concave lateral margins of the pronotum. Synonymy. Brachydirus erosus Bernhauer and B. schadei Scheerpeltz are identical in all respects, including male sexual structures and male and female morphology. B. velitaris Erichson is identical in all respects with the above two “species” except the coloration as noted above. Distribution. Golden-orange/black morph: known from Brazil (Fig. 67). Black morph: known from elevations of 90–700 m in Argentina, Brazil, and Paraguay (Fig. 67). Habitat. Palm fruitfalls. Remarks. Collected in flight intercept traps. Material. Holotype of S. velitaris, ‚ with labels ‘5889’, ‘Type’, ‘velitaris Er. Brazil-Virra’ (in ZMHB). This is the only specimen in the type series and therefore is the holotype. – Lectotype of B. erosus, ‚, here designated, with labels ‘‚’, ‘S. Catharina Rio Capivary, Fruhstorfer’, ‘erosus Bernh. Typus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Lectotype Brachydirus erosus Bernhauer des. Chatzimanolis 2003’ (in FMNH). – Paralectotype of B. erosus, 1 specimen with labels ‘483’, ‘erosus Für. St. Catharina Standinger’, ‘erosus Bernh. Cotypus’, ‘Chicago NHMuseum M. Bernhauer Collection’, ‘Paralectotype Brachydirus erosus Bernhauer des. Chatzimanolis 2003’ (in FMNH). – Holotype of B. schadei, „ [not ‚ as erroneously mentioned by Scheerpeltz] with labels ‘„’, ‘Villa Rica Paraguay’, ‘Fr. Schade leg. 17.x.1924’, ‘Santa Barbara’, ‘Brachydirus schadei n.sp.m.’, ‘ex. coll. Scheerpeltz’, ‘Typus Brachydirus schadei O. Scheerpeltz’ (in NMW). SCHEERPELTZ (1971) explicitely mentioned that there is only one specimen for his description. However, in his collection there are two specimens, one labeled ‘Typus’ (the specimen above) and another, female one labeled ‘Cotypus’. This specimen was not mentioned in the original description and therefore this is not a valid paratype for this species. – (Goldenorange/black morph:) BRAZIL: Bergland leg. (1 in NMW); Bahia (6 in BMNH); Bahia, Fruhstorfer leg. (1 in FMNH, 2 in NMW); Espirito Santo (2 in BMNH); Guanabo Repress de Rio Grando, iii.1970, M. Alvarenga leg. (1 in AMNH); Represa, Rio Grande, Guanabora, viii.1966, M. Alvarenga leg. malaise trap (1 in NMW). (Black morph:) ARGENTINA: ‘gobhisionas’ (1 in FMNH). BRAZIL: (no labels) (1 in BMNH); Itatiaya, Est. do Bio., 700 m, 16.ii.1924 (1 in FMNH); Minus, Kraatz leg. (1 in DEI); Nova Teutonia, 27o11´S 52o23´W, 300– 500 m, xi.1971, Fritz Plaumann leg. (2 in CNCI); Nova Teutonia, Bierig leg. (4 in FMNH); Nova Teutonia, 10.x.1938, Fritz Plaumann leg. (1 in FMNH); Parana Guanapuava, ii.1961, H. Schneider leg. (1 in CNCI); S. Catharina, Rio Capivary, 1888, Fruhstorfer leg. (12 in BMNH, 14 in DEI, 7 in FMNH, 4 in NMW); S. Catharina, Theresopolis, 1888, Fruhstorfer leg. (2 in BMNH); S. Paulo, Barueri, xii.1965, K. Lenko leg. (1 in CNCI); S. Paulo, Karl Bransik leg. (1 in FMNH); Sao Paulo, Braz. Mraz Mus. Fragenze (1 in FMNH); Kraatz leg. (1 in DEI). PARAGUAY: Itapua, Karoney 17 km W San Rafael Reserve, 26o45´53´´S 55o50´37´´W, 90–110 m, 20.xi.2000, Z. Falin leg. palm fruitfall (1 in SEMC); Itapua, San Pedro Mi, San Rafael Reserve, 26o31´24´´S 55o48´18´´W, 90 m, 23– 30.xi.2000, Z. Falin leg. FIT (1 in SEMC); Itapua, Yatai, prop. Hostettler family, San Rafael Reserve, 100 m, 26o38´17´´S 55o39´50´´W, 21–25.xi.2000, Z. Falin leg. FIT (2 in SEMC); S.W. Salto de Guaira, 10.xii.1971, L. Pena Coll (1 in CNCI).

4.38. Nordus xanthocerus (Nordmann, 1837) (Figs. 57, 103) Brachydirus xanthocerus Nordmann, 1837: 131. Staphylinus xanthocerus (Nordmann); ERICHSON (1839: 353). Nordus xanthocerus (Nordmann); BLACKWELDER (1952: 262). Description. L e n g t h . 9.0–10.5 mm. C o l o r a t i o n . Head black; mandibles and labrum orange; antennomeres all golden-orange. Pronotum, elytra, and scutellum black. Abdomen black except segments I, II light

46

brown; VIII and posterior 1/3 of VII golden-orange. Legs light brown except distal parts of pro- and mesofemora and protarsi orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures on anterior 3/4 of epicranium dense, about 6.2 punctures / 0.3 mm; longitudinally elongate; margins of punctures confluent; on posterior 1/4 about 2.7 punctures / 0.3 mm, punctures separated by a distance as wide as 0.5 punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse microlines and micropunctures. Gula narrow near midlength (least width : length = 0.06), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 57): 1st gradually thickened club-like, but narrower near distal end, subequal in length to 2nd plus 3rd, 2nd subequal in length to 3rd, 5th shorter than 4th, 1st– 5th longer than greatest width, 6th–10th almost quadrate; 6th–11th densely covered by microtrichae, 5th–10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum quadrate, width : length = 1.01; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and shallow; medial impunctate area broadest anteriorly (width 0.5–1 punctures); punctures flanking this area in 2–3 moderately dense longitudinal rows separated by distance as wide as 0.5 puncture, punctures of a row contiguous; large, convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum without arch-like carina, with grayish-yellow microsetae and punctures covering entire surface. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, punctures almost contigous, covered with yellow microsetae. Legs: fore and middle legs with yellow macrosetae, except mesotibiae and -tarsi with brown ones; hind legs with brown macrosetae except coxae, trochanters and anterior parts of femora with orange ones. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII extremely punctuated (IV–V slightly less so medially); IV–V and VIII covered with yellow microsetae, VI and VII covered with yellow ones laterally and black ones medially. Paratergites III–VII covered with whitish-yellow microsetae on posterior half. Sterna: III–VII densely punctuated; III–V and VIII with yellow microsetae, VI and VII with yellow and black ones. Segment VII with black iridescent setae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 103) with evenly curved ventral surface, hind margin with very deep V-shaped notch medially; sternum IX (identical to that in Fig. 107) with two long apical processes and a shallow Ushaped emargination medially. Female: with no obvious secondary sexual structures; hind margin of sternum VIII with no obvious modifications, i.e., not particularly produced and evenly convex (Fig. 114). M a l e g e n i t a l i a . Identical to those in Figs. 190–193. Paramere in dorsal outline almost parallel-sided from middle to subapical region, then narrowed to a subacute tip,


apically subequal in length to median lobe and narrower; completely straight in lateral outline. Sensory spinules (few) arranged in two longitudinal bands close to midline and restricted to tip. Median lobe with narrowed subacute apex in dorsal outline, slightly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two short lobes. Diagnosis. Among species with black head, pronotum and elytra, Nordus maculiceps and N. xanthocerus can be recognized by the shape of sternum VIII: in females it is evenly rounded and not strongly produced at its hind margin (as in Fig. 114), while in all other black species it is either sinuate (as in Fig. 118) or more strongly produced into a lobe (as in Fig. 121); in males sternum VIII has a very deep V-shaped notch (as in Figs. 84, 103), while in all other black species it is as in Figs. 87, 88, 92, or 93. N. maculiceps is very similar to N. xanthocerus but can be distinguished from it by the presence of two bright red spots on the middle of the epicranium and the almost parallel lateral margins of the pronotum (margins concave in N. xanthocerus); also, in N. xanthocerus the pronotum is more densely punctuated than in N. maculiceps. Distribution. Known from Brazil and Ecuador (Fig. 21). There are no specimens known from Bolivia and Peru as erroneously mentioned in SCHEERPELTZ (1971) and HERMAN (2001). Habitat. Unknown. Material. Lectotype, ‚, here designated, with labels ‘Type’, ‘Hist. Coll. – (Coleoptera) Nr. 5888 Staphylinus xanthocerus Nordm. Bahia Sello Zool. Mus. Berlin’, ‘Lectotype Brachydirus xanthocerus Nordmann des. Chatzimanolis 2002’ (in ZMHB). The second label above is incorrect; NORDMANN (1837) described this species in Brachydirus and it was ERICHSON (1839) who transferred it to Staphylinus before KRAATZ (1857) re-established the genus Brachydirus. No paralectotypes are designated because the remaining specimens in the types series belong to N. maculiceps. – (No locality labels), 10.x.1938, ex. coll. Scheerpeltz (2 in NMW). BRAZIL: Espirito Santo, A. Heyne Bern-Wilm leg. (2 in FMNH); Rio de Janeiro, Guanabra, xii.1967, M. Alvarenga leg. (1 in AMNH); Rio de Janeiro, Sahlber leg. (1 in FMNH); Rondonia, 62 km S Ariquemes Faz Pancho Grandes, 10o32´S 62o48´W, 5–15.x.1993, C. & K. Messenger leg. (1 in UNSM). ECUADOR: Pichincha, 6 mi E Santo Domingo de los Colorados, 24.ii.1955, E.I. Schlinger & E.S. Ross leg. (2 in CASC).

4.39. Nordus zischkai (Scheerpeltz, 1971) (Figs. 8, 48, 93, 110, 252–255) Brachydirus zischkai Scheerpeltz, 1971: 96. Nordus zischkai (Scheerpeltz); HERMAN (2001: 3582). Description. L e n g t h . 9.0–10.5 mm. C o l o r a t i o n . Head black; mandibles and labrum dark brown; antennomeres 1–2 dark yellow, 3–6 dark brown, 7–9 brownish-yellow, 10–11 light to dark brown. Pronotum, elytra, and scutellum black. Abdomen black except segments VIII and posterior 1/3 of VII golden-orange. Legs black except distal parts of pro- and mesofemora and -tarsi orange. H e a d . Microsetae yellow. Anteclypeus very small. Punctures in anterior half of epicranium dense, about 5.3 punctures / 0.3 mm; longitudinally elongate; margin of punctures confluent; in posterior half about 3.5 punctures / 0.3 mm, distance between punctures as wide as 0.5–1 punctures. Length of eyes ca. 0.6ⴛ length of head, distance between eyes ca. 1.5ⴛ length of eye. Middle of epicranium flat; epicranium without microsculpture of transverse

47


microlines and micropunctures. Gula narrow near midlength (least width : length = 0.05), gular sulci widely separated anteriorly and almost contiguous near midlength and posteriorly. Antennomeres (Fig. 48): 1st thickened only in distal half, but narrower near distal end, longer than 2nd plus 3rd, 2nd subequal in length to 3rd, 5th subequal in length to 4th, 1st–3rd longer than greatest width, 4th–6th subquadrate, 7th–10th wider than greatest length, width of distal margin of 9th and 10th at least 2ⴛ length; 7th–11th and distal part of 6th densely covered by microtrichae, 5th– 10th with single circumferential row of prominent macrosetae near middle, 1st–4th with irregularly scattered macrosetae. T h o r a x . Pronotum subquadrate, width : length = 0.86; anterolateral angles not prominent, subcariniform; lateral margins concave in dorsal aspect; disc laterally without distinct gaps in punctuation; punctures large and deep; medial impunctate area broadest anteriorly (width 0.5–1 punctures); punctures flanking this area in 1–2 sparse longitudinal rows separated by distance as wide as 0.5–1 punctures, punctures of a row not contiguous, but distance varies; large, convex, raised impunctate area in anterolateral quarters, and large impunctate area along posterior and posterolateral margins. Pronotum with yellow microsetae. Metepimeron with few sparse punctures anteriorly, not in rows. Metepisternum with punctures forming more than 2 rows. Scutellum with shallow arch-like carina, with grayish-yellow microsetae and punctures covering posterior 2/3. Elytra not depressed near scutellum, with medium-sized punctures, about 3.1 punctures / 0.3 mm, and interspaces as wide as 0.5–1 punctures, covered with yellow microsetae. Legs with yellow macrosetae. A b d o m e n . Terga: III with 0–4 discal microsetae; V without subbasal carina; IV–VII well punctuated (V slightly less so medially); IV with yellowish-brown microsetae medially and posterolaterally, V with brown ones except posterolaterally, VI with yellow ones on posterolateral corners and anteromedially, VII with brown ones. Paratergites IV–VII covered with whitish-yellow and black microsetae on posterior half. Sterna: VI–VII on anterolateral corners without or with sparse punctures; III–VII densely punctuated otherwise (IV–VI slightly less so posteromedially); III–VI and VIII with brownish-yellow microsetae, VII with brown ones. Segment VII with black iridescent setae throughout. Cerci slightly curved, basal half black and distal half yellow. S e c o n d a r y s e x u a l s t r u c t u r e s . Male: hind margin of sternum VII (identical to that in Fig. 62) with broad and deep emargination medially; sternum VIII (Fig. 93) with evenly curved ventral surface, hind margin with two moderate triangular lobes and a strong V-shaped emargination medially; sternum IX (Fig. 110) with two moderate apical processes and a moderate U-shaped emargination medially. Female: with no obvious secondary sexual structures except hind margin of sternum VIII forming a slightly pointed extension (Fig. 115). M a l e g e n i t a l i a . As in Figs. 252–255. Paramere in dorsal outline consistently converging to a subacute tip, apically subequal in length to median lobe and narrower; slightly dorsally convex in lateral outline. Sensory spinules (many) arranged in a medial distal cluster and a pair of far lateral rows extending basally from it, all restricted to tip. Median lobe with broadly subacute apex in dorsal outline, weakly upcurved apically in lateral outline, forming a dorsomedial carina forked apically into two tiny lobes.

Diagnosis. Nordus antennatus, N. banghaasi, N. picticornis, N. aeneipennis, and N. zischkai can all be distinguished from other species of Nordus by the following characters: punctures on anterolateral corners of sterna VI and VII less dense than on the rest of these sterna; punctures on anterior part of epicranium longitudinally elongate; and shape of sternum VIII in males (shape varies among these species, but overall is distinctive for this group of species). N. antennatus and N. banghaasi can be distinguished from N. picticornis, N. aeneipennis, and N. zischkai by antennomere 4 being longer than wide (subquadrate in other species). In N. picticornis head, pronotum, and elytra are golden-orange, whereas in N. aeneipennis and N. zischkai head and pronotum are reddish-brown to black and elytra are dark brown to black. N. aeneipennis can by distinguished from N. zischkai by the following characters: color of head and pronotum reddish-brown and elytra dark brown (all black in N. zischkai); punctures on anterior part of the epicranium denser in N. aeneipennis than in N. zischkai; punctures on posterior part of epicranium shallow in N. aeneipennis and deep in N. zischkai; and in males the shape of the aedeagus is different (Figs. 134–137 for N. aeneipennis and Figs. 252–255 for N. zischkai). Distribution. Unknown from elevations of 730–1800 m in Bolivia (Fig. 21). Habitat. Not known. Remarks. SCHEERPELTZ (1971) mentioned in the description that he had two specimens from this species, a male holotype and a female allotype. However, only the female specimen was found and that one is therefore designated as the neotype. The neotype is from the type locality and agrees in every respect with Scheerpeltz’s description. Specimens were collected in dung traps and flight intercept traps. Material. Neotype, „, here designated, with labels ‘„’, ‘Bolivia: Yungas 1000 m 2.v.1948 leg. R. Zischka’, ‘ex. coll. Scheerpeltz’, ‘Typus Brachydirus zischkai O. Scheerpeltz’, ‘zischkai Scheerp.’, ‘Neotype Brachydirus zischkai Scheerpeltz des. Chatzimanolis 2003’ (in NMW). – BOLIVIA: Cochabamba Province, Cochabamba, 109 km E Yungas, Cochabamba–Villa Tunari Rd., 1480 m, 17o8´50´´S 65o42´29´´W, 01.–06.ii.1999, 6 specimens R. Hanley leg. and 9 specimens F. Genier leg. (15 in SEMC); Cochabamba Province, Cochabamba, 124 km E Yungas, Cochabamba–Villa Tunari Rd., 730 m, 17o3´54´´S 65o38´43´´W, 06.–08.ii.1999, R. Hanley leg. FIT (1 in SEMC); Cochabamba Province, Cochabamba, 117 km E Yungas, Cochabamba–Villa Tunari Rd., 1040 m, 17o6´32´´S 65o41´12´´W, 01.–06.ii.1999, R. Hanley leg. FIT (2 in SEMC); Cochabamba Province, Cochabamba, 105 km E Yungas nr. Rio Carmen Mayu, Cochabamba–Villa Tunari Rd., 1800 m, 17o8´47´´S 65o43´55´´W, 06.ii.–08.vi.1999, F. Genier leg. dung trap (3 in SEMC); Coroico (2 in BMNH); Songo (1 in BMNH).

4.40. Keys to species of Nordus Below I provide two keys for the identification of Nordus. The first key (A) uses heavily male-specific and coloration characters and can reliably identify male specimens; females can be identified for some species, for the rest they can be identified only through association with a male. Readers are strongly encouraged to use key (A); both male sexual characters and coloration (with the exception of the polymorphic species) show no to minimum variability within species. If the identification of some specimens is problematic, or female specimens must be identified, then the second key (B) can be used. It has been written with minimum reference to coloration characters and almost no

48

reference to sexual characters. Thus, only when there are no other morphological characters that can distinguish two species, there will be reference to coloration and/or male or female sexual characters. It should be mentioned that for almost all species, there are differences in the male genitalia and sterna VII–VIII, and readers are encouraged to look up those figures in addition to the characters provided in key (B).

Key A 1 Anterior part of head metallic green. 2 1’ Anterior part of head golden-orange, brown, or black. 4 2 (1) Head completely metallic green; pronotum and elytra metallic green; shape of aedeagus as in Figs. 234–236. N. speciosus 2’ (1) Head golden-orange with anterior part metallic green; pronotum and elytra golden-orange, shape of aedeagus as in Figs. 122–125, 226–229. 3 3 (2) Length 7 mm; legs and thoracic sterna mostly black; aedeagal paramere in dorsal aspect with pointed tip (Fig. 125). N. aeneiceps 3’ (2) Length 9.5–11 mm; legs and thoracic sterna goldenorange; aedeagal paramere in dorsal aspect with rounded tip (Fig. 229). N. simplex 4 (1) Pronotum brown to black. 5 4’ (1) Pronotum golden-orange. 15 5 (4) Abdomen dark orange to light brown. 6 5’ (4) Abdomen black except hind margin of sternum VII and segment VIII. 7 6 (5) Head and pronotum dark brown to black; middle of epicranium with two bright red spots; tergum V with subbasal carina; sternum VIII of males as in Fig. 98; aedeagus as in Figs. 214–217. N. portokalis 6’ (5) Head and pronotum light red-brown; middle of epicranium without red spots; tergum V without subbasal carina; sternum VIII of males as in Fig. 69; aedeagus as in Figs. 134–137. N. aeneipennis 7 (5) Antenna yellow, golden-orange, or darker orange; sternum VIII of males without lobes on hind margin, medial emargination of varied shape and size but never a very narrow cleft (Figs. 79, 84, 92, 103). 8 7’ (5) Antenna with some antennomeres yellow or golden orange and others brown or black; sternum VIII of males with two moderate or strong triangular lobes on hind margin, medial emargination of varied shape and size (Figs. 72, 76, 93), sometimes a very narrow cleft (the two lobes together then considered to constitute a single lobe deeply split medially; Figs. 87, 88). 11 8 (7) Middle of epicranium slightly to strongly convex; sternum VIII of males with a narrow and shallow V-shaped emargination (Fig. 79) or with a broad and moderately deep U-shaped emargination (Fig. 92). 9 8’ (7) Middle of epicranium flat; sternum VIII of males with a very deep V-shaped emargination (Figs. 84, 103). 10 9 (8) Middle of epicranium very strongly convex; sternum VIII of males with broad and moderately deep U-shaped emargination (Fig. 92); aedeagus distinctive (Figs. 252– 255). N. velitaris (in part) 9’ (8) Middle of epicranium slightly convex; sternum VIII of males with a narrow and shallow V-shaped emargination (Fig. 79); aedeagus as in Figs. 162–165. N. dichromos (in part) 10 (8) Middle of epicranium with two bright red spots; lateral margins of pronotum almost parallel to each other. N. maculiceps


10’ (8) Middle of epicranium without red spots; lateral margins of pronotum concave in dorsal aspect. N. xanthocerus 11 (7) Sternum VIII of males with two very strong triangular lobes, medial emargination very deep and narrow, its inner margins essentially parallel-sided (the two lobes considered to constitute one large lobe split medially; Figs. 87, 88); with brush-like patch of prominent setae on the basal part of each side of emargination; paramere with middle part distinctly widened, disc-like (Figs. 202, 241). 12 11’ (7) Sternum VIII of males with two moderate or strong triangular lobes, medial emargination lower and wider, Vor U-shaped (Figs. 72, 76, 93); without brush-like patch of prominent setae on the basal part of each side of emargination; paramere with middle part not widened and disc-like (Figs. 134, 150, 252). 13 12 (11) Paramere tip in lateral view almost straight. Sensory spinules along distal 1/6 of lateral sides of paramere. Apex of median lobe continuously narrowed, subacute (Figs. 241–244). N. terminalis (in part) 12’(11) Paramere tip in lateral view strongly down-curved. Sensory spinules along distal 1/10 of lateral sides of paramere. Apex of median lobe slightly widened and broadly rounded (Figs. 202–205). N. omios 13 (11) Tergum V with subbasal carina; aedeagal paramere in dorsal outline distally widened, with a broad subtruncate tip produced to a small point medially; apex of median lobe with 2 broad, widely separated teeth, the base of each forming an almost 90o angle with the distal lateral margin of the median lobe (Figs. 150–153). N. cherylae 13’(11) Tergum V without subbasal carina; aedeagal paramere in dorsal outline not widened, with tip neither subtruncate nor produced to a small point medially; apex of median lobe forming a small medial tooth-like carina separated at the apex into two short lobes. 14 14(13) Antennomeres 1–2 dark yellow, 3–6 dark brown, 7– 9 brownish-yellow and 10–11 light to dark brown; sternum VIII of males with two moderate triangular lobes and with strong V-shaped emargination medially (Fig. 93); aedeagal paramere with pointed tip (Fig. 255). N. zischkai 14’ (13) Antennomeres 1–5 dark yellow, 6–9 whitish-yellow and 10–11 brown; sternum VIII of males with two strong triangular lobes and with shallow broad U-shaped emargination medially (Fig. 72); aedeagal paramere with rounded tip (Fig. 137). N. antennatus 15 (4) Head brown; elytra dark orange to brown; aedeagus as in Figs. 170–173. N. elegans 15’ (4) Head completely golden-orange, or golden-orange with anterior part shining black; elytra golden-orange or dark brown to black; aedeagus not as above. 16 16 (15) Elytra dark brown to black. 17 16’ (15) Elytra golden-orange. 18 17 (16) Middle of epicranium flat; sternum VIII of males or with a deep and very narrow emargination (Fig. 88) medially; with brush-like patch of prominent setae on basal part of each side of emargination; aedeagus as in Figs. 241–244. N. terminalis (in part) 17’ (16) Middle of epicranium strongly convex; sternum VIII of males with broad and moderately deep U-shaped emargination (Fig. 92); without brush-like patch of prominent setae on basal part of each side of emargination; aedeagus as in Figs. 248–251. N. velitaris (in part) 18 (16) Scutellum dark brown to black. N. nigroscutellatus 18’ (16) Scutellum golden-orange. 19 19 (18) Head entirely golden-orange. 20 19’ (18) Head golden-orange with anterior part shining black. 37


20 (19) Hind femur and tibia dark brown to black; sternum VIII of males as in Fig. 90; aedeagus distinctive as in Figs. 230–233. N. solitarius 20’ (19) Hind femur and tibia yellow to golden-orange; sternum VIII of males and aedeagus not as above. 21 21 (20) Anteclypeus well developed; temporal area posterior to eyes greatly expanded; pronotum with uniform punctuation almost throughout; lateral margins of pronotum strongly convex in dorsal aspect; aedeagus distinctive as in Figs. 194–197. N. major 21’ (20) Anteclypeus very small; temporal area posterior to eyes not expanded; punctuation on pronotum not uniformly distributed; lateral margins of pronotum concave in dorsal aspect; aedeagus not as above. 22 22 (21) Sternum VII completely black, tergum VII black 23 with less than posterior 1/8 golden-orange. 22’ (21) Sternum and tergum VII black with posterior 1/4–1/2 golden-orange. 24 23 (22) Antennomeres 4–5 longer than wide; lateral margins of pronotum almost parallel to each other in dorsal aspect; sternum VIII of males very distinctive, divided into two broad lobes and with V-shaped emargination medially, distal edge of each lobe with brush-like patch of densely placed, prominent, long setae (Fig. 102); aedeagus distinctive as in Figs. 245–247. N. testaceus 23’ (22) Antennomeres 4–5 subquadrate; lateral margins of pronotum concave in dorsal aspect; sternum VIII of males with broad and moderate U-shaped emargination medially (Fig. 81); hind margin lateral to emargination without such setation; aedeagus as in Figs. 178–181. males of N. facialis 24 (22) Sternum VIII of males with evenly curved ventral surface, with strong triangular lobes and with a very deep and narrow, essentially parallel-sided emargination medially; with brush-like patch of prominent setae on basal part of each side of emargination (Fig. 88); aedeagus as in Figs. 241–244. N. terminalis (in part) 24’ (22) Sternum VIII of males with evenly curved ventral surface, with two small, moderate or strong triangular lobes extending beyond the margin of the sternum (i.e. hind margin lateral to emargination distinctly concave) and with a shallow to moderately deep emargination medially; without brush-like patch of prominent setae on basal part of each side of emargination (Figs. 73, 78, 80, 95, 96, 97, 100). 26 24’’ (22) Sternum VIII of males with evenly curved ventral surface, without lobes (i.e. hind margin lateral to emargination straight or convex) and with a shallow, moderate or strong U- or V-shaped emargination medially; without brush-like patch of prominent setae on basal part of each side of emargination (Figs. 75, 77, 79, 82, 83, 99). 32 24’’’ (22) Sternum VIII of males not with evenly curved ventral surface but rather elevated along the middle, forming a carina, distal end of carina continuing into a discrete lobe upon the hind margin (potentially forked); without brush-like patch of prominent setae around base of lobe (Figs. 70, 91). 25 25 (24) Hind margin of sternum VII of males (Fig. 61) with small truncate lobe extending across midline, with series of black prominent setae at its base; lobe arising from distal end of carina of sternum VIII (Fig. 70) not medially divided and hook-like (upcurved) distally; aedeagus as in Figs. 126–129. N. amazonicus (in part) 25’ (24) Hind margin of sternum VII (Fig. 65) with very strong notch at midline and prominent setae around margin of notch; lobe arising from distal end of carina of sternum VIII (Fig. 91) medially deeply divided, fork-like; aedeagus as in Figs. 237–240. N. styloceros

49

26 (24) Sternum VIII of males with two small to very small triangular lobes extending beyond hind margin of sternum and with a moderate U-shaped or small V- or U-shaped emargination medially (Figs. 78, 80, 95). 27 26’ (24) Sternum VIII of males with two strong triangular lobes extending beyond hind margin of sternum and with a shallow to moderate U-shaped or very deep V-shaped emargination (Figs. 73, 96, 97, 100). 29 27 (26) Distance between eyes subequal to 2ⴛ length of eye; eyes placed more anteriorly on head; sternum VIII of males with two small triangular lobes and a moderate U-shaped emargination medially (Fig.78); aedeagus as in Figs. 158–161. N. densiventris 27’ (26) Distance between eyes subequal to 1.5ⴛ length of eye, eyes placed more posteriorly on head; sternum VIII of males with two very small triangular lobes and a small Vor U-shaped emargination medially (Figs. 80, 95). 28 28 (27) Antennomeres 1–8 yellow, 9–10 darker orange to brown; aedeagal paramere in dorsal view not very wide, as in Figs. 166–169. N. diversiventris 28’ (27) Antennomeres 1–10 yellow; aedeagal paramere in dorsal view very wide, as in Figs. 174–177. N. elytisi 29 (26) Medial emargination of sternum VIII of males shallow to moderate, U-shaped (Figs. 73, 97). 30 29’ (26) Medial emargination of sternum VIII of males very deep, V-shaped (Figs. 96, 100). 31 30 (29) Antennomeres 1–6 yellow, 7–11 whitish-yellow; distal margin of sternum VIII of males (Fig. 73) with a shallow to moderate broad U-shaped emargination medially. N. banghaasi 30’ (29) Antennomeres 1–4 yellow, 5–9 and distal part of 11 pale yellow, 10 and basal part of 11 dark brown to black; distal margin of sternum VIII of males (Fig. 97) with a moderate U-shaped emargination medially. N. picticornis 31 (29) Aedeagus as in Figs. 222–225. N. semelius 31’ (29) Aedeagus as in Figs. 206–209. N. periergos 32 (24) Pronotum very densely punctuated (as in Fig. 15), lateral margins very concave in dorsal aspect. 33 32’ (24) Pronotum less densely punctuated, lateral margins convex or weakly concave in dorsal aspect. 35 33 (32) Middle of epicranium convex; antennomeres 1–6 yellow, 7–11 darker yellow, 6 almost 2ⴛ as wide as greatest length; aedeagus as in Figs. 154–157 or 218–221. 34 33’ (32) Middle of epicranium flat; antennomeres 1–11 uniformly yellow, 6 slightly wider than long; aedeagus as in Figs. 186–189. N. gebieni 34 (33) Aedeagus as in Figs. 154–157, paramere very slender. N. cribricollis 34’ (33) Aedeagus as in Figs. 218–221, paramere very wide. N. seferisi 35 (32) Lateral margins of pronotum concave in dorsal aspect; pronotum sparsely punctuated; sternum VIII of males with narrow and shallow V-shaped emargination medially (Fig. 79); aedeagus as in Figs. 162–165. N. dichromos (in part) 35’ (32) Lateral margins of pronotum slightly to strongly convex in dorsal aspect; pronotum more densely punctuated; sternum VIII of males with broad and moderately deep U-shaped emargination medially (Figs. 75, 82). 36 36 (35) Pronotum moderately to strongly convex; tergum IV with less than 5 discal microsetae. N. fungicola 36’ (35) Pronotum slightly convex; tergum IV with multiple discal microsetae. N. championi 37 (19) Sternum VII completely black, tergum VII black with less than posterior 1/8 golden-orange. females of N. facialis

50

37’ (19) Sternum and tergum VII black with posterior 1/ –1/ golden-orange. 38 4 3 38 (37) Middle of epicranium slightly to strongly convex; antennomeres 2–10 light brown to black. 39 38’ (37) Middle of epicranium flat; antennomeres 2–5 yellow to golden-orange, 8–10 dark brown to black. 41 39 (38) Punctures on pronotum sparse and shallow; punctuation on anterior part of head very dense; antennomere 7 wider than long. N. stomachoponos 39’ (38) Punctures on pronotum dense and deep; punctuation on anterior part of head not very dense; antennomere 7 subquadrate. 40 40 (39) Antennomeres 5 and 6 quadrate, 2–11 dark brown to black; only females known. N. longipes 40’ (39) Antennomeres 5 and 6 longer than wide, 2–10 light brown to black, 11 pale yellow; aedeagus as in Figs. 130–133. N. amphivolos 41 (38) Antennomeres 1–4 whitish-yellow, 7–11 black; sternum VII of males with broad and deep emargination medially; sternum VIII of males evenly rounded, with single broad lobe, rounded at the subapical region (Fig. 74); aedeagus as in Figs. 142–145. N. batesi 41’ (38) Antennomeres 1–7 yellow, 8–10 black, 11 basally black and distally yellow; sternum VII of males (Fig. 61) with small lobe extending across the midline, with series of black prominent setae at its base; sternum VIII of males (Fig. 70) not evenly rounded but elevated towards the middle, forming a carina, distal end of carina continuing into a discrete lobe upon the hind margin; aedeagus as in Figs. 126–129. N. amazonicus (in part)

Key B 1 Head, pronotum and elytra metallic green. N. speciosus 1’ Head, pronotum, and elytra dark brown to black or golden-orange. 2 2 (1) Lateral margins of pronotum convex. 3 2’ (1) Lateral margins of pronotum concave or almost parallel to each other. 4 3 (2) Anteclypeus very small. N. fungicola 3’ (2) Anteclypeus well developed. N. major 4 (2) Tergum V with subbasal carina. 5 4’ (2) Tergum V without subbasal carina. 7 5 (4) Pronotum subquadrate; abdomen dark orange to light brown. N. portokalis 5’ (4) Pronotum longer than wide; abdomen black. 6 6 (5) Head dark brown, pronotum golden-orange and elytra dark orange to brown. In males aedeagus as in Figs. 170– 173. In females sternum VIII with evenly convex hind margin (as in Fig. 114). N. elegans 6’ (5) Head, pronotum and elytra black. In males aedeagus as in Figs. 150–153. In females sternum VIII with sinuate hind margin (as in Fig. 118). N. cherylae 7 (4) Tergum V with 0–4 microsetae on medial disc. 8 7’ (4) Tergum V with >5 microsetae on medial disc. 11 8 (7) Punctures on elytra very sparse (with interspaces of more than 1ⴛ width of punctures); elytra depressed near scutellum. 9 8’ (7) Punctures on elytra denser (punctures confluent or with interspaces of 0.5–1ⴛ width of punctures); elytra not depressed near scutellum. 10 9 (8) Paramere tip in lateral view almost straight. Sensory spinules along distal 1/6 of lateral sides of paramere. Apex of median lobe continuously narrowed, subacute (Figs. 241–244). N. terminalis


9’ (8) Paramere tip in lateral view hook-like. Sensory spinules along distal 1/10 of lateral sides of paramere. Apex of median lobe slightly widened and broadly rounded (Figs. 202–205). N. omios 10 (8) Antennomere 4 longer than wide, 5 longer than 6; epicranium convex; medial impunctate area on pronotum as wide as 2–3 punctures; along flanks of medial impunctate area punctuation less dense; punctures on elytra dense N. dichromos (interspaces 0.5–1ⴛ width of punctures). 10’ (8) Antennomere 4 subquadrate, 5 subequal in length to 6; epicranium rather flat; medial impunctate area on pronotum as wide as 0.5 punctures; along flanks of medial impunctate area punctuation dense; punctures on elytra very dense (confluent). N. solitarius 11 (7) Antennomere 5 longer than wide. 12 11’ (7) Antennomere 5 subquadrate. 14 12 (11) Antennomere 6 longer than wide. N. amazonicus 12’ (11) Antennomere 6 subquadrate. 13 13 (12) Antennomere 7 wider than long; punctures on anterior part of epicranium not longitudinally elongate; punctures on posterior part of epicranium deep; punctures on pronotum flanking medial impunctate area dense; pronotum with strongly raised impunctate area in anterolateral quarters; lateral margins of pronotum almost parallel to each other; punctures on elytra dense (interspaces 0.5–1ⴛ width of punctures). N. diversiventris 13’ (12) Antennomere 7 subquadrate; punctures on anterior part of epicranium longitudinally elongate; punctures on posterior part of epicranium shallow; punctures on pronotum flanking medial impunctate area less dense; pronotum with moderately raised impunctate area in anterolateral quarters; lateral margins of pronotum concave; punctures on elytra very dense (confluent). N. xanthocerus 14 (11) Lateral margins of pronotum almost parallel to each other. 15 14’ (11) Lateral margins of pronotum concave. 17 15 (14) Punctures on anterior part of epicranium longitudinally elongate; punctures on posterior part of epicranium shallow; punctures on pronotum flanking medial impunctate area less dense. N. maculiceps 15’ (14) Punctures on anterior part of epicranium not longitudinally elongate; punctures on posterior part of epicranium deep; punctures on pronotum flanking medial impunctate area dense. 16 16 (15) Medial impunctate area on pronotum as wide as 1– 1.5 punctures; pronotum with strongly raised impunctate area in anterolateral quarters. N. styloceros 16’ (15) Medial impunctate area on pronotum as wide as 0.5 puncture or less; pronotum with moderately raised impunctate area in anterolateral quarters. N. testaceus 17 (14) Antennomere 6 wider than long. 18 17’ (14) Antennomere 6 subquadrate. 22 18 (17) Antennomere 4 longer than wide. N. cribricollis 18’ (17) Antennomere 4 subquadrate. 19 19 (18) Microsculpture of transverse microlines on epicranium absent. 20 19’ (18) Microsculpture of transverse microlines on epicranium present. 21 20 (19) Punctures on pronotum flanking medial impunctate area very dense. N. seferisi 20’ (19) Punctures on pronotum flanking medial impunctate area less dense. N. elytisi 21 (19) Punctures on anterior part of epicranium not longitudinally elongate. N. periergos 21’ (19) Punctures on anterior part of epicranium longitudinally elongate. N. semelius

51


22 (17) Anterior part of epicranium metallic green, posterior part golden-orange. 23 22’ (17) Anterior and posterior part of epicranium goldenorange or dark brown to black. 24 23 (22) Length 7 mm; cerci in both sexes straight. N. aeneiceps 23’ (22) Length 9.5–11 mm; cerci in both sexes slightly curved. N. simplex 24 (22) Middle of epicranium convex. 25 24’ (22) Middle of epicranium flat. 30 25 (24) Antennomere 7 subquadrate. 26 25’ (24) Antennomere 7 longer than wide. 27 26 (25) Antennomere 5 subequal in length to 6. N. longipes 26’ (25) Antennomere 5 longer than 6. N. amphivolos 27 (25) Antennomere 5 longer than 6. N. velitaris 27’ (25) Antennomere 5 subequal in length to 6. 28 28 (27) Antennomere 4 longer than wide; punctures on pronotum shallow. 29 28’ (27) Antennomere 4 subquadrate; punctures on pronotum deep. N. facialis 29 (28) Pronotum with strongly raised impunctate area in anterolateral quarters; medial impunctate area on pronotum as wide as 1 puncture; punctures on pronotum flanking medial impunctate area dense. N. stomachoponos 29’ (28) Pronotum with moderately raised impunctate area in anterolateral quarters; medial impunctate area on pronotum as wide as 3–4 punctures; punctures on pronotum flanking medial impunctate area less dense. N. championi 30 (24) Punctures on anterior part of epicranium not longitudinally elongate. 31 30’ (24) Punctures on anterior part of epicranium longitudinally elongate. 34 31 (30) Antennomere 4 longer than wide. 32 31’ (30) Antennomere 4 subquadrate. 33 32 (31) Scutellum dark brown to black. N. nigroscutellatus 32’ (31) Scutellum golden-orange. N. gebieni 33 (31) Antennomere 5 subequal in length to 6; microsculpture of transverse microlines on epicranium absent; lateral margins of pronotum strongly concave. N. batesi 33’ (31) Antennomere 5 longer than 6; microsculpture of transverse microlines on epicranium present; lateral margins of pronotum concave. N. densiventris 34 (30) Antennomere 4 longer than wide. 35 34’ (30) Antennomere 4 subquadrate. 36 35 (34) Head, pronotum and elytra golden-orange. N. banghaasi 35’ (34) Head, pronotum and elytra dark brown to black. N. antennatus 36 (34) Antennomere 5 subequal in length to 6. 37 36’ (34) Antennomere 5 longer than 6. N. picticornis 37 (36) Punctures on posterior part of epicranium deep; lateral margins of pronotum slightly concave. N. zischkai 37’ (36) Punctures on posterior part of epicranium shallow; lateral margins of pronotum strongly concave. N. aeneipennis

5. Designation of lectotypes and neotypes and synonymies Lectotypes were selected for: Brachydirus aeneipennis Bernhauer, B. antennatus Sharp, B. banghaasi Bernhauer, B. batesi Sharp, B, championi Sharp, B. cribricollis Sharp, B. densiventris Bernhauer, B. divisus Sharp, B. erosus Bernhauer, B. fungicola Sharp, B. gebieni Bernhauer, B. maculiceps Sharp, B. major Bernhauer, B. occipitalis

Bernhauer, B. simplex Sharp, B. solitarius Sharp, B. styloceros Sharp, B. xanthocerus Nordmann, and Staphylinus testaceus Fabricius. Neotypes were designated for B. picticornis Solsky and B. zischkai Scheerpeltz. Lectotype designations are justified under article 74.1.1 and neotypes designations under article 75.3 of the ICZN (1999) in order to fix the status of these specimens as the sole namebearing type of the respective species and for the expressed purpose of nomenclatural stability.

6. Discussion There are three species of Nordus (N. dichromos, N. terminalis and N. velitaris) with different color morphs. These three species are not closely related (see CHATZIMANOLIS in press b). I have chosen to describe these morphs as one species for N. dichromos, and to synonymize the existing species names for each color morph for N. velitaris and N. terminalis. The color morphs thereby assigned to the same species have identical morphology, including genitalia and sternum VIII in males. In addition, there are few specimens of intermediate coloration (brown or brownish-orange) for each of these species, which might be an indication of mating between different color morphs. It should be mentioned here, that all the different color morphs of N. terminalis (former names N. terminalis, N. gracilis, and N. divisus) and N. velitaris (former names N. velitaris and N. erosus) were described based on one or very few specimens and that male specimens were often not available for all “species” for comparisons of the genitalia, and this might explain why they were described as different species. In N. terminalis all color morphs are allopatric, and based on the available limited data the black morph and the golden-orange/black morph are more frequently found on higher elevations (above 800 m), while the orange morph is found on lowlands. The same pattern can be observed in N. dichromos, where almost all specimens belonging to the black morph are found on higher elevations, while specimens belonging to the orange morph are mostly found in the lowlands. For N. dichromos, based on the available data, it is not clear how much geographically separated the black and orange morphs are, besides the difference in the elevation. Finally, the black morph and the golden-orange morph of N. velitaris are allopatric but most specimens were collected without elevation data and thus elevation differences are not as clear for this species as for the other two. It is important to mention that all dark colored species in the genus with elevation data known (N. aeneipennis, N. antennatus, N. cherylae, and N. zischkai) are distributed from lowlands to elevations above 1000 m, whereas the majority of golden-orange species are distributed in the lowlands. Nordus is unusual among members of Xanthopygina in having strong sexual dimorphism in non-genitalic parts of the post-abdomen including sternum VII, VIII, and IX in males and sternum VIII in females, and these parts vary greatly among species. The modifications of the male sternum VIII are particularly interesting (Figs. 69–103), and sternum VIII appears to be slightly to dramatically different in all 36 species for which males are known. The biology of these beetles is completely unknown (except for N. fungicola, see CHATZIMANOLIS 2003) and thus only speculations can be made on the evolutionary role of this structure and why it is so variable. There are a couple of hypotheses to explain the function of these structures. Firstly, in N. fungicola males bend sterna VII–VIII ventrally

52

to mate and thus the sexually selected modifications might assist in mating. The medial emargination of the hind part of sterna VII– VIII in N. fungicola supports this hypothesis; however, there are species (e.g., N. omios, N. styloceros, N. terminalis, and others) where the distal part of sternum VIII is not emarginate but rather elongate, and seems unlikely to facilitate the ventral bending of the abdomen. Another hypothesis has to do with the waving of the abdomen observed in N. fungicola (see CHATZIMANOLIS 2003). Individuals of N. fungicola raise their abdomens over their backs and wave them antero-posteriorly four or five times, followed by a slower side to side movement, and prominently display the brilliant yellow apex of the abdomen. It is likely that males display sternum VIII to females or competitor males in a sexual competition; however females also wave their abdomens but appear to have no interspecific variation in these structures. The forthcoming phylogenetic analysis of Nordus (CHATZIMANOLIS in press b) might provide insight into the evolution of coloration and postabdominal sterna, but more field observations on other species of Nordus are needed to explain their function. The secondary sexual structures on abdominal sterna of Nordus present us many questions on function and design and deserve more attention by evolutionary biologists.

7. Acknowledgments I am grateful to J.S. Ashe, M.S. Engel, C.D. Michener, and C.A. Murphy for comments and support throughout this project. I thank K.-D. Klass and an anonymous reviewer for comments that greatly improved an earlier draft of this paper. I am also grateful to the curators of the museums for loan of specimens: M. Brendell, A. Davies, D. Drugmand, T. Erwin, J. Frisch, L. Herman, D. Kavanaugh, O. Martin, J. Navarrete Heredia, A. Newton, P. Perkins, B. Ratcliffe, H. Schillhammer, M. Thayer, M. Uhlig, and L. Zerche. Financial support was provided by an Ernst Mayr Travel Grant from the Museum of Comparative Zoology, Harvard; a Panorama Grant from the Natural History Museum and Biodiversity Research Center of the University of Kansas; numerous H.B. Hungerford Grants from the program of Entomology at the Department of Ecology and Evolutionary Biology of the University of Kansas; the Snow Entomological Development Fund from the Division of Entomology at Natural History Museum/BRC of the University of Kansas; a 10 week graduate fellowship from the Smithsonian Institution; a Collection Study Grant from the American Museum of Natural History; a Sigma-Xi Grant in Aid of Research; and by NSF PEET Grants (DEB-9521755 and DEB-9978110) awarded to J.S. Ashe. I will also like to thank M. Sharkey and his collaborators in Colombia (NSF grant DEB-0205982) for contribution of specimens from Colombia for this project. This is contribution number 3358 from the Snow Entomological Collection, Division of Entomology, Natural History Museum, University of Kansas.

8. References ASHE, J.S. & S. CHATZIMANOLIS 2003. A revision of the genus Elmas Blackwelder, 1952 (Coleoptera: Staphylinidae: Staphylininae: Xanthopygina), with a preliminary reconstructed phylogeny of the species. – Scientific Papers, Natural History Museum, University of Kansas 28: 1–41. BERNHAUER, M. 1905. Neue Staphyliniden aus Südamerika. – Deutsche Entomologische Zeitschrift 1905: 177–187. BERNHAUER, M. 1917a. Neue Staphyliniden aus den columbischen Cordilleren und dem übrigen Südamerika. – Entomologische Blätter 12: 17–24. BERNHAUER, M. 1917b. Neue südamerikanische Staphyliniden. – Wiener Entomologische Zeitung 36: 112–116. BERNHAUER, M. & K. SCHUBERT 1914. Staphylinidae IV. Pp. 289–408 in: S. SCHENKLING (ed.), Coleopterorum Catalogus 5(57). – Junk, Berlin.


BLACKWELDER, M. 1936. Morphology of the coleopterous family Staphylinidae. – Smithsonian Miscellaneous Collections 94: 1–102. BLACKWELDER, M. 1952. The generic names of the beetle family Staphylinidae, with an essay on genotypy. – United States National Museum Bulletin 200: 1–483. CHATZIMANOLIS, S. 2000. An association between Ptecticus nigrifrons (Diptera: Stratiomyidae) and Nordus fungicola (Coleoptera: Staphylinidae). – Journal of the Kansas Entomological Society 73[2001]: 232–234. CHATZIMANOLIS, S. 2003. Natural history and behavior of Nordus fungicola (Coleoptera: Staphylinidae). – Annals of the Entomological Society of America 96: 225–230. CHATZIMANOLIS, S. in press a. The larva of Nordus fungicola (Sharp) (Coleoptera: Staphylinidae) with notes on natural history. – Deutsche Entomologische Zeitschrift. CHATZIMANOLIS, S. in press b. Phylogeny of the neotropical rove beetle Nordus (Coleoptera: Staphylinidae) with a special reference to the evolution of colouration and the secondary sexual characters. – Systematic Entomology. CHATZIMANOLIS, S. & J.S. ASHE 2004. On the identity of Plociopterus variegatus Sharp (Coleoptera: Staphylinidae). – The Coleopterist’s Bulletin 58: 53–55. ERICHSON, W. 1839. Genera et Species Staphylinorum Insectorum Coleopterorum familiae. – F.H. Morin, Berlin, 400 pp. FABRICIUS, J.C. 1801. Systema Eleutheratorum II. – Bibliopolii Academici Novi, Kiliae, 687 pp. FRANK, J.H. 1982. The parasites of Staphylinidae (Coleoptera): a contribution towards an encyclopedia of the Staphylinidae. – University of Florida Agricultural Experiment Station Bulletin 824: 1–118. GEMMINGER, M. & E. VON HAROLD 1868. Catalogus Coleopterum hucusque descriptorum synonymicus et systematicus. – E.H. Gummi, Monachii, 2: 425–752+6. HAMMOND, P.M. 1972. The micro-structure, distribution and possible function of peg-like setae in male Coleoptera. – Entomologica Scandinavica 3: 40–54. HAYASHI, Y. 1997. Studies on the Asian Staphylininae (Coleoptera, Staphylinidae) III. The characteristics of the Xanthopygini. – Elytra 25: 472–492. HERMAN, L.H. 2001. Catalog of the Staphylinidae (Insecta: Coleoptera). 1758 to the end of the second millennium. – Bulletin of the American Museum of Natural History 265: 1–4218. HERMAN, L.H. 2002. Case 3207. Staphylinidae, Latreille, 1804 (Insecta, Coleoptera): proposed conservation of 65 specific names. – Bulletin of Zoological Nomenclature 59: 99–113. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE 1999. International Code of Zoological Nomenclature, 4th edn., adopted by the International Union of Biological Sciences. – International Trust for Zoological Nomenclature, London, xxix + 306 pp. INTERNATIONAL COMMISSION ON ZOOLOGICAL NOMENCLATURE 2003. Opinion 2053 (Case 3207). Staphylinidae Latreille, 1804 (Insecta, Coleoptera): 65 specific names conserved. – The Bulletin of Zoological Nomenclature 60: 237–246. KRAATZ, G. 1857. Naturgeschichte der Insecten Deutschlands. Abt. 1. Coleoptera. Zweiter Band. – Nicolai, Berlin, Lief. 3–4 pp. 377–768, Lief. 5–6 pp. 769–1080. MOORE, I. 1964. A new key to the subfamilies of the Nearctic Staphylinidae and notes on their classification. – The Coleopterist’s Bulletin 18: 83–91. NAOMI, S.I. 1982. Revision of the subtribe Xanthopygina (Coleoptera: Staphylinidae) of Japan. I. – Kontyû 50: 125–133. NAOMI, S.I. 1983a. Revision of the subtribe Xanthopygina (Coleoptera: Staphylinidae) of Japan. II. – Kontyû 51: 47–55. NAOMI, S.I. 1983b. Revision of the subtribe Xanthopygina (Coleoptera: Staphylinidae) of Japan. III. – Kontyû 51: 582–592. NEWTON, A.F., JR. & M.K. THAYER 1995. Protopselaphinae new subfamily for Protopselaphus new genus from Malaysia, with a phylogenetic analysis and review of the omaliine group of Staphylinidae including Pselaphidae (Coleoptera). Pp. 219–320 in: J. PAKALUK & S. ÅLIPINSKI (eds.), Biology, Phylogeny and Classification of Coleoptera. Papers celebrating the 80th birthday of Roy A. Crowson. – Muzeum i Instytut Zoologii PAN., Warszawa.

53


NEWTON, A.F. JR., M.K. THAYER, J.S. ASHE & D.S. CHANDLER 2001. 22. Staphylinidae. Pp. 272–418 in: R.H. ARNETT & M.C. THOMAS (eds.), American Beetles. Archostemata, Myxophaga, Adephaga, Polyphaga: Staphyliniformia, Vol. 1. – CRC Press, Boca Raton. NORDMANN, A. VON 1837. Symbolae ad monographiam staphylinorum. – Ex Academiae Caesareae Scientiarum 4: 1–167. PAYKULL, G. DE 1789. Monographia Staphylinorum Sueciae. – Johann Edmann, Upsaliae, 89 pp. QUEZADA, J.R., C.A. AMAYA, & L.H. HERMAN 1969. Xanthopygus cognatus Sharp (Coleoptera: Staphylinidae), an enemy of the coconut weevil, Rhynchophorus palmarum L. (Coleoptera: Curculionidae) in El Salvador. – Journal of the New York Entomological Society 20: 264–269. SCHEERPELTZ, O. 1971. Studien an den Arten der neotropischen Gattung Brachydirus Nordman [sic] (Col. Staphylinidae, Subfam. Staphylininae, Tribus Xanthopygini). – Mitteilungen der Entomologischen Gesellschaft Basel 21: 93–109. SHARP, D. 1876. Contribution to an insect fauna of the Amazon Valley (Col. Staph.). – Transactions of the Entomological Society of London 1876: 27–424. SHARP, D. 1884. Staphylinidae. Pp. 313–392 in: Biologia Centrali-Americana. Insecta. Coleoptera 1(2). – Taylor & Francis, London. SMETANA, A. 1995. Rove beetles of the subtribe Philonthina of America north of Mexico (Coleoptera: Staphylinidae). Classification, phylogeny and taxonomic revision. – Memoirs on Entomology, International 3: 1–946.

19

SMETANA, A. & A. DAVIES 2000. Reclassification of the north temperate taxa associated with Staphylinus sensu lato, including comments on relevant subtribes of Staphylinini (Coleoptera: Staphylinidae). – American Museum Novitates 3287: 1–88. SOLSKY, S.M. 1872. Enumération et description des coléoptères de la famille Staphylinidae recueillis par MM. C. Jelski et le Baron de Nolken pendant leurs voyages dans l’Amérique du Sud en 1870 et 1871. – Horae Societatis Entomologicae Rossicae 8[1871]: 289–314. SOLSKY, S.M. 1875. Matériaux pour l’entomographie de l’Amérique du Sud. Staphylinides recueillis par MM. C. Jelski et le Baron de Nolken dans le Pérou el la Nouvelle Grenade. Article III. – Horae Societatis Entomologicae Rossicae 11: 3–36. THAXTER, R. 1900. Preliminary diagnoses of new species of Laboulbeniaceae, II. – Proceedings of the American Academy of Arts and Sciences 35: 153–209. THAXTER, R. 1908. Contribution toward a monograph of the Laboulbeniaceae. Part II. – Memoirs of the American Academy of Arts and Sciences 13: 217–469. THAXTER, R. 1931. Contribution toward a monograph of the Laboulbeniaceae. Part V. – Memoirs of the American Academy of Arts and Sciences 16: 1–432. WENDELER, H. 1926. Neue exotische Staphyliniden (Col.). – Deutsche Entomologische Zeitschrift 1926: 71–75. WILEY, E.O. & R.L. MAYDEN 2000. The evolutionary species concept. Pp. 70–89 in: Q.D. WHEELER & R. MEIER (eds.), Species concept and phylogenetic theory: a debate. – Columbia University Press, New York.

20 Figs. 19–21. Distribution of Nordus species. 19: N. antennatus, N. aeneipennis, N. banghaasi, and N. amazonicus. 20: N. championi, N. fungicola, and N. stomachoponos. 21: N. dichromos, N. xanthocerus, N. semelius, N. zischkai, N. seferisi, N. portokalis, N. amphivolos, N. cherylae, N. elytisi, N. elegans, and N. periergos.

21

54


Figs. 22–57. Antennae. 22: N. aeneipennis. 23: N. amazonicus. 24: N. amphivolos. 25: N. antennatus. 26: N. banghaasi. 27: N. batesi. 28: N. championi. 29: N. cherylae. 30: N. cribricollis. 31: N. densiventris. 32: N. dichromos. 33: N. diversiventris. 34: N. fungicola. 35: N. facialis. 36: N. gebieni. 37: N. maculiceps. 38: N. major. 39: N. nigroscutellatus. 40: N. omios. 41: N. simplex. 42: N. solitarius. 43: N. speciosus. 44: N. stomachoponos. 45: N. styloceros. 46: N. terminalis. 47: N. velitaris. 48: N. zischkai. 49: N. elegans. 50: N. elytisi. 51: N. longipes. 52: N. periergos. 53: N. picticornis. 54: N. portokalis. 55: N. seferisi. 56: N. semelius. 57: N. xanthocerus. Scale in Figs. 24, 36, and 48 valid for all figures.


55

Figs. 61–65. Male abdominal sternum VII. 61: N. amazonicus. 62: N. densiventris. 63: N. dichromos. 64: N. omios. 65: N. styloceros. Scale in Fig. 63 valid for all figures.

56


58

66

59

67

60

68

Figs. 58–60, 66–68. Distribution of Nordus species. 58: N. maculiceps, N. densiventris, N. batesi, and N. cribricollis. 59: N. major, N. longipes, N. gebieni, N. nigroscutellatus, and N. aeneiceps. 60: N. diversiventris, N. facialis, N. picticornis, and N. simplex. 66: N. solitarius. 67: N. styloceros, N. speciosus, and N. velitaris. 68: N. omios and N. terminalis.


Figs. 69–86. Male abdominal sternum VIII. 69: N. aeneipennis. 70: N. amazonicus. 71: N. amphivolos. 72: N. antennatus. 73: N. banghaasi. 74: N. batesi. 75: N. championi. 76: N. cherylae. 77: N. cribricollis. 78: N. densiventris. 79: N. dichromos. 80: N. diversiventris. 81: N. facialis. 82: N. fungicola. 83: N. gebieni. 84: N. maculiceps. 85: N. major. 86: N. nigroscutellatus. Scale in Figs. 71 and 83 valid for all figures.

57

58


Figs. 87–103. Male abdominal sternum VIII. 87: N. omios. 88: N. terminalis. 89: N. simplex. 90: N. solitarius. 91: N. styloceros. 92: N. velitaris. 93: N. zischkai. 94: N. elegans. 95: N. elytisi. 96: N. periergos. 97: N. picticornis. 98: N. portokalis. 99: N. seferisi. 100: N. semelius. 101: N. speciosus. 102: N. testaceus. 103: N. xanthocerus. Scale in Fig. 95 valid for all figures.


Figs. 104–112. Male abdominal sternum IX. 104: N. championi. 105: N. densiventris. 106: N. gebieni. 107: N. maculiceps. 108: N. terminalis. 109: N. velitaris. 110: N. zischkai. 111: N. speciosus. 112: N. portokalis. Scale in Fig. 106 valid for all figures except for Fig. 112.

Figs. 113–121. Female abdominal sternum VIII. 113: N. batesi. 114: N. championi. 115: N. diversiventris. 116: N. simplex. 117: N. solitarius. 118: N. speciosus. 119: N. stomachoponos. 120: N. styloceros. 121: N. terminalis. Scale in Fig. 115 valid for all figures.

59

60


Figs. 122–153. Aedeagus; black dots represent sensory spinules and black long lines macrosetae; sensory spinules are included in the dorsal view despite their ventral position; darker coloration of median lobe apically is shown as darker dotted areas. 122–125: N. aeneiceps; 122: dorsal view; 123: lateral view; 124: detail of apex, lateral view; 125: detail of apex, dorsal view. 126–129: N. amazonicus; 126: dorsal view; 127: lateral view; 128: detail of apex, lateral view; 129: detail of apex, dorsal view. 130–133: N. amphivolos; 130: dorsal view; 131: lateral view; 132: detail of apex, lateral view; 133: detail of apex, dorsal view. 134– 137: N. antennatus; 134: dorsal view; 135: lateral view; 136: detail of apex, lateral view; 137: detail of apex, dorsal view. 138–141: N. banghaasi; 138: dorsal view; 139: lateral view; 140: detail of apex, lateral view; 141: detail of apex, dorsal view. 142–145. N. batesi; 142: dorsal view; 143: lateral view; 144: detail of apex, lateral view; 145: detail of apex, dorsal view. 146–149: N. championi; 146: dorsal view; 147: lateral view; 148: detail of apex, lateral view; 149: detail of apex, dorsal view. 150–153: N. cherylae; 150: dorsal view; 151: lateral view; 152: detail of apex, lateral view; 153: detail of apex, dorsal view.


61

Figs. 154–185. Aedeagus; black dots represent sensory spinules and black long lines macrosetae; sensory spinules are included in the dorsal view despite their ventral position; darker coloration of median lobe apically is shown as darker dotted areas. 154–157: N. cribricollis; 154: dorsal view; 155: lateral view; 156: detail of apex, lateral view; 157: detail of apex, dorsal view. 158–161: N. densiventris; 158: dorsal view; 159: lateral view; 160: detail of apex, lateral view; 161: detail of apex, dorsal view. 162–165: N. dichromos; 162: dorsal view; 163: lateral view; 164: detail of apex, lateral view; 165: detail of apex, dorsal view. 166–169: N. diversiventris; 166: dorsal view; 167: lateral view; 168: detail of apex, lateral view; 169: detail of apex, dorsal view. 170–173: N. elegans; 170: dorsal view; 171: lateral view; 172: detail of apex, lateral view; 173: detail of apex, dorsal view. 174–177: N. elytisi; 174: dorsal view; 175: lateral view; 176: detail of apex, lateral view; 177: detail of apex, dorsal view. 178–181: N. facialis; 178: dorsal view; 179: lateral view; 180: detail of apex, lateral view; 181: detail of apex, dorsal view. 182–185: N. fungicola; 182: dorsal view; 183: lateral view; 184: detail of apex, lateral view; 185: detail of apex, dorsal view.

62


Figs. 186–217. Aedeagus; black dots represent sensory spinules and black long lines macrosetae; sensory spinules are included in the dorsal view despite their ventral position; darker coloration of median lobe apically is shown as darker dotted areas. 186–189: N. gebieni; 186: dorsal view; 187: lateral view; 188: detail of apex, lateral view; 189: detail of apex, dorsal view. 190–193. N. maculiceps; 190: dorsal view; 191: lateral view; 192: detail of apex, lateral view; 193: detail of apex, dorsal view. 194–197: N. major; 194: dorsal view; 195: lateral view; 196: detail of apex, lateral view; 197: detail of apex, dorsal view. 198–201: N. nigroscutellatus; 198: dorsal view; 199: lateral view; 200: detail of apex, lateral view; 201: detail of apex, dorsal view. 202–205: N. omios; 202: dorsal view; 203: lateral view; 204: detail of apex, lateral view; 205: detail of apex, dorsal view. 206– 209: N. periergos; 206: dorsal view; 207: lateral view; 208: detail of apex, lateral view; 209: detail of apex, dorsal view. 210–213: N. picticornis; 210: dorsal view; 211: lateral view; 212: detail of apex, lateral view; 213: detail of apex, dorsal view. 214–217: N. portokalis; 214: dorsal view; 215: lateral view; 216: detail of apex, lateral view; 217: detail of apex, dorsal view.


63

Figs. 218–247. Aedeagus; black dots represent sensory spinules and black long lines macrosetae; sensory spinules are included in the dorsal view despite their ventral position; darker coloration of median lobe apically is shown as darker dotted areas. 218–221: N. seferisi; 218: dorsal view; 219: lateral view; 220: detail of apex, lateral view; 221: detail of apex, dorsal view. 222–225: N. semelius; 222: dorsal view; 223: lateral view; 224: detail of apex, lateral view; 225: detail of apex, dorsal view. 226–229: N. simplex; 226: dorsal view; 227: lateral view; 228: detail of apex, lateral view; 229: detail of apex, dorsal view. 230–233: N. solitarius; 230: dorsal view; 231: lateral view; 232: detail of apex, lateral view; 233: detail of apex, dorsal view. 234–236: N. speciosus; 234: dorsal view; 235: lateral view; 236: detail of apex, dorsal view. 237–240: N. styloceros; 237: dorsal view; 238: lateral view; 239: detail of apex, lateral view; 240: detail of apex, dorsal view. 241– 244: N. terminalis; 241: dorsal view; 242: lateral view; 243: detail of apex, lateral view; 244: detail of apex, dorsal view. 245–247: N. testaceus; 245: dorsal view; 246: lateral view; 247: detail of apex, lateral view.

64


Figs. 248–255. Aedeagus; black dots represent sensory spinules and black long lines macrosetae; sensory spinules are included in the dorsal view despite their ventral position; darker coloration of median lobe apically is shown as darker dotted areas. 248–251: N. velitaris; 248: dorsal view; 249: lateral view; 250: detail of apex, lateral view; 251: detail of apex, dorsal view. 252–255: N. zischkai; 252: dorsal view; 253: lateral view; 254: detail of apex, lateral view; 255: detail of apex, dorsal view.