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smithsonian contributions to zoology • number 641

Review of the Families 

Tanypezidae and 

Strongylophthalmyiidae, 

with a Revision of 

Neotanypeza Hendel 

(Diptera: Schizophora)

Owen Lonsdale
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smithsonian contributions to zoology • numb e r 6 4 1

Review of the Families 

Tanypezidae and 

Strongylophthalmyiidae, 

with a Revision of 

Neotanypeza Hendel 

(Diptera: Schizophora)

Owen Lonsdale

washington d.c.

2013
ABSTRACT

Lonsdale, Owen. Review of the families Tanypezidae and Strongylophthalmyiidae, with a revision of Neotanypeza Hendel 

(Diptera: Schizophora). Smithsonian Contributions to Zoology, number 641, vi + 60 pages, 92 figures, 5 tables, 2013. The 

families Tanypezidae and Strongylophthalmyiidae are reviewed, including Neotanypeza Hendel and Tanypeza Fallén in 

the former and Nartshukia Shatalkin and Strongylophthalmyia Heller in the latter. Neotanypeza is revised, and Tanypezidae is analyzed phylogenetically. Scipopeza Enderlein, syn. nov., Polphopeza Enderlein (syn. Hennig (1936)), Tritanypeza

Enderlein, syn. nov., and Tripolphopeza Enderlein, syn. nov., are treated as junior synonyms of Neotanypeza. The species 

Scipopeza dimorpha Hennig and S. grandis Enderlein are subsequently treated as Neotanypeza for the first time. Twentyfive extant species of Neotanypeza are recognized, 10 of which are described here as new: N. alopecia, sp. nov., N. argentia,

sp. nov., N. leucothrix, sp. nov., N. marshalli, sp. nov., N. micans, sp. nov., N. nigrithrix, sp. nov., N. plotoplax, sp. nov., 

N. posthos, sp. nov., N. symmetros, sp. nov., and N. vexilla, sp. nov. Neotanypeza claripennis (Schiner), stat. reinst., and 

N. abdominalis (Wiedemann), stat. reinst., are resurrected from synonymy. Neotanypeza flavitibia Hennig, syn. nov., is 

included as a junior synonym of N. elegans (Wiedemann); Tritanypeza rufiventris Enderlein, syn. nov., as a synonym of 

N. flavibasis (Enderlein); Polphopeza elegantina Enderlein, syn. nov., as a synonym of N. apicalis (Wiedemann); Tanypeza 

flavohirta Enderlein, syn. nov., as a synonym of N. ornatipes (Bigot); and Tritanypeza cubitofusca Enderlein, syn. nov., and 

N. nigripalpis Hennig, syn. nov., as synonyms of N. callitarsis (Rondani). Twelve lectotypes are designated for the following species (original combinations): Tanypeza abdominalis Wiedemann, Tanypeza apicalis Wiedemann, Tritanypeza cubitofusca Enderlein, Tanypeza claripennis Schiner, Tanypeza dallasi Shannon, Tanypeza elegans Wiedemann, Tritanypeza 

flavibasis Enderlein, Tritanypeza rufiventris Enderlein, Polphopeza montana Enderlein, Tritanypeza ochrifemur Enderlein,

Taeniaptera pallidipennis Bigot, and Tanypeza rutila Wulp.

Cover images (left to right): Strongylophthalmyia angustipennis Melander, male and female in copula; Neotanypeza 

claripennis, tentatively included male; Tanypeza longimana Fallén, female.

Published by SMITHSONIAN INSTITUTION SCHOLARLY PRESS

P.O. Box 37012, MRC 957, Washington, D.C. 20013-7012

www.scholarlypress.si.edu

This work, authored by Owen Lonsdale of the Department of Agriculture and Agri-Food, Government of Canada, was 

prepared as a result of a postdoctoral fellowship awarded by the Smithsonian Institution’s National Museum of Natural 

History. 

© 2013 by Her Majesty the Queen in Right of Canada, as represented by the Minister of Agriculture and Agri-Food 

Canada

The rights to images used with permission as well as the cover and interior designs of the publication are owned either by 

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Library of Congress Cataloging-in-Publication Data

Lonsdale, Owen.

Review of the families Tanypezidae and Strongylophthalmyiidae, with a revision of Neotanypeza Hendel (Diptera: 

Schizophora) / Owen Lonsdale.

 p. cm. — (Smithsonian contributions to zoology ; no. 641)

Includes bibliographical references.

1. Tanypezidae. 2. Strongylophthalmyiidae. I. Title. II. Series: Smithsonian contributions to zoology ; no. 641. 

QL537.T357L66 2013

595.77—dc23

2013020155

ISSN: 0081-0282 (print); 1943-6696 (online)

The paper used in this publication meets the minimum requirements of the American National Standard for 

Permanence of Paper for Printed Library Materials Z39.48–1992.
Contents

List of Tables v

INTRODUCTION 1

Tanypezidae 2

Strongylophthalmyiidae 2

Taxonomic Treatments 2

Life History 2

MATERIALS AND METHODS 3

MORPHOLOGY OF TANYPEZIDAE AND

STRONGYLOPHTHALMYIIDAE 6

Diagnosis 6

Description 6

RESULTS AND DISCUSSION 8

Family Level Support 8

Status of Enderlein’s Genera 10

Tanypeza 10

Neotanypeza 10

Key to the World Genera of Tanypezidae and 

 Strongylophthalmyiidae 11

SYSTEMATICS 11

Neotanypeza Hendel, 1903 11

 Diagnosis 11

 Description 12

Key to the Extant Species of Neotanypeza 13

Descriptions of Extant Neotanypeza Species 15

 Neotanypeza abdominalis (Wiedemann), stat. reinst. 15

 Neotanypeza alopecia, sp. nov. 16

 Neotanypeza apicalis (Wiedemann) 17

 Neotanypeza argentia, sp. nov. 18

 Neotanypeza callitarsis (Rondani) 19

 Neotanypeza claripennis (Schiner), stat. reinst. 20
i v • smithsonian contributions to zoology

 Neotanypeza dallasi (Shannon) 21

 Neotanypeza dimorpha (Hennig), comb. nov. 22

 Neotanypeza elegans (Wiedemann) 23

 Neotanypeza flavibasis (Enderlein) 24

 Neotanypeza flavicalx (Enderlein) 25

 Neotanypeza grandis (Enderlein), comb. nov. 25

 Neotanypeza leucothrix, sp. nov. 26

 Neotanypeza marshalli, sp. nov. 26

 Neotanypeza micans, sp. nov. 27

 Neotanypeza montana (Enderlein) 27

 Neotanypeza nigrithrix, sp. nov. 29

 Neotanypeza ochrifemur (Enderlein) 29

 Neotanypeza ornatipes (Bigot) 30

 Neotanypeza plotoplax, sp. nov. 31

 Neotanypeza posthos, sp. nov. 31

 Neotanypeza quadrisetosa (Enderlein) 32

 Neotanypeza rutila (Wulp) 33

 Neotanypeza symmetros, sp. nov. 34

 Neotanypeza vexilla, sp. nov. 34

Acknowledgments 35

REFERENCES 37

FIGURES 39
Tables

1. Morphological character list 4

2. Character matrix 7

3. Synapomorphies of Tanypezidae and Strongylophthalmyiidae 9

4. Synapomorphies of Strongylophthalmyiidae 9

5. Synapomorphies of the Tanypezidae 10

INTRODUCTION

The Tanypezidae, or stretched-foot flies of von Tschirnhaus (2008), is a readily recognizable family of acalyptrate Diptera (Schizophora). The position of this family in 

the Schizophora has been and still is somewhat uncertain, but there is consensus that 

the Tanypezidae and Strongylophthalmyiidae are sister groups, leading some authors 

(Griffiths, 1972; McAlpine, 1997b) to justifiably include the latter as a subfamily. Although both share a number of external synapomorphies and have highly similar male 

and female genitalia, the general habitus of each is distinctive and conservative, leading 

most authors to keep them separate. Although there has arguably been an unnecessary 

proliferation of family group names for several small clusters of genera in an already 

family-rich Schizophora, the separation of these two families is maintained here, mostly 

for the practical purpose of diagnostics, as discussed below.

The species of Tanypezidae have silvery stripes on the body, a relatively robust thorax, long slender legs, and a large head that is flattened posteriorly (Figures 2–6). Additional features include an apical convergence of veins R4+5 and M1 (Figures 25–33), dorsal 

setulae on vein R1

, and a variably tomentose “ocellar disc” behind the ocelli (Figure 7). 

The species of Strongylophthalmyiidae are comparatively more slender in appearance, 

have reduced chaetotaxy on the frons and thorax, and have a globose sepsid-like head 

with pronounced ocellar bristles and a deep depression behind the ocelli (Figures 1, 34). 

Within the Acalyptratae (Brachycera: Schizophora), McAlpine (1989) relegated the 

sister families Tanypezidae and Strongylophthalmyiidae to a basal position in the Diopsoidea (=Nothyboidea of Hennig, 1958). The remaining Diopsoidea, including Somatiidae, 

Psilidae, Nothybidae, Megamerinidae, Syringogastridae, and Diopsidae, were defined by 

a bare vein R1

 (here treated as plesiomorphic in all diopsoid families), loss of the anterior 

notopleural bristle, and loss of the presutural intra-alar bristle. Conversely, McAlpine 

(1997b) found tanypezids to be more similar in morphology to certain taxa in the Nerioidea, particularly those in the Cypselosomatidae, and suggested a possible relationship 

with that superfamily. In an earlier paper (McAlpine, 1997a), he made similar statements 

regarding the family Megamerinidae, which he moved with more confidence from the 

Diopsoidea to the Nerioidea, following his earlier classification in McAlpine (1966). The 

classification of McAlpine (1989) is maintained here, however, and will be thoroughly 

discussed in a future publication (Lonsdale et al., unpublished). Other parallels with the 

Review of the Families Tanypezidae and 

Strongylophthalmyiidae, with a Revision of 

Neotanypeza Hendel (Diptera: Schizophora)

Owen Lonsdale, Agriculture and Agri-Food 

Canada, 960 Carling Avenue, Ottawa, Ontario K1A 0C6, Canada. Correspondence: owen

.lonsdale@agr.gc.ca. 

Manuscript received 26 September 2012; accepted 6 February 2013.
2 • smithsonian contributions to zoology

neroid families Neriidae and Micropezidae have been frequent 

in the literature since long legs and convergent R4+5 and M1

 veins 

occur in all three families. An alternative to these classifications 

was proposed by Griffiths (1972), which was discussed by McAlpine (1985) and is not considered further here.

Tanypezidae

Tanypeza Fallén was described for the Holarctic T. longimana Fallén (Figure 2) by Fallén (1820). A number of Neotanypeza Hendel and one micropezid (Taeniaptera melanopus

Bigot) have been placed in it since, but the genus is here considered to contain only the type species and the Nearctic T. picticornis Knab and Shannon. 

Hendel (1903) proposed Neotanypeza as a subgenus in 

Tanypeza for the South American species with reduced ocellar 

and postvertical bristles (as in Figures 7, 8). Enderlein (1913) 

then described Polphopeza Enderlein and Scipopeza Enderlein 

for species with differing numbers of frontal orbital bristles. 

Enderlein (1913) also treated Myrmecomyia Robineau-Desvoidy 

(=Cephalia Meigen, Ulidiidae) and Tetradiscus Bigot (=Chyliza Fallén, Psilidae) as tanypezids, but these were removed by 

Hennig (1936), who also synonymized Polphopeza with Neotanypeza, which was given full generic status. Enderlein (1936) 

immediately resurrected Polphopeza and again added two more 

new genera, Tripolphopeza Enderlein and Tritanypeza Enderlein, both of which have three pairs of dorsocentral bristles. 

In the present revision it has been found that the phylogeny 

of the Tanypezide is best represented by a two-genus classification, including Tanypeza and Neotanypeza, with all of Enderlein’s genera reduced to synonyms of the latter. A similar system 

is presented in Apigian and Thompson (2010), although Enderlein’s genera Polphopeza, Tritanypeza, and Tripolphopeza are 

recognized as subgenera, albeit only tentatively. The bulk of the 

family’s diversity is in the neotropical Neotanypeza, which has 

25 species. This genus is here keyed, illustrated, and analyzed 

phylogenetically, including all Tanypeza, Nartshukia Shatalkin, 

several Strongylophthalmyia Heller, and a number of neroid and 

diopsoid exemplars as out-groups. 

The species of Neotanypeza vary subtly in morphology 

and can be difficult to diagnose and delimit; an example of the 

difficulties involved in species circumscription is detailed in 

the comments section for N. claripennis. The numerous other 

species-level problems are discussed in the comments sections for 

those relevant species. As a result of these difficulties, the present 

estimate of species number has remained conservative, despite 

describing a number of new taxa. It must therefore be stressed 

that the species limits presented here must be considered tentative and should be revisited at a later date when additional data 

more sensitive to species boundaries are available. 

Strongylophthalmyiidae

The Strongylophthalmyiidae is also represented by two genera: Nartshukia (known from a single Vietnamese female; Figure 

34) and the widespread and diverse Strongylophthalmyia. Contrasting the predominantly New World Tanypezidae, this group 

has a mostly East Asian distribution (see Shatalkin, 1996), although two species (S. angustipennis Melander [Figure 1] and 

S. pengellyi Barber) are restricted to the Nearctic and several 

species extend into, or are restricted to, the Australian region. 

The only two species described from the Afrotropics were transferred to the Clusiidae by Barraclough (2000). Shatalkin (1996) 

included the genus Longinasus Frey in the family, but Steyskal 

(1968) found this genus to be a junior synonym of Ozaenina

Enderlein, a genus in the tephritoid family Richardiidae. 

Taxonomic Treatments

The two species of Tanypeza were keyed by Steyskal (1965), 

and the two Nearctic Strongylophthalmyia were keyed by Barber 

(2006). The Strongylophthalmyiidae of the Old World, of which 

there are 44 described species, have most recently been treated 

by Steyskal (1971), Krivosheina (1981), Shatalkin (1981, 1993, 

1996), Iwasa (1992, 1998), Yang and Wang (1992, 1996), and 

Papp et al. (2006). The neotropical Tanypezidae was last treated 

in 1936 by Hennig, followed by Enderlein, and the first fossil 

representative of the family, Neotanypeza dominicana Lonsdale 

and Apigian from Dominican amber dating to 17–20 mya, was 

described by Lonsdale and Apigian (2010). An excellent overview of the Tanypezidae was provided by Roháchek (1998). 

Life History

Adults. Very little is known about the life history of 

adult Tanypezidae and Strongylophthalmyiidae, although thorough records of the biology of the two Nearctic Strongylophthalmyia species were provided by Barber (2006), who detailed 

ecology, biology, and behavior. Tanypeza longimana adults have 

been found in low vegetation in humid deciduous woods in Europe, usually at low altitudes in floodplains and submontane 

areas and often around running water (Roháchek, 1998). Unpublished records report the collection of Strongylophthalmyia from 

Alocasia leaves in Australia and ant-like behavior on banana 

leaves in a New Guinea species (Evenhuis, 1989). 

Larvae and Puparia. Palaearctic Strongylophthalmyia species have been reared from under the bark of rotting 

birch, elm (S. stackelbergi Krivosheina), and aspen (S. pictipes 

Frey, S. ustulata (Wahlberg)); larvae were common in the lower 

layers of the bark, whereas puparia were found in the outer layers (Krivosheina, 1984). Nearctic S. angustipennis were also 

reared from aspen bark, where it was presumed they overwintered as mature or nearly mature larvae; the puparia are characterized by biramous anterior spiracles similar to those found in 

S. ustulata (Barber, 2006). Tanypeza longimana was also speculated to live in rotting wood as larvae (Chandler, 1975). There 

is only one documented case of an unsuccessful attempt to rear 

the young of T. longimana, which were thoroughly described in 

the same paper (Foote, 1970): presumably saprophagous third 

instar maggots derived from a gravid female swept from yellow 
number 641 • 3

skunk cabbage (Lysichitum americanum) were fed decaying watermelon rind and pulp.

Eggs. Eggs are approximately 2.3 times longer than 

wide in the Tanypezidae, known from Neotanypeza elegans and 

Tanypeza longimana (Figures 87, 88), and approximately four 

times longer than wide in Strongylophthalmyia angustipennis 

(Figure 89). The micropyle is minute and not strongly raised 

from the egg surface. The surface is covered with minute shallow 

tubercules in Strongylophthalmyia that are grouped within elongate hexagonal cells. Eggs in the Tanypezidae are more roughly 

tuberculate, without a pattern of cells and with several longitudinal furrows converging apically. 

MATERIALS AND METHODS

To discover synapomorphies for the Tanypezidae and

Strongylophthalmyiidae and to polarize characters, out-group 

representatives at the National Museum of Natural History of 

Diopsoidea and Nerioidea were examined, the two superfamilies 

to which the family has been allied. Most of these taxa were 

included in the phylogenetic analysis (see Table 1): Megamerina 

annulifera Bigot (Megamerinidae), Sphyracephala subbifasciata

Fitch (Diopsidae), Centrioncus prodiopsis Speiser (Diopsidae), 

Loxocera collaris Loew (Psilidae), Somatia aestiva (Fabricius) 

(Somatiidae), Nothybus triguttatus Bezzi (Nothybidae), Gobrya 

sp. (Gobryidae), Syringogaster rufa Cresson (Syringogastridae), 

Formicosepsis tinctipennis Meijere (Cypselosomatidae), and 

Pseudopomyza antipoda (Harrison) (Pseudopomyzidae). The 

single male Centrionicus available was not dissected; male genitalic data were taken from Feijen (1983). Data for Nartshukia, 

known only from the female holotype of N. musiva Shatalkin 

(Zoological Museum, Moscow), were taken from Shatalkin 

(1994) and photographs provided by A. Shatalkin.

Most of the terminology used by Steyskal (1987), Roháchek 

(1998), and Barber (2006) is maintained for structures of the 

male genitalia. Deviations from these norms, illustrated in Figures 35–46, are as follows: the “paramere” of Steyskal (1987), 

the “arch-like sclerite of the hypandrium” of Barber (2006), and 

the pregonite of Roháchek (1998) are here treated as the postgonite. The “postgonite” of Barber (2006) is treated as a process 

of the hypandrium. Barber’s (2006) “basiphallus” is possibly an 

elongate basal section of the distiphallus, with the basiphallus 

proper comprising a basal sclerotized ring in this structure, but 

the homology of this region is still partially unclear. The male 

pregenitalic segments are treated as sternites 6, 7, and 8 only. 

The supra-alar bristle of McAlpine (1989) is treated as the intraalar. “Fronto-orbital bristle” is used preferentially over “orbital 

bristle,” as is “phallapodeme” over “aedaegal apodeme.” Abdominal sclerites referred to in the text will belong to the male 

unless stated otherwise. The male cerci are fused and are always 

referred to in the plural.

Male and female genitalia were prepared by macerating the 

abdomen in hot potassium hydroxide (10% solution) for 10–13 

minutes, dissolving the soft tissue, followed by washing in glacial 

acetic acid and water. Genitalia were dissected and then illustrated using a camera lucida while in glycerol. Abdomens were 

placed in genitalia vials pinned with the specimens following examination. Illustrations of the male abdomen, detailing components of the genitalia, are provided for one species of Tanypeza

(T. longimana; Figures 41–45) and Strongylophthalmyia (S. angusticollis Frey; Figures 35–40); the simpler female terminalia 

of these genera are not redrawn as excellent illustrations are already provided in Roháchek (1998) and Barber (2006), with the 

exception of a Tanypeza ventral receptacle (Figure 51), which is 

added for comparison to that of Neotanypeza (Figure 50). Eggs 

and wings were permanently mounted on slides in euparal. 

Phylogenetic hypotheses for the species of Tanypezidae were 

produced through analysis of the character matrix in Table 2 

using PAUP* version 4.0b10 (Phylogenetic Analysis Using Parsimony; Swofford, 2003); a heuristic analysis was performed using 

100 replicate random addition sequences with tree bisection and 

reconnection (TBR). The character matrix was compiled using 

DELTA (Dallwitz et al., 2000). All characters were treated as unordered and unweighted. Trees were drawn, and characters were 

optimized (ACCTRAN optimization) with the aid of MacClade 

(Maddison and Maddison, 2005) and Winclada (Nixon, 2002). 

Because N. grandis exhibits sexual dimorphism in chaetotaxy, 

external characters coded for in the matrix (Table 1) were taken 

solely from the male. Character numbers are listed in square 

brackets in the following phylogenetic discussion.

Material was borrowed from or deposited in the following 

institutions: 

AMNH American Museum of Natural History, New York 

BMNH Natural History Museum, London 

CASC California Academy of Sciences, San Francisco 

CBFC Colección Boliviana de Fauna, La Paz

CNC Canadian National Collection of Insects, Arachnids and Nematodes, Ottawa, Ontario

CSCA California State Collection of Arthropods, California Department of Food and Agriculture, 

Sacramento 

DEBU University of Guelph Insect Collection, Guelph, 

Ontario, Canada 

EMUS Entomological Museum of Utah State University, 

Logan 

INBio Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica 

INPA Colecão Sistemática da Entomologia, Manaus, 

Brazil 

MRSN Museo Regionale de Scienze Naturali, Turin, Italy 

NMW Naturhistorische Museum, Vienna 

PAN Museum and Institute of Zoology, Polish Academy of Sciences, Warsaw

QCAZ Universidad Católica del Ecuador, Quito 

SMT Staatliches Museum für Tierkunde, Dresden, 

Germany 

TAMU Texas A&M University, College Station 

TAU Tel Aviv University, Tel Aviv, Israel 
4 • smithsonian contributions to zoology

TABLE 1. Morphological character list for use in the phylogenetic analysis.

Character Character states (treated as unordered) Character Character states (treated as unordered)

1. Head Back of head flat to shallowly rounded

 Orbicular, back of head rounded, with deep 

depression behind ocelli

2. Frons, posterior Grading onto back of head

margin Flat, strongly delimited

3. Frons Shiny or with thin pilosity

 Velvety, at least in part

4. Male frons As broad as female frons

 Strongly narrowed medially, more so than 

female

5. Reflective Absent

anterolateral Reaching midpoint of frons or beyond

triangles on frons Short, not much longer than wide

6. Ocellar tubercle On posterior margin of frons (closer to level 

of inner vertical) 

 Near midpoint of frons (closer to level of 

anterior fronto-orbital)

7. Postocellar region Shining

 With tomentose patch, sometimes enclosed 

in disc

8. Ocellar disc Absent

 Present, fully tomentose (sometimes shining 

immediately behind ocelli) 

 Present, pilosity reduced to medial spot

9. Occiput Pile short

 Pile long, yellow

 Pile long, white

10. First flagellomere Variable, shorter than three times length of 

pedicel

 Narrow, longer than three times length of 

pedicel

 Thick, width at least half length, longer than 

three times length of pedicel

 Thick, infuscated apically (at least), longer 

than three times length of pedicel

11. Clypeus Narrow, band-like or not as below

 Thick, at least infuscated apically

12. Palpus Brown

 Yellow

13. Buccal cavity Narrow

 Wide

14. Prosternum Narrow, small

 Wide, broadly separating coxae

 Wide, separating coxae and fused to pleuron

15. Hind tibia Straight or evenly bent

 Deviated subbasally

16. Fore tarsal claws Similar to claws on mid and hind legs

 Broad with surrounding region strongly 

setose

17. Fronto-orbital Three or more

number Two

 One 

 None

18. Outer vertical Present

bristle Absent

19. Ocellar bristle Small to absent

 Long, many times length of ocellar tubercle

20. Postvertical bristle Well developed, nearly as long as vertical 

bristle

 Minute to absent

 Of moderate length and longer than tubercle

21. Postvertical bristle Vertical to reclinate or absent

 Slightly to distinctly anteroclinate

22. Postpronotal bristle Present

 Absent

23. Presutural intra-alar Present

 Absent

24. Supra-alar bristle Present (i.e., three bristles in posterolateral 

corner of scutum) 

 Absent (i.e., two bristles in posterolateral 

corner of scutum)

25. Dorsocentral One or none

number Two

 Three or more

26. Lateral scutellar One 

bristle number None

27. Proepisternal Present

bristle Absent

28. Notal setulae Very short, usually brown

 Long, yellow

29. Thorax Without reflective tomentose patches

 With broad silvery tomentose stripes

30. Meron Entirely shining

 Shining with posterior margin tomentose

 Entirely tomentose

31. Long pile Absent (some evident on anepisternum and 

katepisternum in Megamerina) 

 Present on postpronotum, anepisternum, 

anepimeron and katepisternum

32. Hind Narrow at base

basotarsomere With swollen protuberance at base

33. Male hind With pale indistinct setulae

trochanter and With dark, long, stout bristles

femur

34. Greater ampulla Absent

 Present

35. Dark bristles Absent or hairs weak and white

extending to base Present, well developed

of knob on halter
number 641 • 5

TABLE 1. (Continued)

Character Character states (treated as unordered) Character Character states (treated as unordered)

36. Wing Clear

 Dusky, at least apically

 Variably patterned with distinct spots

37. Subcostal break Absent

 Present

38. Subcosta Complete

 Incomplete

39. Vein R1 Bare dorsally

 Setose dorsally

40. Veins R4+5 and M1 Parallel to slightly convergent 

 Strongly convergent

41. CuA2 Shallowly bent or straight

 Strongly bowed

42. Cell dm Posterior margin straight or nearly so

 Posterior margin slightly pointed at dm-cu

 Posterior margin strongly pointed at dm-cu 

(see arrow, Figure 26)

43. Calypter (upper) Linear, nearly absent; sometimes shallowly 

lobate

 Large, broadly rounded

44. Pile on calypter Short, color variable 

(upper) Long, pale

 Long, brown

45. Calypter (lower) Hairs pubescent, relatively sparse

 Hairs short, dense, and “furry”

46. Scutum Entirely dark or with pale shoulder or 

posterolateral spot

 With thick yellowish medial stripe

47. Fore femur Entirely yellow

 With inner distal spot; sometimes also with 

outer distal spot

 Brown apically or predominantly brown

48. Mid femur Yellow

 Brown apically

 Brown on distal third or more

49. Meron, pilose Absent

anteromedial Present

margin

50. Postcoxal bridge Absent

 Present

51. Hind femur Yellow

 Brown medially

 Brown apically

52. Fore tibia Yellow

 Brown on basal half or more

53. Mid tibia Yellow or only brown apically

 Entirely brown

 With basal and apical bands

 Brown on basal half

54. Hind tibia Yellow

 Entirely brown

 With basal and subapical bands

 Brown on basal half

55. Fore basotarsomere Base yellow

 Base brown

56. Mid basotarsomere Base yellow

 Base brown

57. Hind basotarsomere Base yellow

 Base brown

58. Abdomen Entirely brown or only bicolored along lateral 

margins

 Yellow on tergite 2 at least

59. Abdomen Setulose, sometimes smooth, never “pebbled” 

 Without setulae, with “pebbled” texture 

(pronounced posteriorly)

60. Sternite 2 Anterior margin straight, entire

 Anterior margin with short separate sclerite 

with two posterior emarginations

61. Tergite 2 Anterolateral bristles neither long nor thick

 Anterolateral bristles long and thick

62. Male tergite 1 Lateral bristles brown to black

 Lateral bristles yellow

63. Male sternite 6 Symmetrical or nearly so

 Asymmetrical

64. Male sternite 6 Both sides evenly sclerotized

 With heavily sclerotized left sublateral band

65. Male sternite 7 Well developed, distinct

 Atrophied, indistinct

66. Male sternites Encircling abdomen on vertical axis; short

7 and 8 Encircling abdomen obliquely; elongate

67. Male sternite 8 Smooth or pebbled texture

 Wrinkled

68. Male sternite 8 Setulose

 Silvery tomentose

69. Surstylus Separate from epandrium

 Fused to epandrium on posterior half

 Entirely fused to epandrium

70. Surstylus Narrower and shorter than epandrium

 Large and broadly rounded, as large as epandrium (seen laterally)

71. Surstylus, setulae Narrow, curved, not different from 

on outer posterior surrounding setulae

margin Stout, straight, spine-like

72. Bifid hypandrial Absent

process Present

73. Hypandrium Aedeagal guides (hypandrial lobes) separate 

anteromedially

 Aedeagal guides nearly meeting medially

(continued)
6 • smithsonian contributions to zoology

UMO Oxford University Museum of Natural History, 

Oxford, UK 

USNM National Museum of Natural History, Washington, D.C. 

UZMC Zoological Museum, University of Copenhagen, 

Copenhagen 

ZMHU Museum für Naturkunde der Humboldt, Berlin 

ZMUM Zoological Museum of Moscow University, 

Moscow

MORPHOLOGY OF TANYPEZIDAE 

AND STRONGYLOPHTHALMYIIDAE

Diagnosis

Gena with long straight hairs; occiput bulging and with 

long, dense yellow to white pile. Antenna slightly to strongly elbowed. Zero to two (0–2) fronto-orbital bristles. Vibrissa absent. 

Proepisternal and katepisternal bristles absent. Ocellar tubercle 

usually removed from posterior margin of frons. Prosternum 

connected to pleuron laterally (completely fused to pleuron in 

Strongylophthalmyiidae), not separated from pleuron by broad 

membranous region (compare Figure 11 to Figures 9 and 10). 

Calypter broad and lobe-like, with marginal hairs long. Legs 

long and slender. Surstylus completely fused to epandrium (Figures 35, 43, 52). Postgonite long, flexible, band-like, with several 

apical setulae, and fused to inner face of hypandrium basally 

(Figures 39, 44, 56). Female segments 6–10 distinctly narrowed 

and elongate, forming slender tubular ovipositor; female cerci 

long, curved in cross section, and largely fused (Figures 47–49); 

sternite 10 with elongate, well-sclerotized, narrow clavate internal process extending into segment 8 (Figures 47, 48).

Description

Head. Predominantly black, although antenna, face, 

gena, and occiput frequently paler. Antenna at least slightly 

TABLE 1. (Continued)

Character Character states (treated as unordered) Character Character states (treated as unordered)

74. Number of Many

hypandrial bristles One

 None

75. Postgonite Lobate, usually separate, but sometimes fused 

to hypandrium along margin

 Elongate, band-like, curved, and fused to 

hypandrium

76. Basiphallus Distinct, articulating phallapodeme and 

distiphallus

 Indistinct, fused to base of distiphallus

77. Epiphallus Small, lobate (if present) 

 Large, plate-like, composed of two 

articulating sclerites; distal sclerite small

 Large, plate-like, composed of two 

articulating sclerites; distal sclerite large

78. Distiphallus Variable, but not as below

 Elongate, with heavy sclerite in membranous 

apex

 Elongate with apical sclerotized section

 Membranous medially or sublaterally, sides 

toothed; usually not much longer than wide

79. Female tergite 6 Widest basally or apically

 Widest medially

80. Female segment 7 Sternite and tergite separate

 Sternite and tergite partially fused

 Sternite and tergite forming complete 

sclerotized tube

81. Female sternite 8 Entire

 Divided or at least desclerotized medially

82. Female Short

intersegmental At least two times longer than segment 8

membrane

(segments 7 and 8)

83. Sternite and No more than two times longer than wide

tergite 10 Several times longer than wide

84. Female sternite Absent

10 with internal Present

process

85. Female cerci Subcylindrical, round in cross section

 Curved in cross section

86. Spermatheca Three

number Two, well developed

 Two, with one highly reduced

87. Spermatheca Surface smooth

 Surface papillose

88. Spermatheca Base not telescoped

 Base telescoped

89. Female genital Entirely membranous

chamber With vaginal sclerite

90. Ventral receptacle Various, but not as below

 Transversely lobate with surrounding dome

91. Spermathecal ducts Entirely separate 

 Fused near base
number 641

• 

7

TABLE 2. Character matrix.Taxon Character states (see Table 1)Megamerina annulifera 0010011020000000200101110111000001000000020100012011001000000110000000000000000201000100000Sphyracephala subbifasciata 000001000000000020010111011000000002000000000001202111000000000010000000020?00001000000100?Centrioncus prodiopsis 00000010000000002101011101100000000000000200000120210300000000???1?000000000000110000000000Formicosepsis tinctipennis 000000000000000100000000211000000001000000000000222111000000000000000000021000?01??101???0?Loxocera collaris 0000010001010000301101110010000001001100010200000000000000000110?000???0000?0000?0000???00?Nothybus triguttatus 001001100101010020010111001000010002000000000000000111011100000010001000000000?00??001???0?Nartshukia musiva 100?00?0100001001001?10?01?10000?0?011010011?0??2220000000?????????????????????????????????Strongylophthalmyia 1000010010010200101011110111000000001100001100000000000000100110010020010111111211111201111

angustipennis

S. pengellyi 10000100100002001000111?01110000000111000011000022211111101000100100200101111112111?1201111S. spinipalpa 10000100100002001010111101110000000011000012000000200000001000?0010020011111111?11??1??????Tanypeza longimana 011111112(02)10110111000010001011101110001111111111(012)0(01)0020000011111000121001211221010011110010T. picticornis 0111111120101101110000100010111011100011111111110010020000011011000121001211221010??1??????Neotanypeza dominicana 01111111211011011100000020101211011000111211?0?120101111100?11???0?1201??????1?????????????N. argentia 011111112????111110100101010121101??001?1?1110??0?00?00?000?11???0?120???????3?????????????N. elegans 011121122110111111010000101012110110001112121011(12)000(12)(12)(01)(01)(01)0011011100120100111231010011110010N. micans 01112112211111112101001010101211011100111211101?2120220001011011?001201???1123101???1??????N. apicalis 011111112?11111121010010101012110111001112111011000003000001111000012010021123?01???1??????N. nigrithrix 01112112231011113101001010101211011000111212101121102211100?10???????01??????3?????????????N. leucothrix 01112111211011112101001000101211011100111211101121202201110?10???????01??????3?????????????N. quadrisetosa 011111112310111121010010101012110110001112121011212111111001101110012000011123?????????????N. posthos 0111(12)111231011112101011000101211011(01)001112111011112(01)2111100110???0112000011123101???1??????N. symmetros 011111112311111021010110001012110111001112111011122011011001100000012000011123?????????????N. marshalli 011121112310111111010010101012110111001112111011222111111001110000012000001123101???1??????N. dimorpha 011111112310111131010110001012110112001112111011220111110001101100112000011123101???1??????N. vexilla 0111111123101111310101100010121101120011121110111(01)(02)(01)1101100111110001201021123101???1??????N. plotoplax 0111211123101111310101100010121101110011121210?1222111111001100010112010011123?????????????N. grandis 011121112310111131010110001012110112001112111011222111111001101100112010011123101???1??????N. dallasi 01111111231011112102000020101211011000111211101110002201100111111001201??????3101???1??????N. claripennis 011111112310111111020000201012110110001112111011100022011001111110012010011123101??????????N. montana 0111(12)111231011102101001010101211011(01)001112111011(12)00022000001111110012010011123101???1??????N. callitarsis 0111(12)1112(13)1011111102000020101211011(01)001112111011(01)0000(23)0000011111?001201???1123101???1??????N. ornatipes 0111111122111111110200002010121101100011121110?1000022011001111110012010011123101???1??????N. ochrifemur 011111112111111111020000201012110110001112111011100003000001111110012010011123?????????????N. alopecia 0111211223101111110200002010121101110011121(12)10111000(03)3000001101110012010011123101???1??????N. flavicalx 011?2111221111111102000020101211?1100011121110111000330000011?????????????????101???1??????N. abdominalis 011111112(123)1111111102000020101211011000111211101100003200010111???001201?0??123101???1??????N. flavibasis 01112111231111111102000020101211011100111211101110000300010?111110012010011123101???1??????N. rutila 011111112(123)1111111102000020101211011000111211101100000(23)000101110000012000001023101???1??????
8 • smithsonian contributions to zoology

elbowed (pronounced in Tanypezidae, with first flagellomere 

narrower and elongate; Figures 7, 8). Arista subbasal, short 

plumose to bare. Antennae closely spaced, usually touching.

Chaetotaxy: 0–2 reclinate fronto-orbital bristles; 1 ocellar bristle 

(divergent); 1 postvertical bristle (divergent); 1 inner vertical 

bristle; 0–1 short outer vertical bristle; 1–2 subgenal bristles; vibrissa absent. Frons with row of minute lateral setulae on orbital 

plate and with large space between ocellar tubercle and posterior 

margin. Occiput with long pile; gena narrow and with short, 

dense setulae.

Thorax. Black, occasionally with yellowish spots. 

Chaetotaxy: 0–1 postpronotal bristle; 2 notopleural bristles; 0–1 

presutural intra-alar bristle; 1 postsutural intra-alar bristle; 2 

supra-alar bristles; 1–3 dorsocentral bristles; 1 apical scutellar 

bristle (cruciate); 0–1 lateral scutellar bristle; 1 or more anepisternal bristles; proepisternal and katepisternal bristles absent. 

Scutum setulose with bare to sparsely setulose regions above 

wing and on large, distinct postpronotum. Prosternum broad, 

reaching proepisternum. Fore and mid coxae widely separated 

by katepisternum. 

Wing. Usually clouded apically, sometimes clear or 

more elaborately patterned (Figures 24–34). Subcosta complete 

(Tanypezidae) or fading apically (Strongylophthalmyiidae); 

subcostal break absent (Tanypezidae) or present (Strongylophthalmyiidae); costa reaching vein M1

. Anal cell and cell bm short 

and closed. A1

 + CuA2 nearly attaining wing margin. Calypter 

with long marginal hairs. Halter white with dorsal setulae on 

stalk sometimes dark and pronounced (Tanypezidae). 

Legs. Yellow in base color, variably pigmented. Long, 

slender, evenly setulose, and with few outstanding bristles, including several ventroapically on mid tibia. 

Abdomen. Black with anterior tergites sometimes 

variably yellow. Evenly setulose and setose with longer bristles 

on posterior margin of tergites; sternites 3–5 and posterior margin of sternite 2 bare medially in Strongylophthalmyia. Tergite 

1 fused to tergite 2 with suture evident laterally (Figure 49). 

Pleural membrane widely separating tergites and sternites and 

enclosing spiracles; minutely trichose (more dense and shaggy in 

Strongylophthalmyia).

Male Abdomen. Spiracles 1–6 below respective 

tergites (Figures 40, 45); seventh spiracle ventral, lateral to inner 

distal margin of sternite 8 (Figure 42). Sternite 8 dorsal, setose; 

sternite 7 left lateral and fused to sternite 8; sternite 6 subrectangular, ventral, and usually with right margin receding and left 

margin produced (Figures 35, 36, 45). Epandrium setose and 

dome-like (Figures 35, 37, 41–43, 52–53). Surstylus setose and 

fused to epandrium with suture absent. Cerci large, lobate, setose, and fused along most of length. Subepandrial sclerite long, 

narrow, and poorly sclerotized medially. Phallapodeme long, 

widest apically, and with medial process abutting hypandrium. 

Epiphallus composed of 2 sclerites that articulate with posterior margin of hypandrium and base of phallus; unfolds when 

phallapodeme slides distally through hypandrium to project 

distiphallus from abdomen (compare Figures 38 and 57 to Figure 44). Basiphallus absent or fused to distiphallus. Pregonite 

absent. Postgonite long, flexible, and band-like, with several apical setulae (sometimes bare); fused to inner surface of hypandrium at base, extends with apex of epiphallus when unfolded. 

Distiphallus minutely spinulose or with variable texturing, linear and short to very long, with membranous and sclerotized 

components. Ejaculatory apodeme small and narrow with base 

bulbous (Figure 39).

Female Abdomen. Segments 6–10 distinctly narrowed and elongate, forming tubular, slender ovipositor that 

is partially retractable (Figures 47–49). Tergite and sternite on 

segment 8 entirely divided longitudinally; tergite and sternite 7 

partially (Strongylophthalmyiidae) to entirely (Tanypezidae) divided. Tergite and sternite 10 narrow; sternite 10 with elongate, 

narrow clavate internal process extending into segment 8 (Figures 47, 48). Cerci short to relatively long and conical, curved 

in cross section and connected along length by membrane and 

fused along distoventral margin. Ventral receptacle small and 

pouch-like. Spermathecae (2) broadly ovate, well sclerotized, 

and pigmented. Also see Roháchek (1998) and Barber (2006).

Results and Discussion

Family Level Support

The majority rule bootstrap tree (100 replicates; maximum 

trees retained 10,000; TBR branch swapping) recovered the 

Strongylophthalmyiidae with a value of 90 and the Tanypezidae 

and Tanypeza with a value of 100. Tanypezidae + Strongylophthalmyiidae (bootstrap value 63) and Neotanypeza (bootstrap 

value 62) were moderately supported, and no sister group relationships were well supported within Neotanypeza (Figure 90). 

A heuristic search resulted in 45 trees that are 286 steps in length 

(Consistency Index = 0.41; Retention Index = 0.76); the strict 

consensus is presented in Figure 90. The optimized tree used here 

(Figure 91, top) is four steps longer than the most parsimonious cladograms and is discussed in the Neotanypeza section. See 

Tables 3–5 for a summary of the characters interpreted as synapomorphic for the family level groups, with departures from 

the phylogeny explained in the tables.

A sister group relationship between the Tanypezidae and

Strongylophthalmyiidae was recovered in all trees, being moderately supported in the analysis by a number of synapomorphies 

[9, 14, 43, 74, 75, 77, 79, 84, 85, 90; and tentatively 42, 69, 

76], most of which were male and female genitalic. There were 

relatively few external synapomorphies to unite the two families, 

and all of these are regularly seen elsewhere in the Acalyptratae, 

making them somewhat impractical for diagnosis. Furthermore, 

the two families together were not as well defined as each family was by itself: Tanypezidae: [2–5, 7–9, 11, 13, 16, 18, 19, 

22, 24, 29, 30, 31, 35, 37–41, 45, 60, 61, 64, 68, 74, 77, 87]; 
number 641 • 9

Strongylophthalmyiidae: [1, 14(2), 21, 26, 28, 34, 59, 66, 72, 

78, 80, 82, 83, 86, 88, 89, 91]. The increased character support 

for the separate lineages, combined with the increased number 

of external diagnostic features, makes it more useful to retain 

the two families as separate entities, although recognizing their 

sister group relationship. A key to the genera of Tanypezidae

and Strongylophthalmyiidae is presented below in light of these 

reevaluated characters.

A convergence of veins R4+5 and M1

 near the wing apex [40] 

may be another synapomorphy of the two families since it is present in both the Tanypezidae and Nartshukia (Shatalkin, 1993), 

but a revision of the Strongylophthalmyiidae will be necessary to 

determine whether this state is independently derived in the latter. A convergence of these veins is also seen in putatively related 

out-groups (possibly a result of a reduction in wing size), as well 

as in the Micropezidae and Neriidae, which were not included in 

the analysis. An enlarged and expanded prosternum [14] is also 

synapomorphic for the two families, although a similar state is 

seen in the Nothybidae. Strongylophthalmyia was not supported 

as monophyletic, but the analysis was not designed to test this 

hypothesis and should be treated separately in a future study. 

Of the out-groups examined, the morphology of Nothybus Rondani (the only genus in the family Nothybidae) was 

the most interesting, as this taxon shared a number of features 

otherwise only found (or predominantly found) in the Tanypezidae and Strongylophthalmyiidae. The prosternum, like those 

in the latter two families, is large and plate-like, fully extending 

to the proepisternal margin; the ocellar tubercle is located near 

the midpoint of the frons (also found in the Megamerinidae and 

some Diopsidae; reversed in Nartshukia), and the surstylus is 

partially fused to the epandrium. Furthermore, like the Tanypezidae, Nothybus has an elbowed antenna with a long first flagellomere, the lateral margin of the face is tomentose, the hind 

basotarsomere has a slight ventrobasal swelling, some species 

have a reflective stripe (slightly purple iridescent) extending from 

the anterolateral margin of the scutum to the posterior margin of the pleuron, and there are velvety purple patches on the 

anterolateral margin of the frons (a silvery tomentose postocellar 

patch, more similar to that of the Tanypezidae, is also found in 

the Megamerinidae). Although these characters suggest a relationship between the Tanypezidae and Strongylophthalmyiidae, 

the modified prosternum is likely convergent because of the drastic modification of the thorax, and the absence of many of these 

states in the Strongylophthalmyiidae suggests that these characters have been independently derived. 

TABLE 3. Synapomorphies of Tanypezidae and Strongylophthalmyiidae.

Character no. Synapomorphic state

9 Occiput with long pile

14 Prosternum wide, reaching proepisternum

42 Cell dm extending to a point at point of attachment 

to dm-cua

43 Calypter large and lobe-like

69 Surstylus fused to epandriuma

74 One hypandrial bristle (although state often variable)

75 Postgonite elongate, band-like, setulose apically, and 

fused to hypandrium at baseb

76 Basiphallus indistinct, fused to base of distiphallus 

(reversed in N. rutila)

a

77 Epiphallus plate-like and made up of two articulating sclerites

79 Female tergite 6 widest medially

84 Female sternite 10 with elongate internal processb

85 Female cerci curved in cross section and largely to 

completely fused

90 Ventral receptacle with numerous lobes covered by 

dome 

aCharacters only tentatively assigned to the Tanypezidae + 

­Strongylophthalmyiidae (recovered as synapomorphies of the dubious clade 

Tanypezidae + Strongylophthalmyiidae + Loxocera in the analysis). 

bComplex characters also found in Formicosepsis.

TABLE 4. Synapomorphies of Strongylophthalmyiidae.

Character no. Synapomorphic state

1 Head orbicular; back of head concave above foramen 

14 Suture between prosternum and proepisternum 

absent

21 Postvertical bristle anteroclinate

26 Lateral scutellar bristles absent

28 Notal setulae long, erect, and yellow

34 Greater ampulla lost

59 Epandrium, surstylus, and sternites 7 and 8 without 

setulae; surface with “pebbled” texture

66 Male sternites 7 and 8 strongly oblique

72 Hypandrium with bifid process ventrally

78 Distiphallus long, segmented, with spinulose 

membrane and apical sclerotized section

80 Female segment 7 forming complete sclerotized tube

82 Female segments 7 and 8 widely separated by 

membranous region

83 Female segment 10 elongate

86 Only one well-developed spermatheca 

88 Base of spermatheca telescoped

89 Vaginal sclerite present

91 Spermathecal ducts fused near base
1 0 • smithsonian contributions to zoology

Of additional interest were the examined representatives 

of the superfamily Nerioidea in the families Cypselosomatidae (coded for in the matrix) and Pseudopomyzidae, which are 

quite divergent in external appearance but have similar terminalia. Males have a long, band-like postgonite fused to the 

inner posteroventral margin of the hypandrium [75], and females have a rodlike internal process on sternite 10 [84] that 

is longer and more strongly sclerotized in the Tanypezidae and 

Strongylophthalmyiidae. These complex genitalic features are 

strongly suggestive of a much closer relationship with families 

in the Nerioidea, as previously posed by McAlpine (1997b), and 

preliminary analyses show them to be common throughout the 

superfamily. To investigate these and other similarities, further 

studies of family level relationships in the Nerioidea and Diopsoidea are currently being developed by the author.

Status of Enderlein’s Genera

On the basis of the present analysis, none of Enderlein’s 

genera—Polphopeza (synonymized by Hennig), Scipopeza, syn. 

nov., Tripolphopeza, syn. nov., or Tritanypeza, syn. nov.—reflect 

the evolutionary history of the family, and they are here treated 

as junior synonyms of Neotanypeza. In fact, very few sister 

group relationships were well supported within the genus at all. 

Furthermore, the monotypic Tripolphopeza appears to be defined on a misdiagnosed character, as N. elegantina (=N. apicalis) has two pairs of dorsocentrals, not three, and the type species 

of Scipopeza (N. grandis) is polymorphic for its defining feature 

(number of fronto-orbital bristles). 

Tanypeza

Tanypeza was recovered as monophyletic and the sister 

group to Neotanypeza in all trees. It was supported by stout 

black bristles on the hind trochanter and hind femur of the 

male [33], a medial pruinose stripe on the scutum [46], a large, 

broadly rounded surstylus [70], swollen, medially touching aedaegal guides on the hypandrium [73] (Figure 42), and a characteristic phallus [78] (Figure 44). 

Neotanypeza

Neotanypeza was heavily supported as monophyletic in 

all trees derived from the analysis. Contrasting this strong support for monophyly of the genus, none of the internal nodes 

had greater than 50% bootstrap support, as there was a paucity of consistent synapomorphies. As a result, most species were 

grouped on the basis of the relatively more numerous superficial and highly variable color characters, so a slightly longer 

tree (four steps longer; Figure 92) was selected as the working 

phylogenetic hypothesis since it instead favored chaetotaxy and 

other structural characters. A straight (not deviated) hind tibia 

[15] and stout postverticals [20] are interpreted as ancestral and 

result in the basal placement of the fossil N. dominicana. Neotanypeza is now defined by a wide first flagellomere [10] (variably modified), a bent hind tibia [15] (excluding the fossil N. 

dominicana), two dorsocentral bristles [25] (one or three bristles 

secondarily derived in several species), an entirely tomentose 

meron [30], a swollen basal protuberance on the hind basotarsomere [32], a strongly pointed posterior margin of cell dm (not 

a straight line from one end to the other) [42], stout, pointed 

setulae on the surstylus [71] (lost in several species), and a small, 

medially membranous and laterally spinulose distiphallus [78] 

(state unknown in N. dominicana). 

TABLE 5. Synapomorphies of the Tanypezidae (color characters 

excluded).

Character no. Synapomorphic state

2–5, 7, 8, “Tanypezidae-type” frons

9 Occiput with long white pile

11 Clypeus long and narrow but flat and broad 

anteriorly

13 Buccal cavity large

16 Fore tarsal claws larger than those on mid or hind 

legs

18 Outer vertical bristle absent

19 Ocellar bristle reduced

22 Postpronotal bristle present (reversed in some 

species)

24 Supra-alar bristle present (reversed in some species)

29 Thorax with broad silvery tomentose stripes

30 Posterior margin of meron tomentose

31 Thorax with long pile on postpronotum, 

anepisternum, anepimeron, and katepisternum

35 Strong row of dark bristles on stalk of halter 

extending to base of knob (few weak scattered 

brown bristles near base of stalk in Nothybus; many 

Diopsoidea with long row of white hairs)

37 Subcostal break absent

38 Subcosta complete

39 Vein R1 setulose dorsally

40 Veins R2+3 and M1 converging apically

41 Vein CuA2 strongly bowed

45 Lower calypter with hairs short, dense, and “furry”

60 Sternite 2 transversely divided; posterior margin of 

anterior sclerite with two emarginations

61 Tergite 2 with long, stout anterolateral bristles

64 Male sternite 6 with narrow, shining, left sublateral 

sclerotization

68 Male sternite 8 silvery tomentose

74 Loss of all hypandrial bristles (one pair recovered in 

some Neotanypeza)

77 Epiphallus composed of two large folding plates

87 Spermatheca papillose
number 641 • 1 1

Key to the World genera of Tanypezidae and STRONGYLOPHTHALMYIIDAE

1. Head globose and approximately as high as long [Figures 1, 34]. Frons shining, without raised disc behind tubercle. Clypeus band-like. Occiput shining with pile yellow. Outer vertical bristle present. Lateral scutellar bristle absent. Thorax 

long and narrow, without silvery tomentose stripes. Subcosta incomplete. Subcostal break present [Figure 24]. Vein R1

bare. Veins R4+5 and M1

 usually parallel apically but convergent in Nartshukia [Figure 34]. Vein CuA2 slightly bent. Greater 

ampulla absent. Tergite 2 without outstanding lateral bristles. Sternite 2 entire. Male sternite 8 not tomentose. Epandrium 

and surstylus without setulae, only strong bristles, and surface with “pebbled” texture [Figures 35–37]. Female tergite and 

sternite 7 fused, with sternite 7 narrowly divided (unknown for Nartshukia). One well-developed spermatheca. . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongylophthalmyiidae, 2

Head higher than long and back of head flat [Figures 7–9]. Frons extensively tomentose and velvety with tomentose disklike shelf behind tubercle. Anterior margin of clypeus broad. Occiput tomentose with pile white. Outer vertical bristle 

absent. Lateral scutellar bristle present. Thorax stout and broad with silvery tomentose stripes [Figures 2–6]. Subcosta 

complete [Figure 26]. Subcostal break absent. Vein R1

 setulose. Veins R4+5 and M1 strongly converging to apex. Vein 

CuA2

 strongly curved. Greater ampulla present. Tergite 2 with long, stout anterolateral bristles. Sternite 2 usually divided 

transversely near anterior margin. Male sternite 8 silvery tomentose. Epandrium and surstylus with setulae and not textured as above [Figures 41–43]. Tergite and sternite 7 separate and strongly divided longitudinally. Two well-developed 

spermathecae [Figure 50]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Tanypezidae, 3

2. Ocellar tubercle positioned near center of frons. Ocellar bristle usually at least twice length of ocellar tubercle. Veins R4+5

and M1

 parallel apically [Figure 24]. Australian, Nearctic, Oriental, and Palaearctic regions. . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Strongylophthalmyia Heller

Ocellar tubercle near posterior margin of frons. Ocellar bristle as long as ocellar tubercle. Veins R4+5 and M1

 convergent 

apically [Figure 34]. Oriental region. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Nartshukia Shatalkin

3. Postvertical bristle as thick as inner vertical and at least 2⁄3 length. One dorsocentral bristle [Figure 2]. Scutum with wide, 

light, central tomentose stripe. Meron shining with posterior margin pruinose. Base of hind basotarsomere only barely 

swollen. Hind tibia entirely straight. Male with stout black bristles on posterior face of trochanter and posterior base of 

hind femur. Surstylus as long and broad as epandrium (seen laterally), without spine-like setulae [Figures 41–43]. Distiphallus much longer than wide [Figure 44]. Nearctic and Palaearctic regions. . . . . . . . . . . . . . . . . . . . Tanypeza Fallén

Postvertical thin and minute, sometimes not much longer than tubercle (well developed in fossil species). One, two, or 

three dorsocentral bristles [Figures 3–6]. Scutum variably patterned but rarely with central tomentose stripe. Meron covered with dense pruinosity. Base of hind basotarsomere with swollen ventral protuberance. Hind tibia with subbasal bend 

(straight in fossil species). Male without stout black bristles basally on hind legs. Surstylus relatively short and narrow, 

often with minute spine-like setulae along outer posterobasal margin [Figures 52, 53]. Distiphallus approximately two 

times longer than wide, sometimes slightly longer [Figure 56]. Neotropical region. . . . . . . . . . . . . Neotanypeza Hendel

SYSTEMATICS

Neotanypeza Hendel, 1903

Neotanypeza Hendel, 1903:203 [as subgenus]. Type species, Tanypeza elegans Wiedemann, by subsequent designation (Enderlein, 1913:225). 

Hendel, 1935:54; Hennig, 1936:32 [as genus]; Enderlein, 1936:41; 

Steyskal, 1967:1, 1987:774; Griffiths, 1972:132; Rohácek, 1998:170. h

Scipopeza Enderlein, 1913:228. Type species, Scipopeza grandis Enderlein, 

by original designation. Hennig, 1936:37; Enderlein, 1936:40; Steyskal, 

1967:2, 1987:774; Rohácek, 1998:170. Syn. nov. h

Polphopeza Enderlein, 1913:226. Type species, Polphopeza quadrisetosa

Enderlein, by original designation. Syn. Hennig, 1936:32; Enderlein, 

1936:40 [resurrection]; Steyskal, 1967:1 [as subgenus]. 

Tritanypeza Enderlein, 1936:42. Type species, Tritanypeza cubitofusca

Enderlein, by original designation. Steyskal, 1967:2 [as subgenus]. Syn. 

nov.

Tripolphopeza Enderlein, 1936:41. Type species, Polphopeza elegantina 

Enderlein, by original designation. Steyskal, 1967:2 [as subgenus]. Syn. 

nov.

Diagnosis

Thorax with pruinose and silvery tomentose patches; stripe 

extending from notopleuron to meron (Figures 3–6; only extending to katepisternum in Tanypeza, Figure 2). Frons velvety, black 

to purplish black, with silvery anterolateral reflective stripe and 

variably tomentose ocellar disc. Postvertical bristle small and 

thin (long and relatively stout in fossil species; Figures 7, 8). One 

to three dorsocentral bristles (only 1 in remainder of family). 

Base of hind basotarsomere with swollen ventral protuberance. 

Hind tibia with subbasal bend (straight in fossil species and remainder of family). Distiphallus composed of single segment 

with lateral spinulose texturing. 
1 2 • smithsonian contributions to zoology

Description

Head. Back of head and ocellar disc (variably tomentose) black; frons velvety black to purple medially and silvery tomentose anterolaterally, blending onto dark tomentose 

parafacial laterally (Figures 12–23). Male frons narrowed medially with sides converging at midpoint (Figure 12; compare to 

Figure 17). Foramen large, nearly 1⁄3 width of back of head and 

almost reaching buccal cavity. Face slightly convex (more pronounced below antennal base), grading smoothly into buccal 

cavity; pale with dark stripes laterally and dorsomedially (fading ventrally). Gena and occiput silvery tomentose with long, 

white pile. Scape small; pedicel not much longer than wide at 

apex; first flagellomere strongly downturned (i.e., antenna elbowed), length more than twice width, shape ovate to narrow 

and subrectangular (Figures 7, 8). Clypeus long and narrow 

with apex broad and sharply curved. Postocular bristle reduced 

to few indistinct dorsolateral hairs. Inner vertical bristle well 

developed, outer vertical absent. Postvertical bristle usually 

small and hairlike (well developed in fossil species). Ocellar 

bristle as long as ocellar tubercle. Zero to two (0–2) frontoorbital bristles, positioned medially near midpoint of frons. 

Pedicel with dominant dorsal marginal bristle. Vibrissa absent; 

genal bristles reduced. Two (2) subgenal bristles. Remainder of 

head tomentose or velvety. 

Thorax. Black, sometimes with yellowish spots (Figures 3–6). Chaetotaxy: 0–1 postpronotal bristles; 2 notopleural 

bristles; 0–1 presutural intra-alar bristles; 1 postsutural intraalar bristle; 2 supra-alar bristles; 1–3 dorsocentral bristles; 1 apical scutellar bristle (cruciate); 1 lateral scutellar bristle; 1 strong 

anepisternal bristle, often accompanied by a number of weaker 

marginal bristles; proepisternal and katepisternal bristles absent. Notum covered with reflective pruinosity, with pronounced 

silvery tomentose patches on posterior margin of scutum and 

laterotergites (excluding anterior half of katatergite) and with 

stripe extending from lateral margin of notopleuron to anepisternum, katepisternum, and meron. Greater ampulla present 

(present in all Tanypezidae, not absent, as noted in Apigian and 

Thompson, 2010).

Wing. Usually clouded distally, sometimes clear or 

more elaborately patterned (Figures 26–33). Subcosta complete 

and subcostal break absent. Calypter with long yellow or brown 

marginal hairs. Vein CuA2

 strongly bowed. R4+5 and M1 strongly 

convergent. Cell dm strongly pointed at dm-cu. Vein R1

 setulose 

dorsally. Stalk of halter with dark dorsal setulae.

Legs. Yellow in base color and variably pigmented. 

Long, slender, evenly setulose, and with few outstanding bristles: 

mid tibia with several ventroapical bristles. Fore tarsal claws 

larger than those on mid or hind legs. Hind tibia usually bowed 

at base (straight in fossil species). Hind basotarsomere with 

swollen ventrobasal protuberance. 

Abdomen. Black, sometimes with anterior tergites 

variably yellow. Bristles usually black, sometimes yellow on anterior tergites or entirely yellow. Evenly setulose and setose, with 

long, stout anterolateral bristles on tergite 2. Anterior section of 

sternite 2 usually differentiated into short, wide separate sclerite 

with 2 deep posterolateral emarginations (Figure 55).

Male Abdomen. Sternite 8 dorsal, silvery tomentose; sternite 7 left lateral and sometimes indistinct (Figure 52). 

Sternite 6 rectangular, usually with right margin receding and 

left margin produced (sometimes symmetrical) and sometimes 

with narrow, shining, left sublateral sclerotization (Figure 54). 

Epandrium setose and dome-like (Figures 52, 53). Surstylus setose and fused to epandrium with suture absent. Cerci large, 

lobate, setose, and fused along most of length. Subepandrial 

sclerite narrow and band-like. Phallapodeme long and clavate 

with medial process abutting hypandrium (Figures 56, 57). 

Basiphallus U shaped with narrow anterior bridge, fused to 

base of distiphallus. Epiphallus composed of 2 large folding 

sclerites that articulate with posterior margin of hypandrium 

and base of phallapodeme; unfolds when phallapodeme slides 

distally through hypandrium to project distiphallus from abdomen. Pregonite absent. Postgonite long, flexible, band-like, 

with several apical setulae and fused to inner face of hypandrium at base; extends with apex of epiphallus when unfolded. 

Distiphallus minutely spinulose and membranous with sclerotized medial and lateral bands (sometimes fused); length usually twice width, sometimes longer. Ejaculatory apodeme small 

and narrow with base flattened.

Female Abdomen. On the basis of dissections of 

N. elegans (Figures 47–50) and N. montana. Spiracles in lateral 

membrane on segments 1–7. Segments 6–10 distinctly narrowed 

and elongate, forming slender tubular ovipositor with long membranous intersegmental regions (minutely trichose on basal half) 

between segments 6 and 7 and segments 7 and 8. Tergite 6 widest before midpoint (not at base). Tergites and sternites on segments 7 and 8 entirely divided longitudinally; setose on segment 

7 and with several minute setulae on segment 8; membrane on 

segment 7 with numerous longitudinal wrinkles on distal half 

and minutely trichose on basal half. Segment 9 absent. Tergite 

10 narrow, pointed at apex, tapering to base and with 1 pair of 

apical bristles; sternite 10 subtriangular with lateral wrinkles, 2 

apical bristles, and with elongate, narrow, spoon-shaped internal 

process extending into segment 8. Cerci short to relatively long 

and conical; curved in cross section, connected along length by 

membrane, and nearly fused along distoventral margin. Spermathecae (2) dark, spherical, and covered with minute papillae 

(excluding apex). Spermathecal ducts with separate attachment 

to genital chamber, weakly sclerotized, clear, tapered at base and 

apex, and approximately 3 times length of spermathecae. Ventral 

receptacle lightly pigmented with thin dome covering transverse 

series of small lobes.
number 641 • 1 3

Key to the Extant Species of Neotanypeza

Note: Neotanypeza grandis is removed in the first couplet due to a sexual dimorphism in chaetotaxy.

1. Wing with single strong black stripe along anterior margin to level of subcostal cell [Figures 5, 28]. Large species (9.9–12.1 

mm). Face orange in ground color [Figure 22]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. grandis (Enderlein)

Wing clear, with anterodistal infuscation or with shorter brown apical stripe(s); if dark along anterior margin, dark band 

not extending past vein r-m [see Figure 29]. Body length usually less than 9.0 mm. Face yellow to white in ground color.

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 2

2. One dorsocentral bristle. Postvertical bristle minute [Figures 7, 8]. Postpronotal bristle usually absent. . . . . . . . . . . . . 3

Two dorsocentral bristles. Postvertical bristle minute [Figures 7, 8]. Postpronotal bristle present [Figure 8]. . . . . . . . . . 8

Three dorsocentral bristles. Postvertical bristle very thin and slightly longer than tubercle. Postpronotal bristle present 

[Figure 8]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15

3. Wing with wide oblique stripe in male [Figure 31], dark on distal 1⁄4 in female [Figure 30]. Hind basotarsomere light yellow with brown apex. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. dimorpha (Hennig)

Wing, if patterned, with shorter and paler apical stripes (usually much faded to absent). Hind basotarsomere dark brown 

or only yellowish at base. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 4

4. Fronto-orbital bristles absent. Wing dusky to strongly patterned. Distiphallus short with deep, clear distomedial emargination [Figure 72]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5

One pair of fronto-orbital bristles. Wing clear or with anterodistal infuscation. Distiphallus long to relatively short, but 

always dark. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 6

5. Wing dusky with anterodistal margin darker. Fore tibia and distal half of fore and mid femora black. Sternite 6 with long 

left lateral sclerotized bar and short basal right lateral spot [Figure 70]. Surstylus almost entirely bare on outer face and 

slightly constricted at base [Figure 71]. Sides of distiphallus straight, slight converging apically and with lateral sclerotized 

bands meeting medially; with floating medial plate between lobes of hypandrium [Figure 72]. . . . . N. plotoplax, sp. nov.

Wing with small spot around apex of R2+3, sometimes in addition to light stripes along distal veins [Figure 32]. Fore tibia 

yellow, at least medially, and femora at most with dark narrow markings at apex. Sternite 6 with narrow left lateral band 

[as in Figure 54]. Surstylus setulose with posterior marginal setulae stout and pointed; not constricted basally [Figure 85]. 

Distiphallus bowed laterally and with lateral sclerotized bands widely separated; hypandrium without floating medial 

sclerite [Figure 86]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. vexilla, sp. nov.

6. Abdomen yellow with narrow brown stripes medially and posteriorly on tergites. First flagellomere narrow and light yellow. 

Mid and hind basotarsomeres yellow basally. Male sternite 6 asymmetrical and with dark sublateral sclerotized band on 

left side. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. leucothrix, sp. nov.

Abdomen entirely dark brown or pigment over entire body faded. First flagellomere broadly rounded and extensively 

pigmented. Mid and hind tarsi entirely dark brown. Male sternite 6 symmetrical (or nearly so), without dark band on left 

side. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 7

7. Body length 9.0 mm. Palpus brown and male calypter hairs brown to golden. Mid and hind femora usually brown at apex. 

Fore tarsi usually entirely brown, sometimes with basal segment partly yellow. Surstylus broadly rounded, evenly setose, 

and with stout, pointed posterior marginal setulae [Figure 74]. Distiphallus with relatively broad, smooth medial section 

[Figure 75]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. posthos, sp. nov.

Body length 7.5 mm. Palpus and male calypter hairs yellow. Mid and hind femora brown on distal half with apex pale. 

Fore tarsi light yellow with apical segment and apex of penultimate segment brown. Surstylus more abruptly tapered on 

distal half, setose posterodorsally and without pointed setulae along posterior margin [Figure 83]. Distiphallus with very 

narrow, smooth medial section [Figure 79]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. symmetros, sp. nov.

8. Ocellar disc shining with central tomentose spot [as in Figure 20]; sometimes also with very narrow marginal tomentose 

band. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9

Ocellar disc usually entirely tomentose; rarely shiny medially or along anterior margin. . . . . . . . . . . . . . . . . . . . . . . . 10

9. Frons velvety anteromedially. Tomentose spot on ocellar disc longer than wide [as in Figure 20]. Clypeus and palpus dark 

brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. elegans (Wiedemann)

Frons shining anteromedially. Tomentose spot on ocellar disc as wide as long [Figure 19]. Clypeus and palpus yellowish. 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. micans, sp. nov.

10. Hairs on calypter brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11

Hairs on calypter yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 12
1 4 • smithsonian contributions to zoology

11. Fronto-orbital bristle absent. Fore tibia entirely light yellow. Anterior margin of ocellar disc shining. . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. nigrithrix, sp. nov.

Fronto-orbital bristle present. Basal half of fore tibia brown. Ocellar disc entirely tomentose. . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. quadrisetosa (Enderlein)

12. Clypeus yellow, at least in part, and palpus usually also yellow. South of Mexico, specimens with thorax yellow, at least 

on scutellar margin, and with male frons densely golden haired and with deep central sulcus [Figure 14]. Lateral bristles 

on male tergite 1 yellow to light brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. apicalis (Wiedemann)

Palpus, clypeus, and thorax entirely dark brown (clypeus and palpus rarely yellowish). Male frons dark to silvery tomentose and never with open central sulcus. Lateral abdominal hairs sometimes yellow on male tergite 1, occasionally extending to tergite 2 or 6. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13

13. Frons and thorax (excluding anterior margin) silvery tomentose [Figure 23]. Two pairs of fronto-orbital bristles. Femora 

and tibiae entirely yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. argentia, sp. nov.

Frons and thorax with silvery tomentose regions reduced to discrete patches. Usually 1 pair of fronto-orbitals. Femora and 

tibiae brown in part. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 14

14. Distal 2/5 (or more) of fore and mid femora dark brown; apex of hind femur dark. Tibiae dark brown with fore tibia 

yellow on inner distal face. Hind coxa brown (yellow in Mexican specimens). Frons (excluding reflective patches) black. 

Antenna black with base of first flagellomere orange. Abdominal bristles sometimes yellow laterally on tergites 1 and 2. 

Distiphallus longer than surstylus [Figure 68]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. marshalli, sp. nov.

Femora entirely yellow with apices sometimes brown. Fore tibia light yellow to brownish at base and mid and hind tibiae 

with yellow and brown bands. Hind coxa yellow. Frons (excluding reflective patches) purple or black with purple tint. 

Antenna orange to yellow, sometimes with apex of first flagellomere infuscated; pedicel rarely brown in Peruvian males. 

Abdominal bristles sometimes yellow along entire length of pregenitalic tergites laterally. Distiphallus not much longer 

than wide. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. montana (Enderlein)

15. Palpus brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 16

Palpus yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19

16. Ocellar disc shining with medial tomentose spot [Figure 20]; rarely only with anterior margin shining around ocellar 

tubercle. Anterolateral tomentose triangle on frons not extending much past ptilinal suture. . . . . . N. alopecia, sp. nov.

Ocellar disc predominantly or entirely tomentose. Anterolateral tomentose triangle on frons extending nearly to level of 

anterior fronto-orbital or beyond. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17

17. Mid tibia yellow. Fore femur entirely yellow or with inner distal spot. Ocellar disc usually with shining region posterior 

to tubercle as long as diameter of ocellus. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. callitarsis (Rondani)

Mid tibia brown, at least faintly on basal half. Fore femur often with inner distal spot and sometimes with outer distal 

spot. Ocellar tubercle at most with narrow shining stripe along anterior margin. . . . . . . . . . . . . . . . . . . . . . . . . . . . . 18

18. Mid and hind tibiae usually yellow on distal half or more, rarely banded, and never entirely brown. Mid and hind tarsi 

dark brown with basal half or more of basal segment yellow (sometimes gradually fading to yellow). Fore femur almost 

always yellow on inner surface. First flagellomere variable in shape and color. . . . . . . . . . . . . . . N. claripennis (Schiner)

Mid and hind tibiae always with dark distal band (often fainter on mid leg), sometimes entirely dark brown [Figure 4]. 

Mid and hind tarsi entirely dark brown but usually yellowish to base on hind leg. Fore femur usually with inner distal 

spot or complete apical band. First flagellomere broadly rounded and apically infuscated; rarely narrow and orange (some 

Central American specimens). . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. dallasi (Shannon)

19. Abdominal tergites entirely brown. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 20

At least abdominal tergite 2 yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

20. Anterolateral tomentose triangle on frons not extending much past ptilinal suture [Figure 21]. Bristles on abdomen dark 

brown, with lateral bristles on female tergite 1 sometimes yellow. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. flavicalx (Enderlein)

Anterolateral tomentose triangle on frons extending at least to level of anterior fronto-orbital. Pale abdominal bristles 

always present, at least laterally on first tergite. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21

21. Pale lateral abdominal bristles restricted to first tergite. Mid tibia yellow. Mid and hind basotarsomeres yellow to base. 

Ocellar disc with small, shining spots lateral to posterior ocelli. . . . . . . . . . . . . . . . . . . . . . . . N. ochrifemur (Enderlein)

Pale abdominal bristles found along length of abdomen laterally. Mid tibia usually with basal band and sometimes also 

with distal band. Mid and hind basotarsomeres usually entirely brown. Ocellar disc entirely tomentose [Figures 16, 17].

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. ornatipes (Bigot)

22. Anterolateral tomentose triangle on frons not extending much past ptilinal suture (dorsal portion as long as wide at widest

point, as in Figure 21) or ending before level of anterior fronto-orbital. Fore femur always with strong, dark inner distal 
number 641 • 1 5

spot. Abdomen dark brown with only lateral regions of tergites 1 and 2 (anterior half) yellow. N. flavibasis (Enderlein)

Anterolateral tomentose triangle on frons extending to level of anterior fronto-orbital or further [Figure 18]. Fore femur 

entirely yellow, sometimes with dark inner distal spot. Abdomen with yellow spots at least on tergites 2 and 3. . . . . . 23

23. Male with anterolateral reflective triangle on frons white and not extending past level of posterior fronto-orbital [Figure 

18]. Male postvertical bristle brown. Female thorax dark brown. Prosternum brown or yellow. Clypeus dark brown, 

sometimes yellow medially. Hind tibia with 1 pair of brown distolateral spots. Face with distinct dark lateral and dorsomedial stripes. Distiphallus short, undivided, and fused to basiphallus [as in Figure 56]. . . . . . . . . . . . . . . . . . . . . . . . . . 

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. abdominalis (Wiedemann)

Male with anterolateral reflective triangle on frons tan colored and extending past posterior fronto-orbital [Figure 15]. 

Male postvertical bristle yellow. Female pleuron and notum with large yellow spots. Prosternum yellow. Clypeus yellow. 

Hind tibia yellow on distal half. Dark lateral and dorsomedial stripes on face very faint. Distiphallus long, bifid, and 

separate from basiphallus [Figure 78]. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . N. rutila (Wulp)

Descriptions of Extant Neotanypeza Species

Neotanypeza abdominalis

(Wiedemann), stat. reinst.

FIGURE 18

Tanypeza abdominalis Wiedemann, 1830:529. 

Tanypeza (Neotanypeza) abdominalis.—Enderlein, 1913:225.

Neotanypeza abdominalis.—Hennig, 1936:33.

Polphopeza abdominalis.—Enderlein, 1936:40; Steyskal, 1967:2 [as synonym of apicalis Wiedemann].

Male (Figure 18). Body length 5.5–6.9 mm. Anterolateral tomentose triangle on frons extending to level of 

anterior fronto-orbital. Ocellar disc fully tomentose. Anterior 

fronto-orbital relatively stout and 2⁄3 length of posterior frontoorbital. Postvertical bristle minute. First flagellomere narrow 

and light yellow to relatively thick and orange. Clypeus dark 

brown and palpus yellow. Postpronotal and presutural intra-alar 

bristles present. Three dorsocentral bristles. Anepisternum with 

only 1 dark, dominant bristle. Calypter hairs pale. Wing clear. 

Legs yellow with mid coxa brown, fore tibia light yellow, mid 

tibia brown on basal half, and hind tibia with dark basal band 

and faint to distinct distal band; tarsi dark brown with basotarsomeres yellow on basal 2⁄3. Abdominal bristles dark brown 

with lateral bristles on tergite 1 yellow; abdomen dark brown 

with tergites 1 and 2 (excluding lateral spot) and anterior half of 

tergite 3 yellow with narrow medial stripe.

Variation. Nontype males sometimes with yellow 

lateral stripes on tergite 3 and anterior half or more of tergite 4. 

Bolivian male smaller (5.3 mm), with wing infuscated distally, 

mid and hind tibiae with dark basal and apical bands, basal 2⁄3

of fore basotarsomere light yellow, mid and hind basotarsomeres 

dark brown and lateral bristles on tergite 2 yellow. Reinhartz 

specimen from Brazil (UZMC) with anterolateral stripe on frons 

extending past posterior fronto-orbital, mid tibia entirely yellow, 

abdomen entirely yellow, excluding brown tergite 6, epandrium, 

and surstylus, and may represent an undescribed species.

Male Genitalia. Epandrium, cerci, and surstylus 

as described for N. claripennis, although stout setulae along posterior margin of surstylus only evident in CNC male. Phallus as 

described for N. claripennis except distiphallus more transparent 

and slightly tapered at apex.

Female. As described for male except as follows: 

body length 6.7–7.8 mm; first flagellomere rounded on anterior 

margin and distally infuscated; anterolateral tomentose triangle 

on frons slightly longer; clypeus sometimes yellow medially; prosternum sometimes yellow; tergites 2–4 variably yellow but always brown medially. 

Distribution. Bolivia, Brazil, Mexico, Paraguay.

Lectotype. BRAZIL. “Mus., Westerm.,” “TYPE” 

[red label], “T. abdominalis, Wied., Brasils, Lund” (1♂, UZMC).

Additional Material Examined. BOLIVIA. 

La Paz: Chulumani, Apa Apa Reserve, 2000 m, 16°21′15′′S, 

67°30′21′′W, 1.iv.2001, S. A. Marshall (1♂, DEBU). BRAZIL. 

“Sele[?] Lagoas, Reinhartz” (1♂, UZMC), “S. Paulo” [illegible], “x.30” (1♀, USNM), “S. Paulo, V.CRLS [?], Bromalho, 

X-39” (1♂, USNM), Maracajú, Mato Grosso, Brasil, ServicoFebre, Amarela, M. E. S. Bras., v.1937 (5♀, USNM), vii.1937 (2♀, 

USNM), vi.1937 (1♀, USNM), Nova Teutonia, 27°11B, 52°23L, 

300–500 m, F. Plaumann, 2.xii.1959 (1♀, CNC), x.1964 (1♂, 

CNC), 2.xii.1959 (1♀, CNC), 5.ii.1959 (1♀, CNC), 21.xii.1959 

(1♂, CNC), São Paulo: São Jose dos Campos, E. R. DePaula, 

23–30.viii.1997 (1♀, EMUS), 30.viii-6.ix.1997 (1♀, EMUS), 

22–29.x.1999 (1♀, EMUS), PR, Terra Boa, 27.xii.1983, J. A. 

Rafael (1♂, INPA). MEXICO. Morelos: Cuernavaca, 5 mi. N, 

15.viii.1954, 7500’[ft], J. G. Chillcott (1♂, CNC), Porterillos, 

Sin., 15 mi W El Palmito, 8.vii.1964, 5000 ft, J. F. McAlpine (1♀, 

CNC). PARAGUAY. Cazaapá, Hermosa, San Rafael Res[erve]., 

prop.[erty of] Lopez family, 26°18′29′′S, 55°45′03′′W, 80 m, 

FIT, 1–3.xii.2000, Z. H. Falin (1♀, DEBU), Villarica, F. Schade, 

i.1937 (1♂, USNM), v.1937 (1♀, USNM), ix.1937 (1♀, USNM).

Comments. Steyskal (1967) classified Neotanypeza 

abdominalis as a synonym of N. apicalis in his catalog without comment. This species is here recognized as valid, although 

its distinctness from other taxa with a yellow palpus and pale 

abdominal spots should be considered tentative because of a 
1 6 • smithsonian contributions to zoology

relative lack of reliable autapomorphies, particularly those of the 

male genitalia. There are also several “intermediate” phenotypes 

described below under N. rutila that somewhat agree with the 

above description. The color of the abdomen and the shape of 

the reflective triangle on the frons are the most useful features to 

characterize this species as currently defined.

Neotanypeza abdominalis was described from an unstated 

number of specimens from Brazil in the university collection in 

Lund, and the sex was given as female. Currently, there is a single 

male in Copenhagen, where the Lund material was deposited, 

which is here designated as the lectotype.

Neotanypeza alopecia, sp. nov.

FIGURES 6, 20

Male. Body length 5.9–7.6 mm. Reflective anterolateral triangle on frons with dorsal portion not much longer than 

wide at widest point. Ocellar disc shining with relatively large 

tomentose spot that is usually widest posterior to midpoint (spot 

often narrower in Brazilian specimens and usually larger than 

that found in similar species such as N. elegans); 1 pair of minute tomentose anterolateral spots rarely present. Anterior frontoorbital smaller than posterior fronto-orbital. Postvertical bristle 

slightly longer than ocellar tubercle (sometimes as long as disc) 

and thin. First flagellomere very narrow and light yellow. Clypeus and palpus dark brown. Postpronotal and presutural intraalar present. Three dorsocentral bristles. Anepisternum with 1 

dark, dominant bristle. Calypter hairs brown. Wing clear with 

distal third very lightly infuscated. Legs yellow with mid coxa 

brown, fore femur dark brown on distal 2/5, fore tibia light yellow, hind femur usually with narrow brown band past midpoint, 

and mid and hind tibiae brown on basal half with 1 pair of faint 

to distinct lateral subapical spots on hind leg; tarsi dark brown 

with basal 2⁄3 of first segment yellow. Abdominal bristles dark 

brown; abdomen dark brown. 

Male Genitalia. As described for N. claripennis 

except as follows: sternite 6 with sclerotized band extending 

transversely near distal margin. 

Female (Figures 6, 20). As described for male 

except as follows: body length 6.1–7.6 mm; reflective anterolateral triangle on frons barely extending past ptilinal suture; frons 

sometimes with shining medial spot behind reflective anterolateral triangles; calypter hairs yellow; first flagellomere thicker and 

infuscated along margins; band on hind femur sometimes faded, 

disappearing posteriorly; brown pigment on mid tibia and/or 

distal brown band on hind tibia sometimes faint to absent. 

Variation. If outer surface of fore femur yellow in 

females, band on hind femur faded to absent, distal half of hind 

tibia entirely yellow, and mid tibia usually with band faded to 

absent. One Brazilian female with fore femur entirely yellow. 

One Peruvian female with left anterior dorsocentral duplicated. 

One tentatively included Bolivian female (DEBU) with pilosity 

on ocellar disc widened, frons largely shining, with reflective 

anterolateral stripe reduced medially, femora and tibiae yellow, fore tarsi light yellow with terminal 3 segments becoming 

browner distally, and wing with dark subapical spot in second 

radial cell. Three Peruvian females (nontypes) with ocellar disc 

tomentose excluding shining region around tubercle; distal 1⁄3 of 

fore femur brown (except apex), band on hind femur distinct to 

faint, basal half of hind tibia brown, and mid tibia yellow. Three 

females from Bolivia and Peru (nontypes) with ocellar disc shining, with 1 pair of lateral tomentose stripes; frons shining anteromedially; band on hind tibia light brown; fore femur yellow with 

light inner distal spot; hind femur and mid tibia entirely yellow.

Etymology. The specific epithet is derived from the 

Greek for “loss of hair,” denoting the largely shining ocellar disc.

Distribution. Bolivia, Brazil, Colombia, Ecuador, 

Paraguay, Peru.

Holotype. ECUADOR. Napo: Res. Ethnica Waorani, 1 km S Onkone Gare Camp, Trans. Ent., 10.ii.1996, 220 

m, 00°39′10′′S, 76°26′W, T. L. Erwin et al., insecticidal fogging of 

mostly bare green leaves, some with covering of lichenous or bryophytic plants in terre firme [sic] forest, Project MAXUS, at trans. 

10, sta. 9, lot 1499 (1♂, USNM, for eventual deposit in Ecuador).

Paratypes. BOLIVIA. S. Inicua Riv., Alto Beni, 15–

18.i.1976, 1100 m, L. E. Pena (1♀, CNC), La Paz: Mapiri (5 km 

W; 15°17.8′S, 68°15.6′W, 750 m), 16.iii.2001, A. Freidberg 

(1♀, USNM), Guanay (8 km E, 15°30.3′S, 67°50.8′W, 510 m), 

13.iii.2001, A. Freidberg (1♀, USNM), 1 km E Guanay, 580 m, 

15°30.2′S, 69°52.3′W, 14.iii.2001, S. D. Gaimari (1♀, USNM), 

5 km W Mapiri, Arroyo Tuhiri, 750 m, 15°17.8′S, 68°15.6′W, 

18.iii.2001, S. D. Gaimari (1♀, USNM), Arroyo Tuhiri, GPS 

15°17.458′S, 68°15.487′W, 10.iv.2001, A. L. Norrbom (1♀, 

USNM), Cochabamba: Villa Tunari, Hotel Los Tuscanes, 

16°58.39′S, 65°23.79′W, 323 m, 4.ix.2000, S. D. Gaimari, ex. 

along trail, on vegetation (1♀, USNM), La Paz, Heath River Wildlife Centre, ~21 km SSW Puerto Heath, 12°40′S, 68°42′W, 29.iv–

12.v.2007, S. A. Marshall (1♀, DEBU), La Paz, Arroyo Tuhiri W 

Mapiri, 15°17′27′′S, 68°15′29′′W, 10.iv.2001, S. A. Marshall (1♀, 

DEBU; 1♀, CBFC). BRAZIL. Bahia, v.1929, Rock. Found. Lab., 

Shannon (1♀, USNM), São Sebastião, 12–13.x.1976, J. Kugler 

(1♀, TAU), Minas Gerais: nr. Timoteo, 19–26.xi.1997, E. R. DePaula (1♀, EMUS), Rio de Janeiro: x.1937–i.1938, R. C. Shannon 

(1♀, USNM), Rio de Janeiro, Dist. Federal, Servico Febre, Amarela, M. E. S. Bras., x.1937 (3♀, USNM), xii.1937 (1♀, USNM), 

vi.1938 (1♀, USNM), vii.1938 (8♀, USNM), viii.1938 (1♀, 

USNM), ix.1938 (2♀, USNM), Rio de Janeiro, viii.1938, R. C. 

Shannon (3♀, USNM), R. Janeiro, ii.1940, R. C. Shannon (1♀, 

USNM), R[io]. d[e]. Janeiro, Yel. Fev. Serv., M. E. S. Bras., R. C. 

Shannon, ix.1938 (1♀, USNM), x.1938 (2♀, USNM), “obs. 27” 

[country not stated, but morphology similar to that of Brazilian 

females] (1♀, USNM), Rondonia: Ouro Preto do Oestre, Igarapé 

Mandi, 6–12.vii.1995, Arm. Malaise, J. A. Rafael & J. Vidal (1♂, 

INPA). COLOMBIA. Cundina-marca, Finca Bella Vista, nr. Sasaima, 7.vi.1965, F. R. & D. L. Craig (1♀, CASC). ECUADOR. 

Zamora, 1500 m, 27–31.viii.1965, L. E. Pena (1♀, CNC), Napo: 

Misahualli nr. Tena, C. Brammer, 27.iv–2.v.2003 (1♀, EMUS), 
number 641 • 1 7

6–19.x.2001, Mal. Tr. (1♀, EMUS), Yasuni Res. Sta., 0°40.566′S, 

76°23.851′W, 250 m, 30.ix–11.x.2002, M.T., C. Brammer (1♀, 

EMUS), Coca, Napo R., v.1965, 250 m, L. Pena (1♀, CNC), 

Res. Ethnica Waorani, 1 km S Onkone Gare Camp, Trans. Ent., 

10.x.1994, 220 m, 00°39′10′′S, 76°26′W, T. L. Erwin et al., insecticidal fogging of mostly bare green leaves, some with covering of lichenous or bryophytic plants in terre firme forest, Project 

MAXUS, At trans. 3, sta. 7, lot 936 (1♀, USNM), 10.x.1994, at 

trans. 4, sta. 3 lot 942 (1♀, USNM), 4.x.1994, at trans. 2, sta. 3, 

lot 862 (1♀, USNM), 10.ii.1996, at trans. 10, sta. 2, lot 1492 (1♀, 

USNM), 30.vi.1995, at trans. 8, sta. 10, lot 1080 (1♂ 1♀, USNM), 

30.vi.1995, at trans. 8, sta. 8, lot 1078 (1♀, USNM), 21.vi.1994, 

at 9 x-trans, 82 m mark, lot 716 (1♂, USNM), 26.vi.1996, at 

trans. 8, sta. 3, lot 1593 (1♂, USNM), 2.x.1996, at trans. 6, sta. 

1, lot 1711 (1♂, USNM), 4.ii.1996, at trans. 2, sta. 3, lot 1413 

(1♂, USNM), Orellana, Res. Ethnica Waorani, 1 km S Onkone 

Gare Camp, Trans. Ent., 216.3 m, 00°39′25.7′′S, 76°27′10.8′′W, 

T. L. Erwin et al., insecticidal fogging of mostly bare green leaves, 

some with covering of lichenous or bryophytic plants in terre 

firme forest, Project MAXUS, 12.ii.1996, lot 1513 (1♀, USNM), 

23.vi.1996, at trans. 10, sta. 9, lot 1679 (1♀, USNM), 19.i.1994, 

at trans. 6, sta. 5, lot 574 (1♂, USNM), Depto. Orellana: Tiputini Biodiversity Station, 00°37′55′′S, 76°08′39′′W, 220–250 m, 

T. L. Erwin et al., insecticidal fogging of mostly bare green leaves, 

some with covering of lichenous or bryophytic plants in terre 

firme forest, 5.ii.1999, trans. 9, sta. 5, lot 2085 (1♀, USNM), 

9.ii.1999, at trans. 2, sta. 4, lot 2013 (1♀, USNM). PARAGUAY.

Inst. Agr. Nac. Caaoupe, Depto. Cordillera, 12.xii.1980 (1♀, 

USNM). PERU. 40 mi E Tingo Maria, 12.xii.1954, E. I. Schlinger 

and E. S. Ross (1♀, CASC), Monson Valley, Tingo Maria, E. I. 

Schlinger and E. S. Ross, 23.ix.1954 (3♀, CASC), 26.x.1954 (1♀, 

CASC), 11.xii.1954 (1♀, CASC), 23.xii.1954 (2♀, CASC), Huanuco, vic. Tingo Maria, 1–5.vi.1999, W. Hanson and S. Keller 

(2♀, EMUS), Chauchainago, 6.viii.1948, “1200,” J. Schuuke 

(1♀, CNC), Avispas, Madre de Dios, 400 m, L. Pena, 20–30.

ix.1962 (2♀, CNC), 1–15.x.1962 (5♀, CNC), Madre de Dios, 

Manu, Rio Manu, 250 m, Pakitza, 12°7′S, 70°58′W, 9–23.

ix.1988, A. Freidberg (6♀, USNM), Madre de Dios, Manu, Rio 

Manu, Cocha Salvador, 240 m, 14.ix.1988, W. N. Mathis (1♀, 

USNM), A. Freidberg (1♀, USNM), Quincemil, Cuzco, 1–15.

xi.1962, 700 m, L. Pena (2♀, CNC), 15–30.x.1962 (1♀, CNC), 

Quincemil, Cuzco, 13–31.viii.1962, 780 m, L. Pena (1♀, CNC), 

Chanchamayo, Dp. Junin, 1200 m, 26.iv.1918, J. M. Schuuke 

(1♀, USNM).

Additional Material Examined. Country 

unknown. [illegible] (1♂, USNM).

Material with Medially Shining Ocellar 

Disc. BOLIVIA. Huachi Beni, W. M. Mann, ix, “Mulford Bio. Expd. 1921–22” (1♀, USNM), Sta. Cruz, Buena Vista, 

26.ii–8.iii.1999, Mal. Tr., F. Parker (1♀, EMUS). PERU. Avispas, 

Madre de Dios, 1–15.x.1962, 400 m, L. Pena (1♀, CNC).

Material with Predominantly Tomentose 

Ocellar Disc. PERU. Quincemil, Cuzco, 13–31.

viii.1962, 780 m, L. Pena (3♀, CNC).

Comments. Neotanypeza alopecia is a widespread 

and relatively common South American species characterized 

by three pairs of dorsocentral bristles, a predominantly shining 

ocellar disc, a short anterolateral pilosity on the frons, and often 

a medial band on the hind femur. Whereas females of this species 

are regularly collected near the ground, virtually every male was 

collected by canopy fogging.

The specimens from Peru with a largely tomentose ocellar 

disc blur the specific boundaries between Neotanypeza alopecia and N. callitarsis, although the color of the legs (particularly 

the hind femur) allows for tentative placement in the former. 

Although N. alopecia is usually distinct from congeners in coloration and tomentosity, specimens such as these and others 

without bands on the hind femur or mid tibia suggest that refinement of species boundaries is appropriate when additional data 

are available.

Neotanypeza apicalis (Wiedemann)

FIGURES 14, 33, 58–60

Tanypeza apicalis Wiedemann, 1830:529. 

Tanypeza (Neotanypeza) apicalis.—Enderlein, 1913:225.

Polphopeza elegantina Enderlein, 1913:227. Syn. nov.

Neotanypeza apicalis.—Hennig, 1936:33; Steyskal, 1967:2. 

Neotanypeza elegantina.—Hennig, 1936:34.

Polphopeza apicalis.—Enderlein, 1936:40.

Tripolphopeza elegantina.—Enderlein, 1936:39. 

Neotanypeza (Polphopeza) apicalis.—Steyskal, 1967:2. 

Neotanypeza (Tripolphopeza) elegantina.—Steyskal, 1967:2.

Male (T. apicalis Lectotype; Figure 33).

Body length 8.6 mm. Frons, pedicel, and scape with thick yellow tomentosity and long yellow pile, with pilosity extending 

around margins of ocellar disc; ocellar disc entirely tomentose; 

frons with deep central sulcus surrounded by long yellow incurved bristles. Anterior fronto-orbital bristle absent and posterior fronto-orbital between sulcus and tubercle. Postvertical 

bristle minute. First flagellomere narrow and light yellow to 

white. Face white. Palpus yellow and clypeus dark brown. Presutural scutum yellow tomentose excluding anterior margin and 

central stripe. Postpronotal bristle present and presutural intraalar bristle absent. Two dorsocentral bristles. Anepisternum with 

1 dark, dominant bristle. Wing dark along distal margin to M1

, 

with slight extension along R4+5. Calypter hairs yellow. Legs yellow with basotarsomeres yellow and second tarsomere becoming 

yellow toward base. Abdominal bristles dark brown with lateral 

bristles on first tergite yellow; abdomen yellow anterior to tergite 

6 with posterior margin brown on tergites 3–5; sternites yellow, 

becoming brownish closer to terminalia. 

Variation: Males (Figure 14). Nontype males 

differ as follows: body length 7.0; tomentose field on frons extending posteriorly between fronto-orbitals to surround anterior 

margin of ocellar tubercle; clypeus yellow; wing clear with distal 
1 8 • smithsonian contributions to zoology

1⁄4 lightly clouded (fading posteriorly); scutellum brown with 

margin yellowish; scutum brown with notopleuron, postpronotum and supra-alar region yellow and without large transverse 

tomentose patches; pleuron yellow with faint brown mottling on 

anepimeron, meron, and sutures of anepisternum and with dark 

spot on venter of greater ampulla; abdomen yellow from tergite 

1 to posterior 1⁄3 of tergite 3 and with narrow central stripe on 

tergites 2 and 3; tergite 2 brownish on posterior 3⁄4 of USNM 

specimen; posterior margin of tergite 2 brown in DEBU male, 

which is also yellow anterolaterally on tergites 4 and 5.

Male Genitalia (Figures 58–60). Externally 

as described for N. claripennis except as follows: sternite 6 with 

deep distal emargination to right side and without sclerotized 

sublateral band; sternite 7 narrow and with single dorsal bristle; 

surstylus more broadly rounded and with long pronounced field 

of stout pointed setulae along posterior margin; hypandrium 

bare; distiphallus short and weakly sclerotized medially past 

base, with 1 pair of pointed basolateral extensions and long apical membranous lobe.

Female (P. elegantina Holotype). Chaetotaxy as described for male. Body length 8.2 mm. Ocellar disc 

entirely tomentose. Frons black with purple tint, with yellowish 

to white reflective tomentose anterolateral triangle that almost 

reaches level of fronto-orbital. First flagellomere light yellow 

with orange tint. Wing only lightly pigmented along anterodistal 

margin. Palpus and clypeus yellow. Thorax mostly dark brown, 

with postpronotum and anterolateral margin of scutum yellow, 

scutum with long floating medial yellowish stripe, scutellum yellowish, and pleuron with faint yellowish mottling. Legs yellow 

with basal half of mid and hind tibiae dark brown; tarsi dark 

brown with fore basotarsomere light yellow on basal half and 

mid and hind basotarsomeres yellowish to base. Abdominal bristles and abdomen dark brown.

Variation: Females. DEBU and USNM females 

differ from P. elegantina type as follows: tomentose triangle on 

frons reaching level of fronto-orbital; clypeus brown laterally in 

DEBU female; USNM female with lateral margin of scutum yellow to alar base and with most of pleuron yellow with brown 

pattern as described above for nontype males; medial stripe on 

scutum absent; abdomen yellowish with segments 6 and 7 and 

lateral and posterior margins of anterior tergites dark (much 

darker in DEBU female). CNC female from Bolivia differs as follows: ocellar disc shining along midline; frons (excluding reflective anterolateral patch) nearly shining, only sparsely setulose; 

first flagellomere narrow and light yellow; palpus brown; clypeus 

yellowish; thorax brown with postpronotum lighter, margin of 

scutellum yellow, and prosternum and pleuron with yellowish 

mottling; mid coxa mostly yellow. 

Distribution. Bolivia, Brazil.

Lectotype [apicalis]. BRAZIL. “S. Catharina”: 

Lüderwaldl S. (1♂, UZMC). 

Holotype [elegantina]. BRAZIL. Santa Catharina [from description]: “Mus., Wester.”, “TYPE” [red label], 

“T. apicalis, Wied., Brasil, Lund” (1♀, PAN).

Additional Material Examined. BOLIVIA. 

S. Inicua Riv., Alto Beni, 15–18.i.1976, 1100 m, L. E. Pena (1♀, 

CNC), La Paz: Caranavi, ca.[~]10 km NW, road to Entel Tower, 

1400 m, 15°46′35′′S, 67°35′48′′W, 13.iv.2001, S. A. Marshall 

(1♀, DEBU). BRAZIL. Pico de Tijuca, D. Federal, “8-12-1940”, 

R. C. Shannon (1♀, USNM), Rio de Janeiro: Dist. Federal, 

23.iii.1940, R. C. Shannon (1♂, USNM), R.D.J., florestal de Tijuca, Agudeida Solidao, 23.i.1990, S. A. Marshall (1♂, DEBU). 

Comments. Polphopeza elegantina is here treated 

as a junior synonym of Tanypeza apicalis, as they both now appear to belong to a single morphologically variable species with a 

particularly pronounced dimorphism between the sexes, with the 

extreme phenotypes (represented by the primary types) blending 

together when additional specimens are taken into account. This 

synonymy is only tentatively applied here, however, and should 

be corroborated using additional data, preferably behavioral 

and/or molecular data. 

Of particular significance in this new synonymy is the color 

of the wings, thorax, and abdomen. Whereas the wing of the 

Tanypeza apicalis lectotype is strongly marked apically (Figure 

33), the other males (which have the same unique frons) have 

wings more similar to that of the holotype of Polphopeza elegantina. These other males also have the yellow clypeus, marginally 

yellow scutellum, and partially yellow pleuron found in that female lectotype. The DEBU and USNM nontype females are even 

more similar in appearance to the lectotype in having a predominantly yellow pleuron with brownish patches on the katepisternum, anepimeron, and meron, as well as a strong spot on the 

venter of the greater ampulla. These females also have a laterally 

brown clypeus, similar to that found in the T. apicalis lectotype 

male, as well as the partially yellow abdomen found in all males. 

This newly defined species can be diagnosed on the basis 

of an entirely tomentose ocellar disc (medially shiny in CNC female, which may not be conspecific), two dorsocentral bristles, 

an absence of the anterior fronto-orbital, a male frons with a 

deep central sulcus surrounded by long yellow incurved hairs, 

yellow calypter hairs, a yellow palpus (dark brown in CNC 

female), a yellow clypeus (sometimes brown laterally), basally 

yellow basotarsomeres (entirely yellow on fore leg in males), a 

narrow, pale first flagellomere, yellow femora, and a dark brown 

abdomen that sometimes has yellow spots. 

Tanypeza apicalis was described from an unstated number 

of males from Brazil in the university collection in Lund. Currently, there is a single male in Copenhagen (where the Lund 

material was deposited) that is here designated as the lectotype. 

Neotanypeza argentia, sp. nov.

FIGURE 23

Male (Figure 23). Body length 8.0 mm. Frons 

entirely silvery tomentose, excluding spot between ocelli and 

around base of vertical bristle. Only posterior fronto-orbital 

present. Postvertical bristle minute. First flagellomeres, wings, 
number 641 • 1 9

and lower half of head largely missing, with remaining portion 

of wing clear. Postpronotal bristle present and presutural intraalar bristle absent. Two dorsocentral bristles. Anepisternum 

with 1 dark, dominant bristle. Calypter hairs pale. Thorax dark 

brown with scutellum and mottling on prosternum and anepimeron yellow; scutum and scutellum silvery tomentose, excluding 

postpronotum and anterior margin of scutum. Legs yellow with 

posterior half of mid coxa brown and tarsi dark brown, excluding basal half of first segment. Abdominal bristles dark brown 

with lateral bristles on tergite 1 yellow; abdomen dark brown.

Male Genitalia. Not dissected; concealed under dirt.

Female. Unknown.

Etymology. The specific epithet is derived from the 

Latin for silver (argentum), referring to the unique silvery appearance of the head and notum.

Distribution. Brazil.

Holotype. BRAZIL. Chapada, “S. W. Williston 

collection” (1♂, AMNH).

Comments. Although the genitalia of the single 

known specimen are not visible and the head is partially destroyed, Neotanypeza argentia is here described as a new species 

because it has a highly unique, dense silver coating on most of 

the frons and notum that easily distinguishes it from congeners. 

Neotanypeza callitarsis (Rondani)

Tanipeza Callitarsis Rondani, 1850:181. 

Neotanypeza nigripalpis Hennig, 1936:36. Syn. nov.

Tritanypeza nigripalpis.—Enderlein, 1936:44.

Tritanypeza cubitofusca Enderlein, 1936:44. Syn. nov.

[Unplaced] callitarsis.—Steyskal, 1967:3. 

Neotanypeza (Tritanypeza) cubitofusca.—Steyskal, 1967:2.

Neotanypeza (Tritanypeza) nigripalpis.—Steyskal, 1967:2.

Male. Body length 6.5–8.4 mm. Reflective anterolateral triangle on frons extending to level of anterior frontoorbital. Ocellar disc tomentose; type males with shining spots 

posterolateral to tubercle. Anterior fronto-orbital bristle shorter 

than posterior bristle. Postvertical bristle slightly longer than 

ocellar tubercle and thin. First flagellomere very narrow and light 

yellow. Clypeus and palpus dark brown. Postpronotal and presutural intra-alar bristles present. Three dorsocentral bristles. Anepisternum with 1 dark, dominant bristle. Calypter hairs yellow. 

Wing clear, sometimes with distal third very lightly infuscated. 

Legs yellow with mid coxa brown, fore femur with broad inner 

subapical spot and faint to absent outer subapical spot, fore tibia 

light yellow, and hind tibia brown on basal half and with 1 pair 

of faint distolateral spots (often absent, including in T. callitarsis

and T. cubitofusca types); tarsi dark brown with basal half of 

first segment yellow. Abdominal bristles dark brown with lateral 

hairs on tergite 1 yellow; abdomen dark brown.

Variation. Brazilian and Costa Rican males with 

reflective anterolateral patch on frons extending to level of posterior fronto-orbital bristle, and postvertical bristle as long as 

ocellar disc. Brazilian male also with tibiae and femora entirely 

yellow. Mexican males with tibiae and femora yellow, excluding nearly vestigial spot on fore femur. Mexican and Guatemalan males with first flagellomere slightly thicker and infuscated 

past base.

Male Genitalia. External genitalia, phallus, and 

postgonite as described for N. claripennis.

Female. As described for male types except as follows: body length 5.3–7.9 mm; dorsal portion of anterolateral 

reflective stripe on frons not much longer than wide; ocellar disc 

always fully tomentose; first flagellomere slightly thickened and 

infuscated distally and along margins; wing nearly clear; fore 

femur entirely yellow; hind tibia yellow on distal half.

Distribution. Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, Guatemala, Mexico, Paraguay, Peru, 

Venezuela.

Holotype [callitarsis]. VENEZUELA. “Venezuela” (1♂, MRSN) [thorax in poor condition]. 

Holotype [nigripalpis]. PERU. Chanchamayo, 

12.i.1904 (1♂, SMT).

Lectotype [cubitofusca]. PARAGUAY. San Bernadino, K. Fiebrig S.V., ii [2.ii.1906 in original description], 

“Paratype” [red label] (1♂, ZMHU).

Additional Material Examined. ARGENTINA. Bemberg, 10.iv.1927, R. C. Shannon (1♀, USNM). BOLIVIA. La Paz: 5 km W Mapiri, Arroyo Tuhiri, 750 m, 15°17.8′S, 

68°15.6′W, 16.iii.2001, S. D. Gaimari (1♂, USNM), Guanay 

(8 km E; 15°30.3′S, 67°50.8′W, 510 m), 13.iii.2001, A. Freidberg (1♂, USNM). BRAZIL. Rondonia: 62 km SE Ariquemes, 

17–24.iii.1989, 180 m, W. J. Hanson (1♂, EMUS). COLOMBIA. Cundina-marca, Finca Bella Vista, nr. Sasaima, 14.iv.1964, 

P. R. Craig (2♂, CASC). COSTA RICA. Puntarenas: Fila Cruces, 

1200 m, 1.vii.1995, A. Picado (1♂, INBio). ECUADOR. Napo: 

Limoncocha, 15.vi.1977, P. J. Spangler and D. R. Givens (1♀, 

USNM), Res. Ethnica Waorani, 1 km S Onkone Gare Camp, 

Trans. Ent., 3.vii.1995, 220 m, 00°39′10′′S, 76°26′W, T. L. 

Erwin et al., insecticidal fogging of mostly bare green leaves, 

some with covering of lichenous or bryophytic plants in terre 

firme forest, Project MAXUS, at trans. 4, sta. 2, lot 1092 (1♀, 

USNM), Limoncocha, 0°24′S, 76°40′W, 9–16.iii.1976, 250 m, 

G. E. Shewell (1♂, CNC). GUATEMALA. Quezaltenango, 1850 

m, 30–31.vii.1972, J. Helava (1♂, CNC). MEXICO. Zimapan, 

viii.1962, F. M. Hull (1♀, CNC), SE Citlaltepeti, Orizaba, Ver., 

4.viii.1964, el. 4000, L. W. Swan (1♂, CASC), Oaxaca: just W 

of Chiapas border, 19 km W of Rizo de Oro ridge S Cerro Baul, 

1615 m, 28.iv.1972, D. E. Breedlove (1♂, CASC), Chiapas: Lagunas de Montebello, 21.ix.1991, 1580 m, D. M. Wood (1♀, 

CNC). PERU. Madre de Dios: Manu, Rio Manu, Cocha Salvador, 240 m, 14.ix.1988, A. Freidberg (2♂, USNM), Rio Tambopata Res., 30 air km SW Pto. Maldonado, 290 m, 16–20.

xi.1979, J. B. Heppner, subtropical moist forest (1♂, USNM). 

VENEZUELA. Rancho Grande, Edo., Aragua, 18–27.ii.1971, 

G. & M. Wood, 1100 m (1♀, CNC), “Akuriman Gran Sabana 

EB Venez”, xi–xii.1940, P. Anduze (1♂, USNM).
2 0 • smithsonian contributions to zoology

Comments. Tritanypeza cubitofusca and Neotanypeza nigripalpis are here included as junior synonyms of 

Tanipeza callitarsis because the type specimens vary only in pigment intensity on the wing and hind tibia. Other characters used 

by Enderlein (1936) to define these species have been found to be 

absent (width of male frons, other color characters) or insignificant (body length). 

Enderlein described Neotanypeza cubitofusca from a series 

of 11 specimens, 2 males and 9 females, which are here treated 

as syntypes. The only specimen examined from ZMHU is here 

designated as the lectotype.

Neotanypeza claripennis (Schiner), 

stat. reinst.

FIGURES 12, 13, 26, 27, 52–57

Tanypeza claripennis Schiner, 1868:247. Bigot, 1886:380; Wulp, 1897:362. 

Tanypeza (Neotanypeza) claripennis.—Hendel, 1903:203; Enderlein, 

1913:226.

Neotanypeza claripennis.—Hennig, 1936:34; Enderlein, 1936:41 [as a junior 

synonym of T. elegans].

Male (Figures 12, 13, 26, 27). Body length 6.4–

7.7 mm. Reflective anterolateral patch on frons reaching level 

of anterior fronto-orbital. Ocellar disc fully tomentose. Anterior 

fronto-orbital shorter than posterior bristle. Postvertical bristle 

minute. First flagellomere frequently pale in coloration and 

very narrow (as in type), but sometimes broadly rounded and/

or darkly pigmented (excluding paler base). Clypeus and palpus 

dark brown. Postpronotal and presutural intra-alar bristles present. Three dorsocentral bristles. Anepisternum with 1 or 2 dominant bristles. Calypter hairs yellow. Wing lightly clouded on 

distal 1⁄4–1⁄3 toward costa and rarely with distal margin or entire 

surface clear; only material from Venezuela and Trinidad ever 

with wing darker. Legs yellow with tibiae sometimes banded, but 

never entirely brown and often entirely yellow on distal half (as 

in type); fore femur almost never with outer distal spot; base of 

basotarsomeres yellow, but sometimes brownish on either mid or 

hind legs (some specimens from Trinidad and Ecuador with fore 

basotarsomere white). Abdominal bristles dark brown with lateral hairs on tergite 1 yellow, but some males with yellow lateral 

bristles on all pregenitalic tergites; abdomen dark brown with 

sternite 8 sometimes yellow. 

Male Genitalia (Figures 52–57). Sternite 6 

asymmetrical (slanted posteriorly on right side, slightly extended 

on left side, and with narrow left sublateral sclerotization), setulose (excluding right posterior quarter), and with long bristles 

on distal margin. Sternite 7 membranous and narrow. Sternite 

8 large, dome shaped, covered with silvery tomentose hairs, 

and heavily sclerotized and setulose with several bristles. Cerci 

broadest at base and nearly fused to apex (slightly emarginate). 

Surstylus approximately 2⁄3 height of epandrium and nearly as 

long; setulose posteriorly, with marginal setulae stout, pointed, 

and dark. Hypandrium U shaped with anterior margin (excluding medial point of weakness) and arms well sclerotized; 1 hypandrial bristle. Postgonite with 4 apical bristles. Distiphallus 

approximately 2/5 length of phallapodeme, with lateral margins 

sclerotized and toothed and with membrane sclerotized and 

slightly toothed medially. 

Female. As described for male, except several Costa 

Rican females with spot on fore femur faint; body length 5.3–

7.9 mm. 

Distribution. Argentina, British Honduras, Colombia, Costa Rica, Dominica, Ecuador, El Salvador, Guatemala, neotropical Mexico, Panama, Peru, Venezuela.

Lectotype. VENEZUELA. “Lindig, 1864, Venezuela”, “claripennis, Alte Sammlung” (1♂, NMW). 

Additional Material Examined. 101♂

306♀ [AMNH, CASC, CNC, DEBU, EMUS, INBio, NMW, 

QCAZ, TAMU, TAU, USNM].

Comments. Schiner (1868) described Neotanypeza 

claripennis from an unstated number of Brazilian specimens. 

Two identified specimens from Vienna, the original depository, 

were available for this study. One of these types was found to 

be conspecific with N. montana; the remaining specimen is here 

designated as the lectotype of N. claripennis. Both were collected 

in 1864 from Venezuela, not Brazil, as stated in Schiner (1868).

Neotanypeza claripennis and N. dallasi are, as currently 

defined, highly variable species with uncertain boundaries that 

likely contain cryptic species. One such potential “species” (currently classified as N. claripennis because of other specimens 

with intermediate morphologies, as mentioned below) consists of 

those specimens with a sharp subapical band on the wing (Figure 

27), a characteristic reflective anterolateral triangle on the frons 

(tan colored and extending past the posterior fronto-orbital), 

and an almost entirely yellow pruinose notum (Figure 13). A few 

specimens, however, have only the former two characteristics, 

several DEBU specimens of N. claripennis have a similar reflective patch on the frons, and one “typical” specimen of N. claripennis from Trinidad has a similarly yellow pruinose scutum. 

Furthermore, one male from Peru (USNM) with a clear wing 

and entirely yellow legs has the same pruinosity on the head and 

notum, although this pruinosity continues medially onto the abdomen to the midpoint of tergite 4. 

Other problematic specimens include a number of males 

and females collected from Chiapas, Guatemala, and El Salvador 

(TAU, USNM) that have a yellow mid coxa and prosternum, a 

dark brown palpus, and a broad, darkly infuscated first flagellomere. The mid tibia, pleuron, and shoulders are also variably 

yellow, the clypeus is sometimes yellow, and the abdomen is also 

variably yellow, ranging from that typical of other N. claripennis

to that of the palest N. rutila; furthermore, the legs of these pale 

specimens are often identical to those of N. dallasi. These specimens entirely blur the lines between Neotanypeza claripennis, 

N. dallasi, and several other taxa with three dorsocentrals and 

have been only tentatively included in this species; as such, this 

variation is not included in the definition above. The remaining 
number 641 • 2 1

phenotypes within N. claripennis and N. dallasi are characterized by a nearly random assortment of particular characters 

states, including the shape and color of the first flagellomere, the 

presence or absence of the apical and/or basal bands on the mid 

and hind tibiae and fore femur, the color of the basotarsomeres, 

and the relative amount of patterning on the wing. These difficulties typify the problems involved with delimiting many species 

of Tanypezidae, which are rarely entirely distinct from one another and are often separated by slight or continuous differences 

that obscure taxonomic boundaries.

For the current purposes of species circumscription and diagnosis, although Neotanypeza dallasi has darker, thicker antennae and darker tibiae and fore femora, these characters emerge 

independently (singly or in pairs) in specimens that are clearly 

much more similar to the lectotype of N. claripennis. These characters, however, are only entirely present in those specimens with 

complete basal and distal bands on the mid and hind tibiae and 

predominantly dark tarsomeres on the mid and hind legs (i.e., 

basotarsomere brown basally on mid leg and only yellowish to 

base on hind leg), so it is the combination of these characters that 

is here used to define N. dallasi. This tentative circumscription 

is clearly in need of reevaluation, preferably using support from 

independent character sets such as molecular data. Last, nearly 

all of the previously mentioned characters also vary within the 

similar N. callitarsis, which is distinguished by a yellow mid 

tibia and should be considered in the future when delimiting the 

boundaries of N. claripennis and N. dallasi. 

Neotanypeza dallasi (Shannon)

FIGURE 4

Tanypeza dallasi Shannon, 1927:41. 

Neotanypeza dallasi.—Hennig, 1936:37.

Tritanypeza Dallasi.—Enderlein, 1936:43.

Neotanypeza (Tritanypeza) dallasi.—Steyskal, 1967:2.

Male. As described for N. claripennis except as follows: body length 7.6–7.9 mm; first flagellomere broadly rounded 

and darkly infuscated (occasionally light orange in Costa Rican 

males and narrow and darkly infuscated in Argentinian males 

and females); anepisternum with 2 dark, dominant bristles; wing 

clear to lightly clouded on distal 1⁄4–1⁄3 (most Costa Rican specimens with wing lightly clouded, at least past midpoint); fore 

femur with dark inner distal spot, and mid and hind tibiae dark 

brown with yellow subapical band (most Central American material with distal 1⁄3 of fore femur dark brown, and most of these 

with mid and hind tibiae entirely dark brown; most Costa Rican 

males with darker wing and mid femur with dark subapical spot 

and light posterodistal spot); tarsi dark brown with basal half of 

fore basotarsomere yellow to base and very base of hind basotarsomere yellowish; only tergite 1 with yellow bristles. Mexican 

material with distal half of fore femur brownish to brown, and 

prosternum, proepisternum, and tergite 1 yellow (sometimes also 

tergites 2 and 3 yellow with central brown stripe); postpronotum 

yellowish and pleuron with some yellow mottling.

Male Genitalia. As described for N. claripennis. 

Female (Figure 4). As described for male, except 

sternite 8 brown and body length 6.1–8.1 mm.

Distribution. Argentina, Bolivia, Colombia, Costa 

Rica, Dominica, El Salvador, Guatemala, neotropical Mexico, 

Panama.

Lectotype. ARGENTINA. Tucumán, Quebrada de 

Lules, 2.ii.1928, Shannon (1♂, USNM).

Paralectotypes Examined. ARGENTINA. Same 

collection as lectotype (2♂, USNM).

Additional Material Examined. ARGENTINA. Tucuman E. del Ponte (1♂, USNM), Tucumán, 

18.iii.1929, Shannon and Del Ponte (1♀, USNM), Tucuman 

Ao. El Caleo, W. Alpachiri, 1000 m, 1.x.1968, L. Pena (1♀, 

CNC), Salta, 30 km E Salta, Campo Quijano, forest vestige at 

El Alisal, 20.ii.1992, S. A. Marshall (1♂, DEBU), forest remnant at El Alisal, 18.ii.1992, S. A. Marshall (1♂ 1♀, DEBU), 

Salta Rosario de Lerma, INESALT yard, Malaise, 16–28.ii.1992, 

S. A. Marshall (1♀, DEBU), Catamarca, Ao. El Pintado, S. La 

Vina, 650 m, 27–29.ix.1968, Pena (3♀, CNC), Catamarca, El 

Alto, 3.xi.1972, G. E. Bohart (1♀, EMUS). BOLIVIA. La Paz: 8 

km S Chulumani, Apa Apa, 1950–2100 m, 16°22′S, 67°30.4[′]

W, 23.iii.2001, S. D. Gaimari (1♀, USNM), 8 km S Chulumani, 

nr. Apa Apa, 1700–1800 m, 16°22′S, 67°30.4′W, 9.iii.2001, S. 

D. Gaimari (1♂, USNM), Jujuy: San Pedrode, 29.iv.1926 (1♀, 

USNM). COLOMBIA. Cundina-marca, Finca Bella Vista, nr. Sasaima, 7.vi.1965, P. R. & D. L. Craig (1♀, CASC), 19.iv.1965, 

P. R. Craig (1♀, CASC), 16 mi S La Union Narine, 2150 m, 

4.iii.1955, E. I. Schlinger & E. S. Ross (1♂, CASC). COSTA 

RICA. La Suiza de Turrialba, P. Schild (1♀, USNM), La Suiza, 

x.1926, P. Schild (1♀, USNM), Turrialba, 15–19.vii.1965, P. J. 

Spangler (1♀, USNM), Alajuela: 20 km S Upala, F. D. Parker, 

6.xii.1990 (1♀, EMUS), 24.i.1991 (1♀, EMUS), 27–31.iii.1991 

(1♀, EMUS), 1–10.iv.1991 (1♀, EMUS), Cartago: Tapanti Natl. 

Pk., Ranger Sta., 1200 m, pans in kitchen refuse, 7–11.x.1999, 

M. Buck (1♀, DEBU), Tapanti Natl. Pk., Arboles Caidos Trail, 

ca. 1300 m, 11.x.1999, S. A. Marshall (1♀, DEBU), R. Grande 

de Orosi, desde Puente R. Dos Amigos hasta la Represa, 1400–

1800 m, 1–11.ii.1996, D. Delgado (1♀, INBio), R. Grande de 

Orosi, desde Administracion hasta Sendero La Pava, 1150–1600 

m, viii.1996, R. Guzman (1♀, INBio), Res. Ftal. Rio Mache 

Alto El Roble, 2200 m, 10.x.1999, M. A. Zumbado, manual 

(1♀, INBio), Guanacaste: 3 km SE R. Naranjo, F. D. Parker, 

2–8.xi.1991 (1♀, EMUS), 12–17.xi.1991 (1♀, EMUS), 18–28.

xi.1991 (1♀, EMUS), 1–10.vii.1992 (1♀, EMUS), Puntarenas: 

Monteverde area, 6–14.vi.1973, 1400–1700 m, Erwin & Hevel 

(1♀, USNM), Monteverde Biol. Res., 1500 m, cloud forest, 

13.vi.2000, M. Buck (1♀, DEBU), Monteverde, B. V. Brown, 1–5.

vi.1988 (2♂ 2♀, EMUS), 28.v–1.vi.1988 (2♀, EMUS), Monteverde, 1500 m, 24–28.ii.1991, D. M. Wood (1♀, CNC), Monteverde, 1500–1800 m, 24–27.ii.1991, B. J. Sinclair, ex sweeping 

trails (1♀, CNC), Monteverde, Summit Cerro Chomogo, MT, 
2 2 • smithsonian contributions to zoology

1–10.iii.1992, 1800 m, D. M. Wood (2♀, CNC), Est. La Casona, Res. Biol. Monteverde, 1520 m, xii.1990, N. Obando (1♂, 

INBio), Est. La Casona, Res. Biol. Monteverde, 1520 m, 14–21.

iv.1996, Trampa de Clanuro Seco, K. Martinez (1♀, INBio), 

Fila Pittier, ACLA, PILA, 2550 m, 3.vii.1995, A. M. Maroto 

(1♂, INBio), Cerro Amigos, Monteverde, 1840 m, 25.ix.1994, 

M. A. Zumbado (1♀, INBio), Coto Brus, Z. P. Las Tablas, Orilla 

R. Bellavista, 1er Campto, C. Echandi, 1900 m, 16.iii.2000, 

M. Alfaro, manual (1♀, INBio), C. Chomogo, Monteverde, 

1800 m, 1.ix.1996, M. A. Zumbado (1♀, INBio), Est. La Casona, Res. Biol. Monteverde, 1520 m, N. Obando, xii.1990 (1♂, 

INBio), San Luis, 1040 m, R. B. Monteverde, x.1992, Z. Fuentes 

& A. Zumbado (1♂, INBio), San Luis, R. B. Monteverde, A. C. 

Arenal, 1000–1350 m, i.1994, Z. Fuentes (1♂, INBio), San José: 

San Carlos, Cerro Cura, 14 km SSW San Marcos de Tarrazu, 

1800 m, 9°36′31′′N, 84°07′09′′W, 15.ii.2003, S. A. Marshall (1♂, 

DEBU), Fca Zacatales, 2100 m, 8–10.viii.1995, M. A. Zumbado 

(1♂, INBio), Escazú, San Antonio, Quebrada Londres, 1700 m, 

17.iii.1997, E. Zumbado (1♀, INBio), San Gerrardo de Dota, 

2000–2500 m, Curso Tachinidae y Syrphidae, 22–26.ii.1992 

(1♀, INBio). DOMINICA. W.I., Clarke Hall, 11–20.ii.1965, 

W. W. Wirth (1♂, USNM), Pont Casse, 23.xi.1964, P. J. Spangler (1♀, USNM). EL SALVADOR. Monte Cristo, 28.vii.1977, 

Barger (1♂ 1♀, USNM). GUATEMALA. Petén: Ruinas Tikal, 

245 m, 7–10.vii.1977, E. M. & J. L. Fisher (1♀, CASC), Tikal, 

13.ii.1979, G. E. Bohart (1♀, EMUS). MEXICO. Chiapas: 

Município de la Trinitaria, Lagunas de Monte Belle National 

Park, Cinco Lagos, 1524 m, 5.x.1981, D. E. & P. M. Breedlove 

(1♀, CASC), Município Union, Canon Juarez SW slope of Volcán Takaná near Talquían, 2134 m, 14.xii.1976, D. E. & J. A. 

Breedlove (1♂, CASC), Município las Margaritas, 48 km NE 

Las Margaritas on Rd. to Campo Alegré, 2134 m, 25.x.1976, 

D. E. & J. A. Breedlove (1♀, CASC), 7200 ft, S. Crist. Las Casas, 

W. R. M. Mason, 8.vi.1969 (1♀, CNC), 29.v.1969 (2♀, CNC), 

Oaxaca: just W of Chiapas border, 19 km W of Rizo de Oro 

ridge S Cerro Baul, 1615 m, 28.iv.1972, D. E. Breedlove (1♂ 2♀, 

CASC), above Valle Nacìonal, 14.v.1963 (1♂, USNM), Veracruz: Lake Catemaco, 18.vi.1969, B. V. Peterson (1♂, CNC), San 

Rafael, Townsend (1♂, USNM), Chiapas: 6 km N San Cristobal, 

3.viii.1990, R. W. Jones (1♀, TAMU), nr. Pueblo, 24.xii.1962 

(1♀, USNM). PANAMA. Chiriqui, 15 km NW Mato del Volcan, 1200 m, 24–31.v.1977, Peck & Howden (1♀, CNC), R. P.: 

Panama Pr., Cerro Campana, 19.viii.1978, N. E. Woodley collection (1♂, USNM).

Comments. Neotanypeza dallasi is found throughout the Neotropics and is one of the few Neotanypeza known 

from the Caribbean. It is similar in morphology and coloration 

to N. claripennis (see comments for N. claripennis), which has 

a similar distribution but is also found in Ecuador, Peru, and 

Venezuela.

Tanypeza dallasi was described from eight males from two 

locations in Tucumán, Argentina: Quebrada de Lules and Tafi 

Veijo. A lectotype is here designated from one of the three USNM 

males that can be reliably placed in this type series.

Neotanypeza dimorpha (Hennig), 

comb. nov.

FIGURES 30, 31, 61–63

Scipopeza dimorpha Hennig, 1936:37. Hennig, 1936:37; Steyskal, 1967:2.

Male (Figure 31). Body length 9.5–10.3 mm. 

Anterolateral reflective triangle extending to midpoint of frons. 

Ocellar disc fully tomentose, relatively small, and not sharply 

delimited. Fronto-orbitals absent. Postvertical bristle minute. 

Postpronotal and presutural intra-alar bristles absent. One dorsocentral bristle. Anepisternum with 1 dark, dominant bristle. 

Calypter hairs yellow. Wing with broad, oblique medial stripe 

that is paler between veins and darkest to costa. Antenna brownish with first flagellomere darker and relatively thick. Clypeus 

and palpus dark brown. Legs yellow with mid and hind coxae 

brown, fore femur dark on distal 1⁄3, mid femur with dark subapical band, fore tibia brown on basal half, and mid and hind 

tibiae brown; tarsi dark brown with base of fore basotarsomere 

yellow and hind basotarsomere light yellow with apex dark. Abdominal bristles black with lateral bristles on tergites 1–3 and 

bristles on sternites yellow; abdomen dark brown.

Male Genitalia (Figures 61–63). As described for N. claripennis except as follows: external components larger and broader; sternite 6 shifted to left and with right 

margin atrophied; sternite 7 well developed and with single 

bristle; surstylus nearly as wide as epandrium, posterior margin 

bare and with minute bumps; hypandrium bare on left side of 

dissected male; postgonite relatively stout and short, brown (not 

yellow to clear), and with numerous apical setulae; distiphallus 

nearly as long as epandrium + surstylus, slightly widened apically with subapical spinules, and with long basal stalk. 

Female (Figure 30). Differs from male as follows: 

body length 10.5 mm; wing dark on distal 1⁄4 with strong spot 

around apex of R2+3; hind coxa light brown; mid femur yellow; 

fore basotarosmere light yellow with apex dark. Female collected by OL also with wing only lightly infuscated around veins 

on distal 1⁄4 (darker apex R2+3) and with faint anterodistal spot 

on mid femur.

Distribution. Ecuador, Peru.

Holotype. PERU. Pichis-Weg, 3.i.1904 (1♂, SMT).

Additional Material Examined. ECUADOR. Napo: SierrAzul Res., 14 km W Cosanga, 0°41′S, 

77°56′W, 10.v.2002, S. A. Marshall (1♂, DEBU; 1♀, QCAZ), 

9.v.2002, O. Lonsdale (1♀, QCAZ), El Chaco, 4.8 km W, 

1750 m, 7.xi.1999, S. A. Marshall (1♂, DEBU), Baeza, 2000 m, 

1–10.iii.1979, S. A. Marshall (1♂, DEBU), 5 km N El Chaco, 

ii.1983, 2000 m, M. Sharkey, Malaise trap (1♀, CNC), 7 km S 

Baeza, 28.iii.1983, G. & M. Wood (1♂, CNC).

Comments. Neotanypeza dimorpha is unique in the 

genus in having a light yellow hind basotarsomere, a dark hind 

coxa, and an oblique medial band on the male wing. Dark subgenal bristles, a dark antenna, and absence of the fronto-orbitals 
number 641 • 2 3

are also diagnostic, but these characteristics are shared with the 

equally distinct N. grandis. Although the newly described female 

of this species does not share the distinct wing pattern characteristic of the male (thereby staying true to Hennig’s intent for 

the specific epithet), it is here included in this species because it 

agrees in chaetotaxy, the tomentose pattern on the head and thorax, and the color of the hind tarsi. The single female paratype 

examined for this species is now treated as the paratype of the 

new species N. micans.

The male genitalia are easily diagnosed. Sternite 6 is almost 

entirely membranous on the right side and heavily skewed to 

the left, where there is a strong sclerotized band, and there are 

numerous minute bumps instead of stout pointed setulae along 

the posterior margin of the surstylus. The postgonite is also stout 

and brown (not yellow), and the distiphallus is long and only 

minutely textured apically.

Neotanypeza elegans (Wiedemann)

FIGURES 47–50, 87

Tanypeza elegans Wiedemann, 1830:528. Schiner, 1868:247; Bigot, 1886: 

380.

Tanypeza (Neotanypeza) elegans.—Hendel, 1903:203; Enderlein, 1913:225.

Neotanypeza elegans.—Hennig, 1936:35; Enderlein, 1936:42; Steyskal, 

1967:1, 1987:774; Griffiths, 1972:133. 

Neotanypeza flavitibia Hennig, 1936:34. Syn. nov.

Polphopeza flavitibia.—Enderlein, 1936:40. 

Neotanypeza (Polphopeza) flavitibia.—Steyskal, 1967:2.

Male. Body length 7.0–9.0 mm. Dorsal portion of reflective anterolateral triangle on frons as long as wide (or less) at 

widest point. Ocellar disc shining with central ovate tomentose 

spot (longer than wide); Peruvian specimen, Ecuadorian specimens, and most Erwin material with additional small, narrow 

tomentose stripe on lateral edge of disc. Anterior fronto-orbital 

short. Postvertical bristle minute. First flagellomere narrow and 

light yellow. Clypeus and palpus dark brown. Postpronotal and 

presutural intra-alar bristles present. Two dorsocentral bristles. 

Anepisternum with 1 dark, dominant bristle and several subdominant bristles. Calypter hairs yellow to golden brown, sometimes 

brown (Bolivia, Brazil, Ecuador, Peru). Wing clear. Legs yellow 

with mid coxa brown, fore femur with dark inner distal and 

light outer distal spots (sometimes forming nearly complete band 

in Ecuadorian and Brazilian specimens), fore tibia light yellow, 

and mid and hind tibiae with broad basal and short distal (interrupted anteriorly and posteriorly) bands; northern specimens 

with mid and hind tibiae dark brown with yellowish subapical 

band; tarsi dark brown with base of first segment sometimes 

lighter. Abdominal bristles dark brown; abdomen dark brown.

Male Genitalia. As described for N. claripennis.

Female. As described for male except as follows: calypter hairs always yellow to golden, never brown; first flagellomere sometimes thicker, with color sometimes orange, with apex 

(or more) usually infuscated; mid and hind tibiae yellow on distal half with 1 pair of opposing subapical spots that sometimes 

form distinct bands; distal 1⁄3 of wing lightly clouded (excluding 

pale spot near margin at apex of second radial cell) with slightly 

darker spot around R2+3 (paler in flavitibia holotype).

Distribution. Argentina, Bolivia, Brazil, Ecuador, 

Peru.

Lectotype [elegans]. BRAZIL. “Brasilia, Freyreiss” 

(1♂, ZMHU).

Holotype [flavitibia]. PERU. Rosalina, ix.1903, 

Urubambatl (1♀, SMT).

Paratype Examined [flavitibia]. PERU. Same 

data as holotype (1♂, SMT).

Additional Material Examined. ARGENTINA. Misiones 5 km E Puerto Iguazo behind Hotel Orquidaes, 1–6.ii.1992, S. A. Marshall (3♂ 2♀, DEBU). BOLIVIA. 

Mapiri, Sarampioni, 700 m, 23.i.1903 (1♂, SMT), 4.iii.1903 

(1♂, SMT), 6.iii.1903 (1♀, SMT), 11.iii.1903 (1♂, SMT), Mapiri, S. Carlos, 800 m, iv.1903 (2♂, SMT), La Paz: Guanay (8 

km W; 15°30.3′S, 67°50.8′W, 510 m), 13.iii.2001, A. Freidberg 

(1♂, USNM), Mapiri (5 km W; 15°17.8′S, 68°15.5′W; 750 m), 

16.iii.2001, W. N. Mathis (1♂, USNM), 8 km S Chulumani, 

nr. Apa Apa, 1700–1800 m, 16°22′S, 67°30.4′W, 9.iii.2001, 

S. D. Gaimari (1♀, USNM), 1 km E Guanay, 580 m, 15°30.2′S, 

69°52.3′W, 14.iii.2001, S. D. Gaimari (2♀, USNM), 5 km W 

Mapiri, Arroyo Tuhiri, 750 m, 15°17.8′S, 68°15.6′W, S. D. 

Gaimari, 16.iii.2001 (2♂, USNM), 18.iii.2001 (1♂, USNM), Cochabamba Prov.: 20 km SW Villa Tunari, Rio Avispha, 460 m, 

17°01.50′S, 65°31.67′W, 5.ix.2000, S. D. Gaimari (1♀, USNM), 

Ichilo Prov.: Santa Cruz Dept., Hotel Flora y Fauna, 4–6 km 

SSE Suena Vista, 17°29.95′S, 63°33.15′W, 400–500 m, N. E. 

Woodley, 9.xi.2003 (1♂, USNM), 14.xi.2003 (1♂ 1♀, USNM), 

Sta. Cruz, Buena Vista, 20.ii.1999, pan tr., F. D. Parker (1♀, 

EMUS). BRAZIL. Nova Teutonia, 27°11′S, 52°23′W, 300–500 

m, F. Plaumann, 4.ii.1968 (1♀, CNC), 24.xi.1969 (1♂, CNC), 

25.xi.1969 (1♂ 1♀, CNC), 7.xii.1969 (1♂, CNC), Rondonia: 

62 km S Ariquemes, W. J. Hanson, 13–25.iv.1992 (1♀, EMUS), 

8–20.xi.1994 (1♀, EMUS), Sao Paulo, Caraguatatuba, 1–5.

xi.1970, J. W. Boyes (1♂, CNC), “Tinguá, R. Janeiro”, ix.1950, 

Servico Febre, Amarela, M. E. S. Bras. (1♂, USNM), PR, Teno 

Boa, 13.vii.1983, J. A. Rafael (1♀, INPA), Est. S. Paulo, Juquia, 

1–4.xi.1935 [?], J. Lane (1♂, USNM), Cochabamba: Villa Tunari, “Orquidario”, 16°59.64′S, 65°26.10′W, 325 m, 6.ix.2000, 

ex: on vegetation, S. D. Gaimari (1♂, USNM), Rondonia: 62 

km SE Ariquemes, W. J. Hanson, 8–20.xi.1994 (1♂, EMUS) , 

180 m, 17–24.iii.1989 (1♂, EMUS). ECUADOR. Misahualli 

nr. Tena, 6–19.x.2001, C. Brammer (1♂, EMUS), Misahualli, 

nr. Tena, 26.viii–2.ix.2000, S. & P. Keller (1♂, EMUS), Yasuni 

Res. Sta., C. Brammer, 250 m, M.T., 0°40.566′S, 76°23.851′W, 

4–9.v.2003 (1♀, EMUS), 30.ix–1.x.2002 (1♀, EMUS), Napo: 

Res. Ethnica Waorani, 1 km S Onkone Gare Camp, Trans. Ent., 

2.x.1996, 220 m, 00°39′10′′S, 76°26′W, T. L. Erwin et al., insecticidal fogging of mostly bare green leaves, some with covering 

of lichenous or bryophytic plants in terre firme forest, Project 
2 4 • smithsonian contributions to zoology

MAXUS, At x-trans 10, 12 m, lot 1704 (1♀, USNM), 2.vii.1995, 

at trans 7, sta. 5, lot 1066 (1♀, USNM), Trans. Ent., 23.i.1994, 

220 m, 00°38′S, 76°36′W, T. L. Erwin et al., insecticidal fogging 

of mostly bare green leaves, some with covering of lichenous 

or bryophytic plants, Project MAXUS, At x-trans 10, 12 m, lot 

631 (1♂, USNM), Res. Ethnica Waorani, 1 km S Onkone Gare 

Camp, 29.vi.1994, 00°39′10′′S, 76°26′W, insecticidal fogging of 

mostly bare green leaves, some with covering of lichenous or 

bryophytic plants in terre firme forest, at 10 x-trans, 58 m mark, 

lot 746 (1♂, USNM), 20.vi.1996, at trans 2, Sta. 5, lot 1535 

(1♂, USNM), 21.vi.1994, at 9 x-trans, 89 m mark, lot 718 (1♂, 

USNM), 8.ii.1996, at trans. 8, sta. 10, lot 1480 (1♂, USNM). 

PERU. Quincemil, Cuzco, 27–31.viii.1962, 740 m, L. Pena (1♂, 

CNC), Monson Valley, Tingo Maria, 26.x.1954 (1♂, CASC), 

Quincemil, Cuzco, L. Pena, 780 m, 13–31.viii.1962 (1♀, 

CNC), 700 m, 1–15.xi.1962 (3♀, CNC), Pucallpa, 3.xii.1987, 

J. Schuuke (2♀, CNC), Pucallpa, i–ii.1951 (2♂, USNM), Monzon Valley, Tingo Maria, E. I. Schlinger & E. S. Ross, 23.ix.1954 

(1♂ 2♀, CASC), 10.x.1954 (2♀, CASC), 15.x.1954 (1♀, CASC), 

19.x.1954 (2♀, CASC), 26.x.1954 (1♂, CASC), 11.xii.1954 

(1♀, CASC), 23.xii.1954 (1♂, CASC), 15 mi NE Tingo Maria, 

E. I. Schlinger & E. S. Ross, 23.ix.1954 (1♀, CASC), Meshagua, 

Urubambati, 22.ix.1903 (1♂, SMT), 30.ix.1903 (1♂, SMT), 

1.x.1903 (1♀, SMT), 7.x.1903 (1♂ 1♀, SMT), 11.x.1903 (1♀, 

SMT), Ucayalifl, Unini, 21.x.1903 (1♀, SMT), Laristhal, 800 m, 

15.viii.1903 (1♀, SMT), Rosalina, 3.ix.1903 (1♂, SMT). Country unknown. “R B” (1♂, USNM).

Comments. Neotanypeza flavitibia is here included 

as a junior synonym of N. elegans because the genitalia, coloration, and pattern of tomentosity on the head of the two species 

are almost identical and vary similarly across the range of both 

species. The material collected by T. Erwin provide the most convincing support for synonymy, with sympatric representatives of 

both taxa showing the exact same regionally specific pattern of 

tomentosity on the ocellar disc. The primary difference between 

the two “species” is the color of the hairs on the calypter, which 

is brown (not yellow to golden brown) in the male lectotype 

of N. elegans; this color state is here considered an uncommon 

male-restricted phenotype found in Ecuador, Peru, Bolivia, and 

southern Brazil. 

Neotanypeza elegans was described from an unstated number of specimens from Brazil in the Senckenbergische Museum, 

Frankfurt. Enderlein (1936) noted that there was one male and 

one female in Berlin from the Loew collection, collected in Brazil 

on the Frey Expedition (“Freyreiss”), and marked these as types; 

the examined male is here designated the lectotype. There are no 

syntypes in Vienna, where Wiedemann’s collection was deposited. 

Neotanypeza flavibasis (Enderlein)

Tritanypeza flavibasis Enderlein, 1936:45.

Tritanypeza rufiventris Enderlein, 1936:46. Syn. nov.

Neotanypeza (Tritanypeza) flavibasis.—Steyskal, 1967:2.

Neotanypeza (Tritanypeza) rufiventris.—Steyskal, 1967:2.

Male. Body length 6.4–7.1 mm. Anterior reflective 

triangle on frons nearly extending to level of anterior frontoorbital. Ocellar disc fully tomentose. Anterior fronto-orbital 

thin and much smaller than posterior fronto-orbital. Postvertical bristle longer than tubercle and very thin. First flagellomere 

light yellow and narrow. Clypeus dark brown and palpus yellow. 

Postpronotal and presutural intra-alar bristles present. Three 

dorsocentral bristles. Anepisternum with 2 dark, dominant bristles. Calypter hairs pale. Wing clear. Legs yellow with mid coxa 

brown and tibiae light yellow with basal half of mid and hind 

tibiae brown; tarsi dark brown with basal half of hind basotarsomere, basal 2⁄3 of mid basotarsomere, and basal 4/5 of fore 

basotarsomere light yellow. Abdominal bristles dark brown with 

lateral bristles on tergite 1 yellow. Abdomen mostly yellow anteriorly with tergite 1 and posterolateral spots on tergites 3 and 4 

brownish; tergite 5 brownish posteriorly or entirely brown; abdomen past segment 5 dark brown, excluding yellow sternite 8.

Male Genitalia. As described for N. claripennis.

Female. As described for male except as follows: 

body length 6.8–8.4 mm; anterolateral triangle on frons not 

extending much past ptilinal suture; first flagellomere broad 

and orange, becoming lightly infuscated apically; tergites 1–4 

yellow and tergites 1–3 with narrow medial stripe (widest on 

tergite 1). Females from Brazil further differ as follows: abdomen dark past anterior 1⁄3 of tergite 3, anterior half of tergite 

2 (Sao Paulo), or entirely brown with anterior half of tergite 

2 orange laterally (southern Brazil and Argentina); wing very 

lightly clouded apically; mid tibia lighter and hind tibia with 

faint distolateral spots. 

Distribution. Argentina, Brazil, Trinidad, and Tobago, Venezuela.

Lectotype [flavibasis]. BRAZIL. “v. Olfers” (1♀, 

ZMHU).

Paralectotype [flavibasis]. BRAZIL. Same data 

as lectotype (1♀, ZMHU).

Lectotype [rufiventris]. VENEZUELA. “Bolivia, Caracas” (1♂, ZMHU).

Additional Material Examined. ARGENTINA. Misiones, 5 km E Puerto Iguazo behind Hotel Orquidaes, 1–6.ii.1992, S. A. Marshall (1♀, DEBU). BRAZIL. Nova 

Friburgo, Sitio Edelweiss, 1100 m, Malaise trap, disturbed 

for., 26.i.1990, S. A. Marshall (1♀, DEBU), PR, Terra Boa, 

27.xii.1983, J. A. Rafael (1♀, INPA), Rio de Janeiro: x.1937–

i.1938, R. C. Shannon (2♀, USNM), Mangaritiba, viii.1938–

ii.1939, R. C. Shannon (1♀, USNM), Rio de Janeiro, i.1939, 

R. C. Shannon (1♀, USNM), Rio de Janeiro, Dist. Federal Brasil, 

Servico Febre, Amarela, M. E. S. Bras., x.1937 (2♀, USNM), 

iv.1938 (2♀, USNM), vi.1938 (8♀, USNM), viii.1938 (2♀, 

USNM), v.1938 (2♀, USNM), ix.1938 (2♀, USNM), S. Paulo, 

[illegible], J. Lane (2♀, USNM), Nova Teutonia, S.C., v.1961, 

F. Plaumann (1♀, CNC), Nova Teutonia, 27°11′B, 52°23′L, 

F. Plaumann (1♀, CNC). TOBAGO. St. Paul: Roxborough (6 

km NNW, 11°16′N, 60°35.4′W), 20.iv.1994, W. N. Mathis 

(1♂, USNM). TRINIDAD. “Trinidad I.”, ASA Wright N.C., 
number 641 • 2 5

29.iii.1980, A. Graves (1♀, EMUS), Simla Res. Sta., 2–15.

vi.1981, Hanson, Clemens (4♀, EMUS).

Comments. Tritanypeza rufiventris is here included 

as a junior synonym of Neotanypeza flavibasis. The availability 

of new material has revealed a gradient of color intermediate between two extremes that are typified by the two lectotype specimens. Paler specimens appear to the north, where the Venezuelan 

type of T. rufiventris was collected, and darker specimens appear to the south in Brazil, where the type of N. flavibasis was 

collected.

Neotanypeza flavicalx (Enderlein)

FIGURE 21

Tritanypeza flavicalx Enderlein, 1936:46.

Neotanypeza (Tritanypeza) flavicalx.—Steyskal, 1967:2.

Male. Body length 5.8 mm. Anterior tomentose triangle on frons not extending much past ptilinal suture. Ocellar disc entirely tomentose. Anterior fronto-orbital bristle half 

length of posterior bristle and very narrow. Postvertical narrow 

and as long as tubercle. First flagellomere relatively narrow and 

orange. Clypeus dark brown (sometimes yellowish in Trinidad) 

and palpus yellow. Postpronotal and presutural intra-alar bristles present. Three dorsocentral bristles. Anepisternum with only 

1 dark, dominant bristle. Calypter hairs pale. Wing clear. Legs 

yellow with mid coxa brown, fore femur with brown inner subapical spot, fore tibia light yellow, and mid tibia only slightly 

darker yellow near base; hind tibiae and mid tarsi missing; fore 

tarsi dark brown with basal 2⁄3 of basal fore tarsomere light yellow. Abdominal bristles and abdomen entirely dark brown. 

Male Genitalia. Single known male (Panama) not 

dissected.

Female (Figure 21). As described for male except 

as follows: body length 5.9–7.7 mm; ocellar disc tomentose with 

shining anterolateral spots; postvertical bristle longer than tubercle and very thin; mid tibia with small basal band; hind tibia 

light yellow with basal half brown; tarsi dark brown with basal 

2⁄3 of fore basotarsomere light yellow and base of mid and hind 

basotarsomeres yellow; abdominal bristles dark brown with lateral bristles on tergite 1 yellow to brown.

Variation. Several females from Brazil (EMUS, 

INPA) with reduced tomentosity on ocellar disc, shining either 

along anterior 1⁄3 or with tomentose region reduced to ovate medial spot. 

Distribution. Bolivia, Brazil, Colombia, Ecuador, 

Panama, St. Lucia, Trinidad, Venezuela.

Holotype. COLOMBIA. “Cordillere, terra caliente” (1♀, ZMHU).

Additional Material Examined. BOLIVIA. 

Ichilo Prov.: Santa Cruz Dept., Hotel Flora y Fauna, 4–6 km SSE 

Suena Vista, 17°29.95′S, 63°33.15′W, 400–500 m, 11.xi.2003, 

N. E. Woodley (1♀, USNM). BRAZIL. Rondonia: 62 km SE 

Ariquemes, 8–20.xi.1994, W. J. Hanson (1♀, EMUS), Roraima: Serra Grande, 11–20.x.1992, D. W. Davis (1♀, EMUS), 

São Paulo: São Jose dos Campos, 29.viii–5.ix.1999, E. R. DePaula (1♀, EMUS), Pará: Óbidos, Serra da Escama, 1°55′00′′S, 

55°00′24′′W, 33.viii.2001, J. A. Rafael & J. F. Vidal (1♀, INPA), 

Amazonas: Campus Univarsitario Manaus, “21–30/2x/1988”, 

Marcia Castilho, arm., Shannon, Isla Fezes (1♀, INPA), Pq. 

N. Jau, Ig. Miratucu/Ig. Do Gerlei, 015700S-614900W, 23–28.

vii.1995, Arm. Malaise, J. A. Rafael & J. Vidal (1♀, INPA), Rio 

de Janiero: x.1937–i.1938, R. C. Shannon (1♀, USNM). ECUADOR. Rio Palenque, 0°35′S, 79°22′W, 22–26.ii.1976, 150 m, 

G. E. Shewell (1♀, CNC). PANAMA. Tabernilla, Canal Zone, 

“July 16-07”, A. Busck (1♂, USNM). ST. LUCIA. Barre d’Isle, 

north side (09–07), 13.93416°N, 60.95863°W, 320 m, S. D. 

Gaimari, A. R. Cline & R. Winton, 18–20.v.2009, Sharkey malaise (1♀, CSCA). TRINIDAD. Simla Res. Sta., 2–15.vi.1981, 

Hanson, Clemens (1♀, EMUS), Verdant Vale, “Oct.28,30.’18.

A345”, Ulrich and Morrison (1♀, USNM), Morne Blue, 

6.vii.1969, H. A. Howden, 2700’[ft] (1♀, CNC). VENEZUELA. T.F.A. Basecamp, 0°51′N, 66°10′W, Cerro d.l. Neblina, 

140 m, 20–24.iii.1984, O. Flint & J. Louton, Malaise trap over 

small stream at east side of base camp (2♀, USNM), T. F. Amaz. 

Cerro de la Neblina Basecamp, 0°50′N, 66°9′44′′W, 140 m, 

4–12.ii.1984, D. Davis & T. McCabe (1♀, USNM), T. F. Amaz. 

Cerro de la Neblina Basecamp, 0°50′N, 66°10′W, 140 m, Malaise trap in rainforest, P. J. & P. M. Spangler, R. A. Faitoute, & 

W. E. Steiner, 10–20.ii.1985 (1♀, USNM), 21–28.ii.1985 (2♀, 

USNM), Valera, E. P. deBellard (1♀, USNM).

Comments. Although the characters used to distinguish Neotanypeza flavicalx from similar species (short reflective tomentose triangle on frons, dark brown abdominal bristles 

that are sometimes paler) are relatively consistent, the status of 

this species should remain tentative since it is unknown how 

significant these relatively weak characters are in the indication 

of specific status. Furthermore, the Brazilian females of N. flavicalx with reduced tomentosity on the ocellar disc reflect an 

intermediate morphology between N. callitarsis and N. alopecia, 

which only otherwise differ in the color of the palpus (here given 

relative import for species diagnosis) and the hind femur. Only 

a single male is known for this species, suggesting that the males 

may inhabit a microhabitat less commonly sampled, as canopy 

fogging may have revealed for N. alopecia.

Neotanypeza grandis (Enderlein), 

comb. nov.

FIGURES 5, 22, 28, 64–66

Scipopeza grandis Enderlein, 1913:228. Hennig, 1936:37; Enderlein, 1936: 

40; Steyskal, 1967:3.

Male (Figures 22, 28). Body length approximately 10.2–12.1 mm. Dorsal portion of anterior tomentose triangle 2 times longer than wide at widest point. Ocellar disc fully 
2 6 • smithsonian contributions to zoology

tomentose. Frons black, excluding reflective parts. Fronto-orbital 

bristles absent. Postvertical bristle minute. Subgenal bristles 

sometimes black, brown to yellow in nontypes. First flagellomere 

narrow; antenna black and face orange. Clypeus and palpus dark 

brown. Postpronotal and presutural intra-alar bristles absent 

(postpronotal present in one nontype). One dorsocentral bristle. 

Anepisternum with 1 dark, dominant bristle, sometimes with numerous subdominant bristles. Calypter hairs yellow. Wing dark in 

first radial cell, excluding base, and around distal 2⁄3 of R2+3; distal half of R4+5 surrounded by light infuscation. Legs yellow with 

mid coxa brown, distal 1⁄3 of fore and mid femora dark brown 

(reduced in nontypes), apex of hind femur brown, and tibiae and 

tarsi dark brown with distal half of fore tibia yellow; nontypes 

with yellowish subapical bands on mid and hind tibiae. Abdominal bristles dark brown; abdomen dark brown.

Male Genitalia (Figures 64–66). As described for N. claripennis except as follows: terminalia stouter, 

with surstylus broader and strongly curved on posteromedial 

margin; sternite 6 more extensively setulose, with weakly sclerotized regions on right distal and left basal margins and with reverse N-shaped sclerotized band on left side; sternite 7 relatively 

large and well developed; setulose portions of sternites 7 and 8 

heavily wrinkled; distiphallus not much longer than wide, without sclerotized medial section, and with sides strongly produced 

at midpoint.

Female (Figure 5). As described for male except 

as follows: body length 9.9–11.0 mm; postpronotal bristle and 

1 fronto-orbital bristle present; dorsocentral bristles closely 

spaced; subgenal bristles brown; first flagellomere more ovate 

with base orange.

Distribution. Ecuador, Venezuela.

Holotype. Ecuador. Baños, R. Haensch S. (1♂, 

PAN).

Additional Material Examined. ECUADOR. Napo: SierrAzul Lodge, 14 km W Cosanga, 2200 m, 

8–11.v.2002, S. M. Paiero (1♀, DEBU), 0°40′S, 77°56′W, S. A. 

Marshall (7♂ 2♀, DEBU; 1♂ 1♀, QCAZ), 0°40′55′′S, 77°56′W, 

O. Lonsdale (1♀, DEBU), 0°41′S, 77°56′W, 10.v.2002, S. A. 

Marshall (1♂ , DEBU; 1♂ 1♀, QCAZ), Reserva SierrAzul, 14 km 

W Cosanga, 2200 m, 5.xi.1999, S. A. Marshall (2♂, DEBU; 2♂

1♀, QCAZ). VENEZUELA. Lindig, 1864, “elegans, Alte sammlung” (2♂ 1♀, AMNH).

Comments. Neotanypeza grandis is easily diagnosed not only by its size but also by its dark antenna, face, 

femora, and wing, the latter of which is heavily infuscated along 

the anterior margin. The male genitalia are also characteristic in 

that the sixth sternite has a dark reversed N-shaped band and the 

epandrium and sternite 8 have a skin-like surface texture (sometimes found in other species with reduced chaetotaxy).

Neotanypeza leucothrix, sp. nov.

Male. Body length 8.5 mm. Dorsal portion of anterior tomentose triangle short, slightly longer than wide at widest 

point; frons shining anteromedially. Ocellar disc tomentose with 

posterior margin shining. Anterior fronto-orbital bristle absent. 

Postvertical bristle minute. First flagellomere relatively narrow 

and light yellow. Clypeus dark brown and palpus brown with orange tint. Postpronotal bristle present and presutural intra-alar 

bristle absent. One dorsocentral bristle. Anepisternum with 1 

dominant bristle. Calypter hairs yellow. Wing clouded on distal 

third with darker spot around R2+3. Legs yellow with mid coxa 

brown, fore femur with dark inner distal and light outer distal 

spots, mid and hind femora with apex brown, fore tibia light 

yellow, and mid and hind tibiae brown with faint yellow subapical bands (nearly indistinct on hind leg); tarsi dark brown with 

basotarsomeres lighter to base. Abdominal bristles dark brown; 

abdomen yellow with narrow medial stripe on tergite 2, tergite 

3 with thick medial stripe and brown posterior margin, tergites 

4 and 5 infuscated sublaterally and with brown posterior and 

anterior margins, tergite 4 with narrow central stripe, tergite 6 

brown excluding lateral margin, and epandrium and posterior 2⁄3

of surstylus brown with yellow mottling.

Male Genitalia. Externally as described for N. 

claripennis (sole male not dissected), except sternite 6 approximately 1⁄3 longer and sternite 7 well developed.

Female. Unknown.

Etymology. The specific epithet refers to the color 

of the hairs on the calypter.

Distribution. Peru.

Holotype. PERU. Quincemil, Cuzco, 1–15.

xi.1962, 700 m, L. Pena (1♂, CNC).

Comments. Of those Neotanypeza with a single 

dorsocentral bristle, N. leucothrix is unique in having a single 

fronto-orbital, a lightly clouded wing with a small subapical 

spot, a posteriorly shining ocellar disc, and a predominantly yellow abdomen.

Neotanypeza marshalli, sp. nov.

FIGURES 7, 8, 29, 67–69

Male (Figures 7, 8, 29). Body length 7.3–7.9 

mm. Dorsal portion of anterior tomentose triangle slightly longer than wide; frons (excluding reflective stripe) black, without 

purple tint. Ocellar disc fully tomentose. Anterior fronto-orbital 

bristle smaller than posterior fronto-orbital. Postvertical bristle 

minute. Clypeus and palpus dark brown. Antenna dark brown 

with base of first flagellomere (relatively narrow) orange. Postpronotal bristle present and presutural intra-alar bristle absent. 

Two dorsocentral bristles. Anepisternum with 2 to 3 dark, dominant bristles. Calypter hairs yellow. Wing clouded apically and in 

anterodistal half. Legs yellow with mid and hind coxae brown, 

fore femur dark brown on distal 3/5, mid femur dark on distal 2/5, 

hind femur dark on distal 1/5, and tibiae and tarsi dark brown 

with fore tibia yellow on inner distal face and yellowish on outer 

apical face. Abdominal bristles dark brown, sometimes with lateral bristles on tergites 1 and 2 yellow; abdomen dark brown.
number 641 • 2 7

Male Genitalia (Figures 67–69). As described 

for N. claripennis except as follows: sternite 6 symmetrical, without sublateral sclerotization, evenly setose and setulose, and with 

distal margin shallowly emarginate; surstylus only 3/5 height of 

epandrium, evenly curved on both margins, more heavily setose, 

with smaller setulose region and without stout conical setulae 

along posterior margin; epandrium and sternite 8 with shallow 

wrinkles; hypandrium with 2 pairs of bristles; postgonite with 

6 apical setulae; distiphallus long (nearly as long as epandrium 

+ surstylus), evenly dark and sclerotized, slightly curved in cross 

section, textured distolaterally, and with several medial fossae. 

Female. As described for male except as follows: 

body length 7.6–8.2 mm; first flagellomere more orange at base; 

only 1 dominant bristle on anepisternum; basal 4/5 of fore basotarsomere and distal 1⁄3 of fore tibia light yellow; distal half of 

fore femur and distal 1⁄3 of mid femur dark brown; abdominal 

bristles entirely dark brown. 

Etymology. The specific epithet honors Dr. Stephen Marshall, who not only collected much of the type material of this species (including the holotype) but also provided all 

photographs of live specimens used here (Figures 1–6).

Distribution. Ecuador, Venezuela.

Holotype. ECUADOR. Napo: SierAzul Res., 14 

km W Cosanga, 2200 m, 0°40′55′′S, 77°56′09′′W, 8–11.v.2002, 

S. A. Marshall (1♂, QCAZ).

Paratypes. ECUADOR. Same data as holotype (4♂

2♀, DEBU; 2♂ 1♀, QCAZ), 10.v.2002, M. Buck (1♂, DEBU), 

SierAzul Res., 14 km W Cosanga, 2200 m, 5.xi.1999, S. A. Marshall (1♂, DEBU), 7 km S Baeza, 28.iii.1983, G. & M. Wood (1♀, 

CNC). VENEZUELA. Aragua Colonia Tovar, 2300 m, sweep, 

steep forest trail, 10.iii.1995, S. A. Marshall (3♂ 1♀, DEBU), 

Mérida (city), Rio Albarregas, 2500 m, cloud for., Los Chorros, 

1.v.1988, S. A. Marshall (1♀, DEBU), Mérida, La Azulita, 20 km 

SE, ULA Biol. Res., La Carbonera, 2300 m, podocarp forest, Malaise, 28.vi–3.viii.1989, S. & J. Peck (1♀, DEBU), Mérida, Tabay, 

La Mucuy, Truchicola trail, 2300 m, cloud forest, FIT, 17.vi–3.

vii.1989, S. & J. Peck (1♀, DEBU), Lara, Prq. Nac. Yacambu, 

10 km S Sanare, 28.ii.1985, Kovarik & Jones (1♂, TAMU), 

Ara.: trail near Yacambu National Park, 5.ix.2008, 9°41′59′′N, 

69°38′52′′W, 1904 m, J. Skevington (1♀, CNC).

Comments. The long, entirely sclerotized phallus 

and dark color of Neotanypeza marshalli easily differentiates it 

from congeners. Other taxa that have the ends of the femora extensively pigmented, including N. dimorpha and N. grandis (both 

of which also have black antennae and tibiae), differ in having no 

postpronotal or fronto-orbital bristles (at least in the male) and 

have strikingly different male genitalia. Neotanypeza symmetros

also has predominantly brown mid and hind tibiae and similar 

genitalia, but the fore leg and antenna are predominantly yellow.

Neotanypeza micans, sp. nov.

Male. Body length 7.9 mm. Dorsal portion of anterior tomentose triangle short, as long as wide at widest point; 

frons shining anteromedially and medial furrow slightly raised 

and widened. Ocellar disc shining with wide, subquadrate, medial tomentose spot. One fronto-orbital bristle. Postvertical bristle minute. First flagellomere narrow and orange. Clypeus yellow 

and palpus light brown to yellowish. Postpronotal bristle present and presutural intra-alar bristle absent. Two dorsocentral 

bristles. Anepisternum with 2 or more dark, dominant bristles. 

Calypter hairs yellow. Wing clear with distal 1⁄3 lightly clouded 

around R2+3 and R4+5. Legs yellow with mid coxa brown, distal 1⁄3 of fore femur dark brown, apex of mid and hind femora 

dark brown, fore tibia light yellow and mid and hind tibiae dark 

brown with faint yellowish subapical band; tarsi dark brown 

with base of first segment yellowish. Abdominal bristles dark 

brown; abdomen light brown with tergites 1–3 yellow, excluding 

dark medial stripe.

Male Genitalia. As described for N. claripennis.

Female. As described for male except as follows: 

body length 10.5–13.7 mm; abdomen dark brown; wing lightly 

clouded on distal 1⁄3; first flagellomere slightly thicker.

Etymology. The specific epithet, which is Latin for 

“shiny,” refers to the bare shining patches on the frons and ocellar disc.

Distribution. Peru.

Holotype. PERU. Meshagua, Urubambafl, 

7.x.1903 (1♂, SMT).

Paratypes. PERU. Meshagua, Urubambafl, 

12.x.1903 (1♀, SMT) [paratype of N. dimorpha], Madre de 

Dios, Manu, Rio Manu, 250 m, Pakitza, 12°07′S, 70°58′W, 

9–23.ix.1988, A. Freidberg (1♀, USNM).

Comments. In addition to having two pairs of dorsocentral bristles, this newly described Peruvian species can be 

readily diagnosed by a shining (not velvety) anteromedial patch 

on the frons, pale mouthparts, and a wide (not narrow) tomentose spot on an otherwise shining ocellar disc. The externally 

similar Neotanypeza apicalis, which also has a pale male abdomen, a yellow palpus, and similar chaetotaxy, can be diagnosed 

by having yellow spots on the thorax (excluding the type), clearer 

wings (excluding the type), entirely yellow mid and hind femora, 

a yellow-haired male frons with a deep, broad central sulcus, and 

an entirely tomentose ocellar disc. 

Neotanypeza montana (Enderlein)

Polphopeza montana Enderlein, 1936:40. 

Neotanypeza (Polphopeza) montana.—Steyskal, 1967:2.

Neotanypeza elegans. Misidentification.—Steyskal, 1987:774.

Female: Lectotype. Body length nearly 9.0 mm. 

Dorsal portion of anterior tomentose triangle approximately 1.5 

times longer than wide; color white, but dorsal portion golden 

yellow if extending past anterior fronto-orbital. Ocellar disc 

fully tomentose. Anterior fronto-orbital bristle absent. Postvertical bristle minute. Clypeus dark brown; palpus yellowish. 

First flagellomere relatively narrow and very lightly infuscated 
2 8 • smithsonian contributions to zoology

apically. Postpronotal bristle present and presutural intra-alar 

bristle absent. Two dorsocentral bristles. Anepisternum with 1 

dark, dominant bristle. Calypter hairs yellow. Wing faintly infuscated to apex. Legs yellow with mid coxa brown, fore femur 

with faint inner distal spot, and mid and hind tibiae with broad 

basal and narrow distal brown bands; tarsi dark brown with 

fore basotarsomere yellow on basal half and mid and hind basotarsomeres yellowish to base. Abdominal bristles dark brown; 

abdomen dark brown.

Variation: Females. As described for lectotype 

except as follows: body length 6.6–9.0 mm; presutural intra-alar 

bristle present in females from Costa Rica, Panama, Brazil, and 

one specimen from Ecuador; anterior fronto-orbital bristle present in specimens outside of Ecuador; palpus dark brown; wing 

varying from clear (Ecuador) or lightly clouded apically (Brazil, 

Costa Rica, Ecuador) to darkly clouded on distal 1⁄3, becoming 

darker to costa (Bolivia, Venezuela); one female from Peru with 

distal half of mid and hind tibiae yellow; apex of mid and hind 

femora brown in females from Bolivia, Venezuela, and Peru; 

basal half of fore tibia brown in females from Bolivia and Venezuela; female from Ecuador with brown distal band on mid and 

hind tibiae; mid tarsi entirely dark in Central American females 

and mid and hind tarsi entirely brown in Bolivian and Venezuelan females; fore basotarsomere light yellow with apex brown 

in Ecuadorian, Bolivian, and Venezuelan females; abdominal 

bristles entirely dark in Bolivian and Brazilian females, yellow 

on tergite 1 in Colombian paralectotype and some females from 

Costa Rica and Panama, and yellow on tergites 1 and 2 in females from Venezuela and Ecuador. 

Male. As described for female lectotype except as follows: body length 8.2–8.5 mm; anterior tomentose triangle on 

frons variable in length (very short to extending to midpoint of 

frons). Ocellar disc tomentose with some Peruvian and Brazilian 

(CNC) males with anterior and posterior (Peru only) margins 

shining. First flagellomere narrow to broad, pale to infuscated 

(infuscation usually darker and more extensive in specimens 

south of Ecuador and Brazil); pedicel brown in Peruvian male. 

Palpus yellow in one Brazilian and one Colombian male that 

have faded or absent distal bands on mid and hind tibiae. Presutural intra-alar present in Brazilian males. Anepisternum with 

1 or 2 dark, dominant bristles. Wing clear to lightly clouded on 

distal 1⁄3. Legs yellow with mid coxa brown, fore femur with 

light outer distal spot and broad, dark inner distal spot (sometimes forming broad band), and mid and hind tibiae dark brown 

with yellowish subapical band (entirely yellow on distal half 

in specimens from Ecuador); basal half of fore tibia sometimes 

brown; tarsi dark brown with basal segment usually yellow to 

base (entirely to predominantly dark in Peruvian and Brazilian 

males and entirely light yellow with apex brown in some South 

American specimens, including most Ecuadorian material); apex 

of mid and hind femora dark in Peruvian male. Abdominal bristles dark brown with lateral hairs yellow on pregenitalic tergites 

of Ecuadorian males and lateral hairs of tergite 1 of Peruvian 

males. Mexican male with lateral bristles on abdomen yellow, 

tergite 3 with lateral margin broadly yellow, and golden tomentose triangle on frons reaching level of anterior fronto-orbital. 

Brazilian males with most putatively ancestral states, having 2 

fronto-orbitals, 1 presutural intra-alar, entirely brown abdominal bristles, a clear wing, yellow femora, and strongly banded 

tibiae. 

Male Genitalia, As described for N. claripennis

except as follows: medial sclerotization absent from distiphallus 

in males with predominantly dark abdominal bristles (that is, 

leaving only 1 pair of pointed lateral sclerites); sternite 6 approximately 50% longer in males with apex of femora and basal 

half of fore tibia brownish. Mexican male with lateral sclerotizations on distiphallus faint and with medial sclerotization narrow 

but pronounced.

Distribution. Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, Panama, Peru, Venezuela.

Lectotype. Colombia. “Cordillieren, Terra fria, 

Thieme S” (1♀, ZMHU).

Paralectotype. Colombia. Moritz (1♀, ZMHU).

Additional Material Examined. ARGENTINA. Salta, 30 km E Salta, Campo Quijano, forest vestige at 

El Alisal, S. A. Marshall, 20.ii.1992 (2♂ 1♀, DEBU), 18.ii.1992 

(2♂, DEBU), Salta, 10 km N La Caldera forest, 1500 m, Camino 

la Cornisa, 27.ii.1992, S. A. Marshall (1♂ 2♀, DEBU). BOLIVIA. 

La Paz: Apa Apa (8 km S Chulumani, 16°35.6′S, 68°51.2′W, 

1960 m), 9.iii.2001, A. Freidberg (2♀, USNM), Apa Apa, 8 km 

S Chulumani, 1950–2100 m, 16°22′S, 67°30.4′W, S. D. Gaimari, 

23.iii.2001 (2♀, USNM), 24.iii.2001 (3♀, USNM), La Paz, Caranavi, ca. 10 km NW, road to Entel Tower, 1400 m, 15°46′35′′S, 

67°35′48′′W, 13.iv.2001, S. A. Marshall (1♀, DEBU), dung pans 

(1♀, DEBU), La Paz, Chulumani, Apa Apa Reserve, 2000 m, 

16°21′15′′S, 67°30′21′′W, 1.iv.2001, S. A. Marshall (3♀, DEBU), 

Cochabamba, 109 km E, 1400 m, traps, mountain/cloud forest, FIT, 1–6.ii.1999, F. Génier (2♀, DEBU). BRAZIL. Rio de 

Janeiro: Dist. Federal, Serviço Febre Amarela, M. E. S. Bras., 

ix.1938 (3♀, USNM), x.1937 (1♀, USNM), Rio de Janeiro, 

R. C. Shannon (1♀, USNM), Rio de Janeiro, i.1939, R. C. Shannon (1♀, USNM), Sao Paulo: Caraguatatuba, 1–5.xi.1970, J. W. 

Boyes (4♂, CNC), Est. Biol Boraceia, Salesopolis, 14–19.x.1970 

(1♂, CNC). COLOMBIA. 3 mi W Villavicencie, Meta, 920 m, 

11.iii.1955, E. I. Schlinger & E. S. Ross (1♂, CASC). COSTA 

RICA. Alajuela: 20 km S Upala, F. D. Parker, 10–19.iii.1991 

(1♀, EMUS), 19.ii.1991 (1♀, EMUS), Limon: Est Hitoy Cerere, 

100 m, R. Cerere, Res. Biol. Hitoy Cerere, R. Guzman, “28–12 

abr 1992” (1♀, INBio), Puntarenas: 2000 m, 12.viii.1996, S. A. 

Marshall (1♀, DEBU), Las Alturas Bio. Stn., 8°57′N, 82°58′W, 

Cerro Chai, 2500 m, 15.viii.1995, S. A. Marshall (1♀, DEBU), 

Monteverde, sweep nr. Biol. Stn., 25.v.1998, S. A. Marshall (1♀, 

DEBU), San Jose: San Gerrardo de Dota, 9.33′N, 8.48′W, Bosque 

Nuboso, 2400–2600 m, 7.viii.1995, D. C. Caloren (1♀, DEBU). 

ECUADOR. Biblián (1♀, USNM), Napo, Baeza, 2000 m, 1–10.

iii.1979, S. A. Marshall (2♀, DEBU), SierrAzul Res., 14 km W 
number 641 • 2 9

Cosanga, 2200 m, 0°40′55′′S, 77°56′09′′W, 10.v.2002, M. Buck 

(2♀, DEBU), 8–11.v.2002, S. A. Marshall (1♀, DEBU), SierrAzul 

Lodge, 14 km W Cosanga, 2200 m, 0°40′55′′S, 77°56′09′′W, Malaise, 9–10.v.2002, Buck & Paiero (1♀, DEBU), Napo, El Chaco, 

2000 m, 15–23.ii.1983, Mal. Tp., L. Masner & M. Sharkey (1♀, 

DEBU), Napo, 5 km N El Chaco, ii.1983, 2000 m, M. Sharkey, Malaise trap (1♀, CNC), Tandapi, 40 km SW Quito, 1300–

1500 m, 15–21.vi.1965, Pena (5♂ 5♀, CNC), Tandapi, Pichincha, 

1300–1700 m, 21.vi.1965, L. Pena (5♂ 3♀, CNC), Rio Margajitas, Rio Pastaza, 1250 m, 20.iii.1939, F. M. & H. Brown (1♀, 

AMNH), Pichincha, Maquipucuna Res., main trail, 1400–1600 

m, 0°07′34′′N, 78°37′57′′W, 27.iv.2002, S. A. Marshall (2♂ 1♀, 

DEBU), Maquipucuna Res., 1200 m, 0°07′34′′N, 78°37′57′′W, 

28.iv.2002, M. Buck (1♂, DEBU), Pichincha, Nanagalito, 7 km 

SE, trout farm, 1500 m, 27.x.1999, S. A. Marshall (1♂, DEBU). 

MEXICO. Oaxaca: 15.1 mi N San Gabriel, Mixtepec, 3850 ft, 

Hwy 131, 11.vii.1987, R. Wharton (1♂, TAMU). PANAMA. 

Chir., Boquete Trail, 28.iii.1982, W. J. Hanson (1♀, EMUS).

PERU. Monson Valley, Tingo Maria, 10.x.1954, E. I. Schlinger 

and E. S. Ross (1♀, CASC), Pasco, 10°35′S, 75°35′W, 30–31.

xii.1972, 1700 m, S. Helava (2♂, CNC). VENEZUELA. Lindig, 1864, “claripennis, Alta Sammlung” (1♀, AMNH), Mérida, 

Mucuy Fish Hatchery, 7 km E Tabay, 6600 ft, 10–13.ii.1978, 

J. B. Heppner (2♀, USNM), Mérida, Mucui, 10 km E Tobay, 

2000 m, 28.iv.1981, H. Townes (4♀, CNC), Yacambo, 1200 m, 

10.v.1981, H. K. Townes (1♀, CNC), Merida (city), Rio Albarregas, 2200 m, cloud for., 24.iv.1988, S. A. Marshall (1♀, DEBU), 

Lara, Yacambu N.P., 17 km E Sinaré, 9°42′N, 69°34′W, 1650 m, 

v.1998, J. Ashe, R. Brooks, R. Hanley, FIT (1♀, CNC).

Comments. As is evident from the above description, Neotanypeza montana as currently defined is a highly 

variable and widespread species that likely contains at least one 

cryptic taxon (hence description of the name-bearing female lectotype first). The reason for not dividing this species into a number of discrete taxa is that there are only a few characters that 

vary among morphotypes, and these vary in relatively nondiscrete ways. Although a number of differences could be compiled 

for any two of the above specimens (for example, when comparing the females from Ecuador and the female from Peru, which 

are highly dissimilar) that would make them appear to belong 

to separate species, an additional series from other neotropical locations would reveal a spectrum of intergradations that 

blur the boundaries of these potential “species.” Some of these 

characters even vary between otherwise identical specimens collected at the same location; in Ecuador, for example, one female 

has a presutural intra-alar bristle (absent in other Ecuadorian 

material), and several other uncharacteristic specimens have a 

dark wing stripe and a basally brown fore tibia. Furthermore, 

although the materials from Venezuela and Bolivia are nearly 

identical in morphology, the lateral hairs on the first tergite (a 

character otherwise useful in specific circumscription) are yellow in the former and brown in the latter. As also seen for N. 

dallasi and N. claripennis, additional data will be necessary to 

determine what components of this variation are indicative of 

actual interspecific boundaries. 

Neotanypeza nigrithrix, sp. nov.

Male. Body length 7.3 mm. Dorsal portion of anterolateral tomentose triangle short, slightly longer than wide at widest point. Ocellar disc tomentose with anterior margin around 

tubercle shining. Fronto-orbitals absent. Postvertical bristle minute. Clypeus and palpus dark brown. First flagellomere relatively 

narrow and brown with base orange. Postpronotal bristle present and presutural intra-alar bristle absent. Two dorsocentral 

bristles. Anepisternum with several dark, dominant bristles. Calypter hairs brown. Wing clear with anterodistal margin faintly 

infuscated. Legs yellow with mid coxa brown, fore femur with 

dark inner distal and light outer distal spots, mid and hind femora with apices brown, fore tibia light yellow, and mid and hind 

tibiae brown with faint yellow subapical bands (nearly indistinct 

on hind leg); tarsi dark brown. Abdominal bristles dark brown; 

abdomen dark brown.

Male Genitalia. Not dissected; hypandrium, 

phallapodeme, and sternite 6 not visible. As described for N. 

claripennis, except sternite 7 well developed, surstylus more extensively setose, and distiphallus emarginate apically and without medial sclerotization (like that of N. plotoplax).

Female. Unknown.

Etymology. The specific epithet refers to the color 

of the hairs on the calypter.

Distribution. Peru.

Holotype. PERU. Pasco, 10°35′S, 75°35′W, 30–31.

xii.1972, 1700 m, S. Helava (1♂, CNC).

Comments. Neotanypeza nigrithrix resembles the 

Peruvian N. plotoplax in having no fronto-orbitals, dark calypter hairs, and a similar distiphallus, but it is smaller and less 

robust, there are two pairs of dorsocentrals (not one), and the 

wing, hind coxa, antenna, and legs are much darker.

Neotanypeza ochrifemur (Enderlein)

Tritanypeza ochrifemur Enderlein, 1936:45.

Neotanypeza (Tritanypeza) ochrifemur.—Steyskal, 1967:2.

Male. Body length 5.7–7.0 mm. Anterior tomentose 

triangle on frons reaching past level of anterior fronto-orbital. 

Ocellar disc tomentose with shining anterolateral spots. Anterior fronto-orbital bristle narrow and half the length of posterior 

bristle, sometimes absent. Postvertical bristle longer than tubercle and very narrow. First flagellomere pale yellow and with 

length slightly more than twice width. Clypeus dark brown and 

palpus yellow. Postpronotal and presutural intra-alar bristles 

present. Three dorsocentral bristles. Anepisternum with only 1 

dark, dominant bristle. Calypter hairs pale. Apex of wing dusky. 

Legs yellow with mid coxa brown, fore tibia light yellow, mid 
3 0 • smithsonian contributions to zoology

tibia grayish to base, and hind tibia with dark basal and light distal bands; tarsi dark brown with base of basotarsomeres yellow 

(paler on fore leg). Abdominal bristles dark brown with lateral 

bristles on tergite 1 yellow; abdomen dark brown.

Variation. One Argentinian male differs as follows: 

first flagellomere darker; fore femur with long inner distal spot; 

mid tibia brownish on basal third; hind tibia with 1 pair of dark 

distolateral spots. Other Argentinian male with brownish basal 

band on hind tibia. Costa Rican male differs as follows: reflective stripe on frons extending to level of posterior fronto-orbital; 

basal half of fore basotarsomere as pale as tibia; mid and hind 

tibiae with dark basal band and dark distolateral spots. 

Male Genitalia. As described for N. claripennis. 

Female. Unknown.

Distribution. Paraguay, Guatemala, Ecuador.

Lectotype. Paraguay. San Bernardino, II [11 December in original description], K. Fiebrig, S.V. (1♂, ZMHU).

Additional Material Examined. ARGENTINA. Misiones, 5 km E Puertos Iguazo behind Hotel Orquidaes, 

1–6.ii.1992, S. A. Marshall (2♂, DEBU). BRAZIL. Roraima: Rio 

Uraricoera, Ilha de Maraca, 21–30.x.1987, J. A. Rafael e equipe, 

Armadilha de Malaise (1♂, INPA). COSTA RICA. Heredia: 

Santo Domingo, INBio Pk., 19–21.viii.2001, S. A. Marshall (1♂, 

DEBU). PANAMA. Canal Zone, Chiva-Chiva Rd., 8 air km N 

Ft. Clayton, 23.vii.1978, N. E. Woodley (1♂, USNM).

Comments. See comments for Neotanypeza ornatipes.

Neotanypeza ornatipes (Bigot)

FIGURES 16, 17

Taeniaptera ornatipes Bigot, 1886:380. 

Taeniaptera pallidipennis Bigot, 1886:380. Syn. Steyskal (1967) [not 

explicit].

Tanypeza mexicana Giglio-Tos, 1893:14. Giglio-Tos, 1895:61; Wulp, 1897: 

363. Syn. Hennig (1936).

Tanypeza ornatipes.—Wulp, 1897:363.

Tanypeza pallidipennis.—Wulp, 1897:363. 

Tanypeza (Neotanypeza) ornatipes.—Enderlein, 1913:225.

Tanypeza (Neotanypeza) pallidipennis.—Enderlein, 1913:225.

Tanypeza (Neotanypeza) mexicana.—Enderlein, 1913:225.

Tanypeza (Neotanypeza) flavohirta Enderlein, 1913:226. Syn. nov.

Neotanypeza ornatipes.—Hennig, 1936:34; Enderlein, 1936:42; Steyskal, 

1967:1.

Neotanypeza flavohirta.— Hennig, 1936:37. 

Tritanypeza flavohirta.—Enderlein, 1936:43.

Neotanypeza (Tritanypeza) flavohirta.—Steyskal, 1967:2.

Male (Figure 16). Body length 6.1–7.9 mm. Anterior tomentose triangle on frons yellowish to tan colored and 

nearly reaching level of anterior ocellus. Ocellar disc fully tomentose. Anterior fronto-orbital bristle thin and 1⁄2–2⁄3 length posterior 

bristle. Postvertical bristle slightly longer than tubercle and very 

thin. First flagellomere pale yellow with length slightly more than 

twice width. Clypeus dark brown and palpus yellow. Postpronotal 

and presutural intra-alar bristles present. Three dorsocentral bristles. Anepisternum with only 1 dark, dominant bristle. Notum, 

excluding anterior margin, with relatively thick yellow pruinosity. Calypter hairs yellow. Wing clear. Legs yellow with hind coxa 

brown, fore tibia light yellow, mid and hind tibiae with dark basal 

and apical bands; fore femur sometimes with faint inner distal 

spot; tarsi dark brown with basal half of fore basotarsomere yellow. Abdominal bristles yellow, becoming brown medially.

Variation. The questionably identified male from 

Costa Rica differs from the N. ornatipes holotype in having the 

anterior dorsocentral reduced and hairlike (possibly as a developmental abnormality) and in having the mid tibia entirely yellow. Erwin material with mid tibia light yellow on distal half and 

fore femur with dark inner subapical spot; INBio specimen with 

mid tibia only brownish at base. Guatemala female with mid 

coxa mostly yellow, mid tibia yellow with only faint basal band, 

fore femur yellow, and basal 2⁄3 of fore basotarsomere yellow.

Male Genitalia. As described for N. claripennis, except posteromedial margin of surstylus sometimes slightly 

more broadly rounded. 

Female (Figure 17). As described for male except as follows: body length 7.0–9.2 mm; first flagellomere more 

broadly ovate in outline, darker and faintly infuscated on distal 

half; yellow abdominal bristles sometimes only extending to tergite 2 or 3; fore basotarsomere lighter on basal 2⁄3; T. flavohirta

and T. pallidipennis types with small shining anterolateral spots 

on ocellar disc; anterolateral reflective triangle on frons usually 

slightly longer and mid and hind tibiae slightly darker.

Distribution. Costa Rica, Ecuador, Mexico.

Holotype [ornatipes]. MEXICO [no label]. “T. 

ornatipes, ex coll. Bigot” (1♂, UMO).

Lectotype [pallidipennis]. MEXICO [no label].

“T. pallidipennis, ex coll. Bigot” (1♀, UMO).

Paralectotypes [pallidipennis]. MEXICO [no 

label]. “T. pallidipennis, ex coll. Bigot” (2♀, UMO).

Holotype [flavohirta]. COSTA RICA. H. Schmidt 

S. (1♀, PAN).

Syntypes [mexicana]. MEXICO. Orizaba; Tuxpango (type information unknown, ?MRSN). [Not examined.]

Additional Material Examined. “T. ornatipes, ex coll. Bigot” [white label] (1♂, UMO). COSTA RICA. 

Zarzero, Schild, & Burgdorr (1♂, USNM), Guanacaste: Est 

Cacao, 1100–1200 m, 15.ii.1996, S. Marshall (1♀, DEBU), Heredia: Santo Domingo, Santa Rosa, INBio, 1100 m, 14.ii.2003, 

F. C. Thompson, manual (1♂, INBio), Santo Domingo, INBio Pk., 

21.viii.2001, K. N. Barber, pooter, leaves along trails, 09°58.4′N, 

84°05.6′W (3♂ 1♀, DEBU), Santo Domingo, INBio Pk., 

18.viii.2001, K. N. Barber, pooter, on Heliconia leaves over long 

lawn grass, 09°58.4′N, 84°05.6′W (1♀, DEBU), Santo Domingo, 

INBio Pk., 9°58′23′′N, 84°06′30′′W, 12–14.ii.2003, O. Lonsdale 

(1♀, DEBU), INBio Pk., 19–21.viii.2001, S. A. Marshall (1♀, 

DEBU), Puntarenas: Monteverde, San Luis Ecolodge, Espajito 

trail, Malaise, 1–24.iii.2007, P. D. Careless (2♂ 3♀, DEBU), 
number 641 • 3 1

Punta Leona, forest around hotel, 21–23.ii.2001, P. D. Careless (1♂, DEBU), San José: Farm La Caja, “ii-10”, H. Schmidt 

(1♂, USNM), San Carlos, Cerro Cura, 14 km SSW San Marcos 

de Tarrazu, 1800 m, 9°36′31′′N, 84°07′09′′W, 15.ii.2003, S. A. 

Marshall (1♀, DEBU). ECUADOR. Napo: Res. Ethnica Waorani, 1 km S Onkone Gare Camp, Trans. Ent., 4.x.1994, 220 

m, 00°39′10′′S, 76°26′W, T. L. Erwin et al., insecticidal fogging 

of mostly bare green leaves, some with covering of lichenous or 

bryophytic plants in terre firme forest, Project MAXUS, at trans. 

2, sta. 1, lot 860 (1♂, USNM), 4.x.1996, 216.3 m, 00°39′25.7′′S, 

76°27′10.8′′W, at trans. 10, sta. 73, lot 1757 (1♂, USNM). GUATEMALA. El Naranjo, Chicacao, “vii 6–45”, E. J. Hambleton 

(1♀, USNM), Coban, Alta Vera Paz., 22.v.1926, J. M. Aldrich 

(1♂ 1♀, USNM), La Providencia, C. Rouillard (2♂ 7♀, USNM), 

Biotopo del Quetzal, near Purulha, at light and sweeping, 21–22.

viii.1995, D. Langor (1♀, CNC). MEXICO. Nr. Chapulhuacan, 

Hidalgo, 22.xii.1961 (1♀, USNM), Chiapas: SE side of Cerro 

Tres Picos, montane rainforest, 1524–1829 m, 28.v.1972, D. E. 

Breedlove (1♀, CASC), Veracruz: 3 mi N. Huatusco, 17.vii.1980, 

Schaffner, Weaver, & Freidlander (1♂, TAMU).

Possibly N. ornatipes. COSTA RICA. San Jose: 

Escazu, 8.ii.1987, G. E. Bohart (1♂, EMUS).

Comments. Neotanypeza ochrifemur is most similar in morphology to N. ornatipes and may eventually prove to 

be conspecific, along with its existing synonyms Tanypeza mexicana, Taeniaptera pallidipennis, and Tanypeza flavohirta. The 

characters used to delimit these two species include the pattern 

on the ocellar disc and the color of the abdominal bristles, mid 

tibia, and mid and hind basotarsomeres (see couplet 21 of the 

key). Those taxa with a yellow palpus and an entirely brown 

abdomen overlap widely in morphology, and the only species 

that appears to be reliably distinct from all others is N. flavicalx, 

having a very short anterolateral reflective triangle on the frons 

and tibiae that are entirely yellow on the distal half; existing specific boundaries should be reexamined when additional data are 

available. Three of the four described syntypes of T. pallidipennis

were examined, one of which is here designated as the lectotype.

In addition to the material listed above, one male and several 

females from Ecuador (CASC) and Chiapas (TAU) tentatively 

identified as this species largely agree with the above description, 

but their color reveals affinities to N. claripennis and N. ochrifemur. The male has yellow hairs along the length of the abdomen 

laterally, but the palpus is light brown, the fore femur is entirely 

yellow, the hind tibia is yellow distally, and the base of the basotarsomeres of all tarsi is yellow. The females also have a palpus 

that varies from relatively dark to mottled yellow, and the yellow 

abdominal bristles are restricted to the first tergite. 

Neotanypeza plotoplax, sp. nov.

FIGURES 70–72

Male. Body length 10.3 mm. Dorsal portion of anterior tomentose triangle approximately 1.5 times longer than 

wide at widest point; dark portion of frons with purple tint. 

Ocellar disc fully tomentose. Narrow bronze pruinose stripe on 

notum in front of scutellum. Fronto-orbital bristles absent. Postvertical bristle minute. Subgenal bristles yellow and nearly indistinct. Antenna entirely dark brown/black, with first flagellomere 

relatively wide but not broadly rounded. Clypeus and palpus 

dark brown. Postpronotal and presutural intra-alar bristles absent. One pair of dorsocentral bristles (both missing: pin through 

right socket, but left socket visible). Anepisternum with 1 dark, 

dominant bristle. Calypter hairs dark brown. Wing dusky with 

anterodistal margin darker and first radial cell yellowish. Legs 

entirely dark brown with fore coxa, basal half of fore and mid 

femora, and all but apex of hind femur yellow. Abdominal bristles dark brown; abdomen dark brown.

Male Genitalia (Figures 70–72). As described for N. claripennis except as follows: sternite 6 with irregular right margin, soft and convoluted distal margin, narrow 

basal right sublateral sclerotization, and long left sublateral band; 

sternite 7 not evident; sternite 8 with shallow skin-like texture; 

surstylus broadly rounded, narrowing to base, only setulose near 

base and without spine-like setulae; hypandrium bare; apex of 

postgonite clear and only membranously attached to base, which 

is almost entirely atrophied on right side; flat arrowhead-shaped 

plate between lobes of hypandrium in front of phallapodeme, 

curving ventrally to apex; distiphallus short and dark with deep, 

clear distomedial emargination and 1 pair of spinulose dorsobasal processes.

Female. Unknown.

Etymology. The specific name compounds the 

Greek words for floating (plotos) and plate or tablet (plax), referring to the characteristic free sclerite between the lobes of the 

hypandrium.

Distribution. Ecuador.

Holotype. ECUADOR. Pichincha: Bellavista Reserve, trail “B”, 2200 m, 30.x.1999, S. A. Marshall (1♂, QCAZ).

Comments. Externally, this species is distinctive 

in having one pair of dorsocentrals, no fronto-orbitals, dusky 

wings that are slightly darkened anterodistally, dark brown calypter hairs, and similarly dark abdominal bristles. Regarding 

the male genitalia, sternite 6 has a second small sclerotized patch 

on the right side basally, the distiphallus is distinct, the surstylus 

is broad and largely bare, and the postgonite is atrophied. Most 

unique, however, is the ventral, floating, arrowhead-shaped plate 

between the halves of the hypandrium, which is likely homologous with the anterior process of the phallapodeme.

Neotanypeza posthos, sp. nov.

FIGURES 73–75

Male. Body length 9.0 mm. Dorsal portion of anterior tomentose triangle approximately 1.5 times longer than 

wide at widest point. Ocellar disc tomentose with portion bordering ocellar tubercle shining. Frons (excluding reflective parts) 
3 2 • smithsonian contributions to zoology

black with purple tint. One fronto-orbital bristle. One subgenal 

bristle brown, 1 yellow. Postvertical bristle minute. First flagellomere slightly thickened and dark with base orange. Clypeus and 

palpus dark brown. Postpronotal and presutural intra-alar bristles absent. One dorsocentral bristle. Anepisternum with 1 dark, 

dominant bristle. Calypter hairs brown (golden in some Bolivian 

specimens). Wing very lightly infuscated anterodistally (clear in 

Ecuadorian male), with overall hue yellowish. Legs yellow with 

mid and hind coxae brown, fore femur with broad inner distal 

spot, mid and hind femora dark apically, fore tibia sometimes 

lightly infuscated on basal half (if so, distal half light yellow), 

and mid and hind tibiae dark brown with yellowish subapical 

band on mid tibia; tarsi dark brown. Abdominal bristles dark 

brown; abdomen dark brown.

Variation. Two smaller paratype males from Ecuador and Venezuela (7.7–8.1 mm) with outer face of fore femur 

and all of mid and hind femora yellow, mid and hind tibiae entirely dark brown, hind coxa yellow, subgenal bristles yellow, 

and antenna entirely black with orange base. 

Male Genitalia (Figures 73–75). As described for N. claripennis except as follows: sternite 6 nearly 

symmetrical and without sclerotized band; sternite 7 not evident; sternite 8 with shallow skin-like texture dorsally; spinulose 

setulae on surstylus restricted to narrow basal patch; distiphallus 

long and dark, slightly expanding apically, and with minute sublateral spinules converging to basal 1⁄3.

Female. As described for male except as follows: 

length 6.8–8.0 mm; reflective triangle on frons extending past 

fronto-orbital in Peruvian female; ocellar disc entirely tomentose; 

subgenal bristles yellow (slightly darker in Peruvian female); first 

flagellomere broadly ovate and orange with dorsal and apical 

margins dark brown; calypter hairs yellow; hind coxa sometimes 

yellow; yellowish subapical band faint on hind tibia in Bolivian 

female, and bands on both legs strong in Peruvian female; fore 

basotarsomere light yellow on basal 3/5– 2/3 (entirely light with 

apex brown in Peruvian female, which is also weakly pigmented 

on thorax and abdomen but not head or legs); Colombian female 

with femora broadly dark brown apically and bristles on first and 

second tergites yellow; wing more darkly and widely infuscated.

Etymology. The specific epithet is Greek for “one 

with a large penis.”

Distribution. Bolivia, Colombia, Ecuador, Peru, 

Venezuela.

Holotype. BOLIVIA. La Paz: Coroico, Cerro 

Uchumachi, 2550 m, 16°12′43′′S, 67°42′49′′W, cloud forest, 

5.iv.2001, S. A. Marshall (1♂, CBFC).

Paratypes. BOLIVIA. Incochoca, Cochab, 25–31.

viii.1956, L. E. Pena (1♂, CNC), La Paz: Coroico, Cerro Uchumachi, 2550 m, 16°12′43′′S, 67°42′49′′W, cloud forest, 5.iv.2001, S. 

A. Marshall (1♂, DEBU). ECUADOR. 40 mi S Alausl, Chimborazo, 20.ii.1955 (1♂, CASC), Napo: Baeza, 2000 m, 1.iii.1979, 

S. A. Marshall (1♂, QCAZ). PERU. Cusco, Wayqecha Biol. 

Stn., ~9 km NE Challabamba, 13°10′S, 71°35′W, ~2800 m, 13–

15.v.2007, S. A. Marshall (1♂, DEBU). VENEZUELA. Aragua 

Colonia Tovar, 2300 m, sweep, steep forest trail, 10.iii.1995, S. 

A. Marshall (1♂, DEBU).

Additional Material Examined. BOLIVIA. 

Br. Ichilo R., rain forest, 15–16.x.1966, B. & K. Burks (1♀, 

USNM), Paracti, Chapare, 1–4.ii.1976, 2200 m, L. E. Pena (1♀, 

CNC), La Paz, Cumbre Alto Beni, 28 km E Caranavi, ~1400 

m, 15°40′31′′S, 67°29′21′′W, cloud forest, 14.iv.2001, S. A. 

Marshall (1♀, DEBU). COLOMBIA. Ant. Queb. La Iguana, 

17 km NW Medellin, 22.ii.1984, C. M. & O. S. Flint, Jr. (1♀, 

USNM). ECUADOR. Napo, Baeza, W along road, 1500 m, 

montane rain forest/pasture, Malaise trap, 16–19.v.1987, Coote 

& Brown (1♀, DEBU), SierrAzul Res., 14 km W Cosanga, 2200 

m, 0°40′55′′S, 77°58′09′′W, 10.v.2002, M. Buck (5♀, DEBU).

PERU. Cuzco, Pueblo, Machu Picchu, 21.iii.1947, 6491 ft, D. C. 

Pallister coll., F. Johnson donor (1♀, AMNH), Cusco, Wayqecha 

Biol. Stn., ~9 km NE Challabamba, 13°10′S, 71°35′W, ~2800 m, 

13–15.v.2007, S. A. Marshall (1♀, DEBU). 

Comments. Neotanypeza posthos is a relatively 

large species with one dorsocentral, one fronto-orbital, a nearly 

symmetrical sternite 6, and a long, dark distiphallus (hence the 

specific name) with a smooth medial plate surrounded by minute 

spinules. The abdominal bristles are also usually entirely dark, 

and the calypter hairs are brown to golden in color. Since the 

most diagnostic characters of this new species are male genitalic, the females listed above are only tentatively included in this 

taxon and are not included in the type series. 

Neotanypeza quadrisetosa (Enderlein)

Polphopeza quadrisetosa Enderlein, 1913:227. Enderlein, 1936:40.

Neotanypeza quadrisetosa.—Hennig, 1936:35.

Neotanypeza (Polphopeza) quadrisetosa.—Steyskal, 1967:2.

Male. Body length 8.7–9.4 mm. Dorsal portion of 

anterior tomentose triangle approximately 2 times longer than 

wide at widest point. Ocellar disc tomentose with anterior 

margin around tubercle shining. Anterior fronto-orbital bristle 

absent. Postvertical bristle minute. First flagellomere relatively 

narrow and brown with base orange. Clypeus and palpus dark 

brown. Postpronotal bristle present and presutural intra-alar 

bristle absent. Two dorsocentral bristles. Anepisternum with 1 

dark, dominant bristle. Calypter hairs brown. Wing clear, sometimes with infuscation around distal half of R2+3. Legs yellow 

with mid coxa brown, fore femur dark on distal quarter (more 

extensive on inner face), mid and hind femora with apex brown, 

fore tibia brown on basal half, and mid and hind tibiae dark 

brown, sometimes with faint yellow subapical bands; tarsi dark 

brown. Abdominal bristles dark brown; abdomen dark brown.

Male Genitalia. As described for N. claripennis, 

except sternite 61⁄3 longer. 

Female. Unknown.

Distribution. Ecuador.

Holotype. Ecuador. Baños, R. Haensch S. (1♂, 

PAN).
number 641 • 3 3

Additional Material Examined. Ecuador. Baños, 19.ii.1937, S. W. Frost (2♂, USNM), Napo: SierrAzul Res., 14 km W Cosanga, 2200 m, 0°40′55′′S, 77°56′09′′W, 

8.v.2002, O. Lonsdale (1♂, DEBU), 10.v.2002, M. Buck (1♂, 

DEBU), SierrAzul Lodge, 14 km W Cosanga, 2200 m, 0°40′55′′S, 

77°56′09′′W, Malaise, 9–10.v.2002, Buck & Paiero (1♂, QCAZ).

Comments. Neotanypeza quadrisetosa is a weakly 

defined Ecuadorian species with relatively undifferentiated male 

genitalia. At present it can only be diagnosed by using the combination of characters presented in the key.

Neotanypeza rutila (Wulp)

FIGURES 15, 76–78

Tanypeza rutila Wulp, 1897:362. Wulp, 1897: tab. ix. 

Tanypeza (Neotanypeza) rutila.—Enderlein, 1913:225.

Neotanypeza rutila.—Hennig, 1936:37.

Tritanypeza rutila.—Enderlein, 1936:43.

Neotanypeza (Tritanypeza) rutila.—Steyskal, 1967:2.

Male (Figure 15). Body length 7.0–8.2 mm. Anterior tomentose triangle on frons tan colored and reaching level of 

posterior fronto-orbital. Ocellar disc fully tomentose. Anterior 

fronto-orbital approximately half length of posterior bristle and 

thin. Postvertical bristle yellow, thin, and slightly longer than tubercle. First flagellomere light yellow and narrow, tapering to 

apex. Clypeus yellow (dark brown in males with more extensively pigmented abdomens) and palpus yellow. Postpronotal and 

presutural intra-alar bristles present. Three dorsocentral bristles. 

Thorax dark brown. Anepisternum with only 1 dark, dominant 

bristle. Calypter hairs pale. Wing clear. Face mostly white with 

dorsal and lateral markings on face faded. Legs entirely yellow 

with mid coxa variably orange to brown, fore tarsi darker yellow 

distally (and pale basally in holotype), and mid and hind tarsi 

brown with mid basotarsomere yellow and hind basotarsomere 

yellow basally; fore and hind tarsi sometimes light brown with 

base paler. Abdominal bristles dark brown with lateral bristles 

on tergite 1 yellow; abdomen dark brown with sternite 8 and 

segments 1–5 yellow (darker on segments 4 and 5), sometimes 

with posterior margin brown on tergites 3–5 and sometimes with 

brown medial stripe on tergites 3 and 4.

Male Genitalia (Figures 76–78). As described for N. claripennis except as follows: sternite 6 nearly 

symmetrical and membranous on left anterior margin, without 

sclerotized sublateral band and with 2 long distomedial bristles; 

sternite 7 well developed and with 2 bristles; surstylus broad, 

doubly sinuate on posterior margin, densely setose, and without stout pointed setulae; hypandrium with multiple bristles on 

each side; postgonite with subapical setulae; distiphallus pointed 

at base, bifid, loosely associated with separate basiphallus, and 

heavily spinulose laterally and medially.

Female. As described for male except as follows: 

body length 7.1–8.2 mm; postvertical black; first flagellomere 

orange with posterior margin broadly rounded and sometimes 

with distal margin lightly infuscated; thorax yellow on anepisternum, postpronotum, notopleuron, prosternum, anterior half of 

meron (mottled, with suture brown), and dorsal half of katepisternum; legs yellow with mid coxa brown, fore tibia light yellow, and hind tibia light yellow with basal half dark brown; fore 

tarsi brown with basotarsomere yellow, excluding apex (basal 

half yellowish in TAMU female); abdomen dark brown, with 

segments 1–6 predominantly yellow; tergites (1)2–4(5) with narrow central stripe (stripe sometimes wide and flaring posteriorly 

on tergites 3–5), posterior margin of tergites (3)4–6 brown, and 

tergites 5 and 6 sometimes brownish to brown.

Variation. Brazilian males: postvertical brown, prosternum dark yellow with brownish medial spots, hind tibia with 

very light subapical band, tarsi brown with basal 2⁄3 of basotarsomeres yellow, clypeus brown, and first flagellomere broader; abdominal coloration indistinct past segment 2. One Costa Rican 

male (INBio) with thorax entirely dark brown. TAMU female 

with thorax (excluding prosternum) brown, hind tibia with distal brown band, basotarsomeres yellow to base on mid and hind 

legs and light yellow on basal half of fore basotarsomere, and abdomen dark with tergites 1–3 yellow (excluding narrow medial 

stripe and brown posterior margin on tergite 3).

Distribution. Costa Rica, Mexico.

Lectotype. MEXICO. Guerrero, Sierra de las 

Aguas Escondidas, 3000 ft, “July”, “cotype”, “Cent. America, 

F. D. Godmando, O. Salvin, B. M. 1903-172” (1♂, BMNH). 

Paralectotypes. MEXICO. Same data as lectotype (2♀, BMNH).

Additional Material Examined. COSTA 

RICA. San Jose: Escazu, F. D. Parker, 24–30.i.1988 (1♂, EMUS), 

13–15.iv.1988 (1♀, EMUS), Guanacaste: 3 km SE R. Naranjo, 

12–14.v.1993, F. D. Parker (1♀, EMUS), Liberia, P. N. Sta. 

Rosa, Est. Santa Rosa, Bque Húmedo, 300 m, viii.1999, Janzen, 

manual (5♀ 1♂, INBio), P. N. Sta. Rosa, Stor Santa Rosa, Area 

Adm., 300 m, J. Sullivan & A. A. Pérez, Malaise, vi.1999 (1♀, 

INBio), iii.1999 (1♂, INBio), Estacion Pitilla, 9 km S de Santa 

Cecilia, 700 m, iii.1995, P. Rios (1♀, INBio), Los Almendros, 

P. N. Guanacaste, 7–26.i.1993, E. López (1♂, INBio). MEXICO. 

Jalisco: 16 mi SW Autlan, 22.viii.1976, Hanson, Schwartz (1♀, 

EMUS), Jalisco: rd. to Parque Nacional de Volcan de Colima, 11 

mi W Hwy. junct. 54 (near Atenquique), 11–12.vii.1984, Carroll, Schaffner, & Freidlander (1♀, TAMU). 

Comments. Neotanypeza rutila is highly variable in 

its color pattern, although all females and some males have a yellow prosternum and those males with a brown prosternum have 

a largely yellow abdomen and clypeus. Northern specimens also 

tend to be paler in overall pigmentation. This species is similar in 

appearance to N. abdominalis, and there is a significant overlap 

in coloration; however, females of N. rutila are much paler on 

the thorax, and males have a different pattern on the frons and 

an unusual distiphallus. 

The male syntype of Neotanypeza rutila is here designated 

as the lectotype.
3 4 • smithsonian contributions to zoology

Neotanypeza symmetros, sp. nov.

FIGURES 79–84

Male. Body length approximately 7.5 mm. Dorsal 

portion of anterior tomentose triangle not much longer than 

wide at widest point; frons (excluding reflective patches) black 

with purple tint anteromedially. Ocellar disc fully tomentose. 

One fronto-orbital. Postvertical bristle slightly longer than tubercle and thin. First flagellomere broadly rounded and infuscated along distal and dorsal margins. Clypeus dark brown and 

palpus yellow. Postpronotal and presutural intra-alar bristles absent. One dorsocentral bristle. Anepisternum with 1 dark, dominant bristle. Calypter hairs yellow. Wing infuscated apically. 

Legs yellow with mid coxa brown, fore femur lightly pigmented 

subapically on inner dorsal surface, mid and hind femora dark 

brown on distal half excluding apex, fore tibia and tarsi light 

yellow with distal tarsomere and apex of penultimate tarsomere 

brown, and mid and hind tibiae and tarsi dark brown. Abdominal bristles dark brown; abdomen dark brown.

Male Genitalia (Figures 79–84). As described for N. claripennis except as follows: sternite 1 apparently fused to sternite 2, elongate; sternite 3 relatively elongate; 

sternite 6 virtually symmetrical, shallowly emarginate at base, 

without sublateral sclerotization, evenly setose, and without 

anteromedial setulae; sternite 7 well developed; surstylus with 

bristles restricted to posterior half and with setulae restricted to 

posterodorsal corner (none stout and pointed); phallapodeme 

and distiphallus slightly longer with distiphallus wide and 

toothed on distal half with relatively long, narrow base and no 

dark sclerotized medial region.

Female. Unknown.

Etymology. The specific epithet refers to the nearly 

symmetrical male sternite 6.

Distribution. Colombia.

Holotype. COLOMBIA. Valle Paramo Las Hermosas, 26.x.1975, 2700 m, J. E. Lattke, montane rainforest, Malaise trap (1♂, CASC).

Comments. Neotanypeza symmetros is a relatively 

small (7.5 mm), pale species with a single dorsocentral and 

fronto-orbital, a broad, dark first flagellomere, dark legs, a yellow palpus, pale calypter hairs, and an unusually long, narrow 

abdomen. The first three sternites are particularly elongate, with 

sternite 1 apparently fused to sternite 2, which form a single 

plate. 

Neotanypeza vexilla, sp. nov.

FIGURES 32, 85, 86

Male. Body length approximately 9.5 mm. Dorsal 

portion of anterior tomentose triangle approximately 1.5 times 

longer than wide at widest point. Ocellar disc fully tomentose. 

Fronto-orbital bristles absent. Postvertical bristle minute. First 

flagellomere relatively wide (but not broadly rounded) and orange with distal 2⁄3 dark. Clypeus and palpus dark brown. Frons 

black with light purple tint. Postpronotal and presutural intraalar bristles absent. One dorsocentral bristle. Anepisternum with 

1 dark, dominant bristle. Calypter hairs yellow. Wing dusky with 

dark apical spot around R2+3. Legs yellow with mid coxa brown, 

fore femur with dark inner distal spot and light outer distal spot, 

mid femur light brown apically, hind femur brownish at apex, 

fore tibia with broad basal and narrow apical light brown bands, 

and mid and hind tibiae and all tarsi dark brown. Abdominal 

bristles dark brown with lateral bristles on first tergite yellow; 

abdomen dark brown.

Male Genitalia (Figures 85, 86). As described for N. claripennis except as follows: sternite 7 well developed but not strongly differentiated from sternite 8; surstylus 

nearly as wide as epandrium, broadly rounded, and with stout 

setulae partially extending around margin of surstylus; hypandrium bare; distiphallus slightly narrower and without medial 

sclerotization. Male with pale wings with distiphallus narrow 

and very slightly narrowing apically, with spinulose margins less 

textured and meeting medially.

Female. As described for male except as follows: 

body length 9.5–11.0 mm; frons black, without purple tint; antenna black; fore femur light brown apically; hind femur and fore 

tibia entirely yellow; fore basotarsomere entirely light yellow.

Variation. CNC female from Peru (Figure 32) differs from above females as follows: body length approximately 

10.0 mm; wing yellowish anteriorly, with light infuscation along 

CuA1

 continuing up dm-cu and then down M1; R4+5 infuscated 

along distal section and R2+3 with dark apical spot; fore femur 

yellow with light inner distal spot; fore basotarsomere only yellowish at base. DEBU female similar to CNC female except mid 

and hind tibiae with yellowish subapical band and base of hind 

basotarsomere yellow. 

Variation: Nontype Specimens. One Ecuadorian male (CNC) largely agrees with above description but 

differs as follows: body length 8.1 mm; fore femur yellow on 

outer surface and becoming darker yellow basally; sternite 8 yellow; epandrium and surstylus with yellowish mottling; distiphallus only 1⁄4 length at widest point, with lateral margin sclerotized 

and center with narrow, tapered sclerite extending for 2⁄3 length 

of distiphallus. This male possibly belongs to a separate species, 

but additional material would be needed before it could be determined whether or not the differences observed represent normal intraspecific variation. A similar Venezuelan female from the 

AMNH may also be conspecific: length 8.4 mm; margins of ocellar disc shiny; first flagellomere broad; wing slightly darker; fore 

femur brownish on outer apex; fore tibia light brown on basal 

half; mid and hind tibiae with yellowish subapical band; sternite 8 brown; yellow lateral abdominal bristles extending along 

length of abdomen.

Etymology. The specific name is derived from the 

Latin for banner, flag, or standard, referring to the characteristic 

wings, which are likely used for conspecific signaling.
number 641 • 3 5

Distribution. Ecuador, Colombia, Peru, Venezuela.

Holotype. ECUADOR. SierrAzul Res., 14 km W 

Cosanga, 2200 m, 0°40′55′′S, 77°58′09′′W, 8–11.v.2002, S. A. 

Marshall (1♂, QCAZ). 

Paratypes. COLOMBIA. Boyacá, SFF Iguaque, 

5°25′12′′N, 73°27′24′′W, 2850 m, 1–19.iv.2000, Malaise, 

P. Reina (1♀, DEBU). PERU. Cuzco, Machu Picchu, 21.xii.1967, 

G. E. Bohart (1♀, CNC), Cusco, Aguas Calientes, 15–16.vi.2006, 

Paiero & Klymko (1♀, DEBU).

Additional Material Examined. ECUADOR. Tandapi, 40 km SW Quito, 1300–1500 m, 15–21.vi.1965, 

Pena (1♂, CNC). VENEZUELA. Lindig, 1864, “elegans”, “Alte 

Sammlung” (1♀, AMNH).

Comments. Although somewhat variable in coloration, Neotanypeza vexilla can be diagnosed by a subapical 

spot on vein R2+3 and a reduction in chaetotaxy. Neotanypeza 

leucothrix (Peru) also has an apical spot on R2+3, but the frons is 

shining anteromedially, the posterior margin of the ocellar disc 

is shining, there is one pair of fronto-orbital bristles, the postpronotal bristle in present, the fore tibia is not banded as above, 

the basotarsomeres are yellow basally, the abdominal bristles are 

brown on the first tergite, and the abdomen has yellow spots.

Acknowledgments

I thank the following curators who loaned material or 

helped to locate types: D. Grimaldi (AMNH); R. Vane-Wright 

and N. Wyatt (BMNH); K. Ribardo (CASC); J. M. Cumming 

(CNC); V. Pechumann, Department of Entomology, Cornell University, Ithaca, New York; M. Buck and S. A. Marshall (DEBU); 

P. Mourie and L. Marinoni, Departmento de Zoologia, Universidade Federal do Parana, Curitiba, Brazil; W. J. Hanson (EMUS); 

M. Zumbado (INBio); A. Henriques (INPA); A. O. Bachman and 

A. Oliva, Museo Argentino de Ciencias Naturales “Bernardino 

Rivadavia,” Buenos Aires; C. Pesarini, Museo Civico di Storia 

Naturale, Milan, Italy; E. Cevallos Saenz, Museo de Historia 

Natural “Javier Prado,” Lima; M. Daccordi (MRSN); N. Papavero and F. C. do Val, Museu de Zoologia, Universidade de São 

Paulo, São Paulo, Brazil; R. Contreras-Lichtenberg and P. Sehnal 

(NMW); R. Bankowska, M. J. Sapiejewski, and T. Huflejt 

(PAN); W. Tobias, Senckenbergisches Museum, Frankfurt, Germany; U. Kallweit and C. Kehlmaier (SMT); E. Riley (TAMU); 

H. Toro, Instituto de Biologia, Departmento de Ciencias, Universidad Católica de Valparaíso, Valparaíso, Chile; A. Pont (UMO); 

A. Contreras-Ramos, Instituto de Biología, Universidad Nacional Autónoma de Mexico, Mexico City; V. Michelson (UZMC); 

and W. Mey (ZMHU). C. Micheli and H. Williams (USNM) 

were extremely helpful with the handling of material. 

Photographs of live specimens (Figures 1–6) were provided 

courtesy of S. A. Marshall, who also collected much of the material examined here. Photographs of the Nartshukia musiva holotype (Figure 34) were provided by A. Shatalkin (ZMUM), who 

also kindly reviewed this manuscript. K. Barber and an anonymous reviewer also provided extremely helpful comments on 

the submitted manuscript. Lastly, I thank F. Christian Thompson (USNM), who ordered type material, examined the types of 

N. rutila on my behalf, and provided much nomenclatural advice 

during my stay at the Smithsonian as a postdoctoral fellow. Funding for this study was provided by a Schlinger Postdoctoral Grant.

Apigian, K., and F. C. Thompson. 2010. “58. Tanypezidae (Tanypezid Flies).” In Manual of Central American 

Diptera, volume 2, ed. B. V. Brown, A. Borkent, J. M. Cumming, N. E. Woodley, and M. A. Zumbado, pp. 

827–831. Ottawa: NRC Research Press. 

Barber, K. N. 2006. Strongylophthalmyia pengellyi n. sp., a Second Species of Nearctic Strongylophthalmyiidae 

(Diptera). Journal of the Entomological Society of Ontario, 137:81–109.

Barraclough, D. A. 2000. The Identity of Strongylophthalmyia Heller Species (Diptera: Schizophora: Strongylophthalmyiidae) Described from the Afrotropical Region, and Their Transfer to the Family Clusiidae. Annals of 

the Natal Museum, 41:103–106.

Bigot, J. M. F. 1886. Diptères nouveaux ou peu connus. 29e. partie. XXXVII. 1er. Essai d’une classification synoptique de groupe des Tanypezidi (mihi) et description de genres et d’espèces inédits. Annales de la Societe 

Entomologique de France, 6:287–392. 

Chandler, P. J. 1975. Notes on the British Status of Three Unusual Acalypterate Flies (Diptera). Proceedings of the 

British Entomological and Natural History Society, 8:66–72. 

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delta-intkey.com/www/refs.htm (accessed January 2008). 

Enderlein, G. 1913. Dipterologische Studien II. Zur Kenntnis der Tanypezinen. Zoologischer Anzeiger, 42:224–

229.

———. 1936. Klassifikation der Tanypeziden. Deutsche Entomologische Zeitschrift, 1936:39–47. 

Evenhuis, N. L. 1989. “Family Strongylophthalmyiidae.” In Catalog of the Diptera of the Australasian and Oceanian Regions, ed. N. L. Evenhuis, p. 472. Honolulu: Bishop Museum Press.

Fallén, C. F. 1820. Opomyzides Sveciae. Lund, Sweden: Berlingianis. 

Feijen, H. R. 1983. Systematics and Phylogeny of Centrioncidae, a New Afromontane Family of Diptera 

(Schizophora). Zoologische Verhandelingen (Leiden), 202:1–137. 

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Giglio-Tos, E. 1893. Diagnosi di nuovi generi e di nuove specie di Ditteri. IX. Bollettino del Museo di Zoologia ed 

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Krivosheina, N. P. 1981. New Palaearctic Species of Genus Strongylophthalmyia Hell. (Diptera Strongylophthalmyiidae). Entomologischeskoe Obozrenie, 60:183–186.

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number 641 • 3 9

FIGURES

FIGURES 1–6. Figure 1: Strongylophthalmyia angustipennis Melander, male and female in copula, Ontario, Canada. Figure 2: Tanypeza 

longimana Fallén, female, Ontario, Canada. Figure 3: Neotanypeza posthos, sp. nov., female, Wayqecha Biological Station, Peru. Figure 

4: N. dallasi (Shannon), female, Savegre Lodge, Costa Rica. Figure 5: N. grandis (Enderlein), female, SierrAzul Research Station, Ecuador.

Figure 6: N. alopecia, sp. nov., female, Heath River Wildlife Center, Bolivia. All photos courtesy S. A. Marshall.
4 0 • smithsonian contributions to zoology

FIGURES 7–11. Figures 7–8: Neotanypeza marshalli, sp. nov.; Figure 7: head, anterior; Figure 8: body, lateral. Figure 9: 

Tanypeza longimana Fallén, ventral. Figure 10: Strongylophthalmyia angustipennis Melander, prosternum and base of 

fore coxae. Figure 11: Centrioncus prodiopsis Speiser (Diposidae), prosternum and base of fore coxae. Abbreviations: fo 

a = anterior fronto-orbital bristle; fo p = posterior fronto-orbital bristle; npl = notopleural bristle; oc = ocellar bristle; pprn 

= postpronotal bristle; pv = postvertical bristle; sgen = subgenal bristles; v s = vertical bristle.
number 641 • 4 1

FIGURES 12–17. Heads, anterodorsal view. Figure 12: Neotanypeza claripennis (Schiner), male lectotype. Figure 13: N. claripennis, tentatively included male. Figure 14: N. apicalis (Wiedemann), male. Figure 15: N. rutila (Wulp), male. Figure 16: N. ornatipes

(Bigot), holotype, male. Figure 17: N. ornatipes, female (Taeniaptera pallidipennis Bigot, lectotype).
4 2 • smithsonian contributions to zoology

FIGURES 18–23. Heads, anterodorsal view. Figure 18: Neotanypeza abdominalis (Wiedemann), female. Figure 19: N. micans, sp. 

nov., holotype, male. Figure 20: N. alopecia, sp. nov., paratype, female. Figure 21: N. flavicalx (Enderlein), holotype, female. Figure 

22: N. grandis (Enderlein), male. Figure 23: N. argentia, sp. nov., holotype, male.
number 641 • 4 3

FIGURES 24–34. Figures 24–32: Left wing, ventral; Figure 24: Strongylophthalmyia angustipennis

Melander (abbreviations: sc, subcostal; c br, subcostal break; all other labeling = actual vein or cell 

names); Figure 25: Tanypeza longimana Fallén; Figure 26: Neotanypeza claripennis (Schiner), with 

arrow indicating pointed deviation along margin of cell dm referred to in character 42; Figure 27: N. 

claripennis, tentatively included specimen; Figure 28: N. grandis (Enderlein); Figure 29: N. marshalli,

sp. nov.; Figure 30: N. dimorpha (Hennig), female; Figure 31: N. dimorpha, male; Figure 32: N. 

vexilla, sp. nov., Peruvian female. Figure 33: N. apicalis (Wiedemann), holotype, dorsal. Figure 34:

Nartshukia musiva Shatalkin, holotype, lateral with wing inset; photo courtesy A. Shatalkin.
4 4 • smithsonian contributions to zoology

FIGURES 35–40. Strongylophthalmyia angustipennis Melander, male abdomen. Figure 35: External components of genitalia, left lateral. Figure 36: Sternites 6–8, ventral. Figure 37: External genitalia, posterior. Figure 38: Hypandrial complex, 

left lateral, showing details of membrane texture on phallus. Figure 39: Hypandrial complex, ventral, phallus removed. 

Figure 40: Ventral (not to scale; dotted lines delimiting medially bare, desclerotized regions). Abbreviations: S = sternite; T 

= tergite. Scale bar = 0.1 mm.
number 641 • 4 5

FIGURES 41–46. Figures 41–45: Tanypeza longimana Fallén, male abdomen; Figure 41: external, posterior; Figure 42: external, 

anterior; Figure 43: external, left lateral; Figure 44: hypandrial complex, left lateral; Figure 45: ventral. Figure 46: T. picticornis 

Knab and Shannon, ventral. Abbreviations: S = sternite; T = tergite. Scale bar = 0.1 mm.
4 6 • smithsonian contributions to zoology

FIGURES 47–51. Figures 47–50: Neotanypeza elegans (Wiedemann), female abdomen; Figure 47: segments 6 to cerci, dorsal (scale bar 

= 0.5 mm); Figure 48: segments 6 to cerci, ventral; Figure 49: left lateral (scale bar = 2.0 mm); Figure 50: internal genitalia (scale bar = 

0.2 mm). Figure 51: Tanypeza longimana Fallén, ventral receptacle (scale bar = 0.2 mm). Abbreviations: S = sternite; T = tergite; IP = internal process of sternite 10.
number 641 • 4 7

FIGURES 52–57. Neotanypeza claripennis (Schiner), male abdomen. Figure 52: External genitalia, left lateral. Figure 53: 

External genitalia, posterior. Figure 54: Ventral, detailing segment 6. Figure 55: Ventral. Figure 56: Hypandrial complex, 

ventral. Figure 57: Hypandrial complex, left lateral. Abbreviations: S = sternite; T = tergite. Scale bar = 0.1 mm.
4 8 • smithsonian contributions to zoology

FIGURES 58–60. Neotanypeza apicalis (Wiedemann), male genitalia. Figure 58: Sternite 6. Figure 59: Left lateral. Figure 60: Hypandrial 

complex, ventral. Scale bar = 0.1 mm.
number 641 • 4 9

FIGURES 61–63. Neotanypeza dimorpha (Hennig), male genitalia. Figure 61: Sternite 6. Figure 62: Left lateral. Figure 63:

Hypandrial complex, ventral. Scale bar = 0.1 mm.
5 0 • smithsonian contributions to zoology

FIGURES 64–66. Neotanypeza grandis (Enderlein), male genitalia. Figure 64: Sternite 6. Figure 65: Left lateral with detail of surface texture. 

Figure 66: Hypandrial complex, ventral. Scale bar = 0.1 mm.
number 641 • 5 1

FIGURES 67–69. Neotanypeza marshalli, sp. nov., male genitalia. Figure 67: Sternite 6. Figure 68: Left lateral. Figure 69: Hypandrial complex, ventral. Scale bar = 0.1 mm.
5 2 • smithsonian contributions to zoology

FIGURES 70–72. Neotanypeza plotoplax, sp. nov., male genitalia. Figure 70: Sternite 6. Figure 71: Left lateral. Figure 72:

Hypandrial complex, ventral. Scale bar = 0.1 mm. Arrow indicates characteristic floating medial plate.
number 641 • 5 3

FIGURES 73–75. Neotanypeza posthos, sp. nov., male genitalia. Figure 73: Sternite 6. Figure 74: Left lateral. Figure 75: Hypandrial complex, 

ventral. Scale bar = 0.1 mm.
5 4 • smithsonian contributions to zoology

FIGURES 76–78. Neotanypeza rutila (Wulp), male genitalia. Figure 76: Sternite 6. Figure 77: Left lateral. Figure 78: Hypandrial complex, 

ventral. Scale bar = 0.1 mm.
number 641 • 5 5

FIGURES 79–84. Neotanypeza symmetros, sp. nov., male abdomen. Figure 79: Hypandrial complex, ventral. Figure 80: 

Hypandrial complex, left lateral. Figure 81: Ventral, detailing sternites 6–8. Figure 82: Ventral. Figure 83: External genitalia, 

left lateral. Figure 84: External genitalia, posterior. Scale bar = 0.1 mm.
5 6 • smithsonian contributions to zoology

FIGURES 85–86. Neotanypeza vexilla, sp. nov., male genitalia. Figure 85: Left lateral. Figure 86: Hypandrial complex, 

ventral. Scale bar = 0.1 mm.
number 641 • 5 7

FIGURES 87–89. Eggs. Figure 87: Neotanypeza elegans (Wiedemann). Figure 88: Tanypeza longimana Fallén. Figure 89:

Strongylophthalmyia angustipennis Melander. Scale bar = 0.3 mm.
5 8 • smithsonian contributions to zoology

FIGURE 90. Strict consensus of the 45 most parsimonious trees resulting from the phylogenetic analysis.
number 641

• 5 9

FIGURE 91. Phylogeny of the Tanypezidae, including out-groups, treating Neotanypeza as an operational taxonomic unit. Characters are listed in Table 1; solid circles indicate unique characters, and open circles are homoplasies or character reversals. 
6 0 

• smithsonian contributions to zoology

FIGURE 92. Detail of the working phylogenetic hypothesis for the species of Neotanypeza.
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