A review of the Archiceroptera Papp genus complex (Diptera:

Sphaeroceridae: Limosininae)

by

Steven Mark Paiero

A Thesis presented to The University of Guelph

In partial fulfilment of requirements for the degree of Doctor of Philosophy in Environmental Sciences

Guelph, Ontario, Canada

© Steven Mark Paiero, December, 2017

ABSTRACT:

A review of the Archiceroptera Papp genus complex (Diptera: Sphaeroceridae:

Limosininae)

Steven Mark Paiero Advisor:

University of Guelph, 2017 Dr. S.A. Marshall

This thesis has two parts. The first part investigates the relationships between the Archiceroptera genus complex and other members of the Limosininae (Diptera: Sphaeroceridae). A focus is placed on the relationships within the larger epandrial process group, which contains Bitheca, Bromeloecia,

Pterogramma, Aptilotella, and Robustagramma, along with Archiceroptera, Rudolfina and several previously unplaced species groups. Molecular and morphological data sets provide the first phylogeny of the group, and were used to support the inclusion of several unplaced species groups within Rudolfina and

Archiceroptera, while one new genus is described. Pectinosina gen. nov. includes two species: P. prominens (Duda), previously placed in Rudolfina, and P. carro n. sp. The second part of the thesis deals with revisions of Archiceroptera Papp and Rudolfina Roháček. Rudolfina now includes 13 described species, nine of which are newly described here (R. bucki, R. exuberata, R. howdeni, R. megepandria, R. pauca, R. pilosa, R. newtoni, R. remiforma, and R. tumida). Archiceroptera now includes 29 species, of which 27 are newly described here (A. adamas, A. addenda, A. barberi, A. basilia, A. bilobata, A. bisetosus, A. braziliensis, A. brevivilla, A. browni, A. caliga, A. calligraphia, A. cobolorum, A. crenulata,

A. curvivilla, A. dolabra, A. llama, A. maniba, A. masoni, A. megacercus, A. megavilla, A. mexicorona, A. mitarakai, A. paracercus, A. pussula, A. ternum, A. triclavus, and A. uncinata). ACKNOWLEDGEMENTS

My wife, partner and best friend, Jordan, for her support and patience throughout my study, especially the last gruelling months. - - -

My parents, the rest of my family and many close friends, who supported my passion over the years, along with bringing me or saving countless insects in vials just in case they were of interest!

To my advisor, Dr. Stephen Marshall, for (most importantly) sharing his passion of entomology in a field course in

1999 that changed the course of my studies, but also for sharing his immense expertise and experience in Diptera.

My committee members, Dr. Gary Umphrey and Dr. Alex Smith, for their input and suggestions during the course of my study. Access to sequence data extracted from material collected by Dr. Smith, obtained with the support of various grants, was important for the molecular examination of the broader EPG clade.

The members of my defense committee: Dr. J. Cumming, Dr. J. Fu and Dr. C. Scott-Dupree.

Val Levesque-Beaudin, for helping with the BOLD database and access to barcoded material.

Dr. Matthias Buck, who was the first person to help me appreciate sphaerocerids and many other Diptera families, during his time at the University of Guelph Insect Collection.

Everyone from the lab, past and present, with special thanks to D. Cheung, M. Jackson, and T. Yau.

I would like to thank the various individuals and institutions that made material available for my study. A. Newton

(FMNH), B. Hubley (ROME), J. Skevington and J. Kits (CNCI), N. Penny (CASC), A. Norrbom (USNM), B.

Brown (Zurqui project material) and M. Pollett (MHNM). It is only through the support of these institutions, along with their willingness to make material available to other researchers, that these types of studies can succeed. I would also like to thank A. Brunke (CNCI), Z. Soltes (HMNH) and N. Penny (CASC) for making digital photographs of type specimens available to me.

This study was funded by an NSERC Discovery Grant awarded to Dr. S. Marshall, and a PGS-D NSERC grant awarded to myself.

NOMENCLATURAL DISCLAIMER:

This thesis is not issued for public and permanent scientific record and for purposes of zoological nomenclature.

Hence it is not published within the meaning of the International Code of Zoological Nomenclature, 4th edition, article 8.2. New names and nomenclatural acts included in this thesis will enter in force only at the time of their publication in the relevant scientific periodicals iii

TABLE OF CONTENTS

ABSTRACT: ...... ii

ACKNOWLEDGEMENTS ...... iii

NOMENCLATURAL DISCLAIMER: ...... iii

List of Tables ...... vii

List of Figures ...... vii

CHAPTER 1 - INTRODUCTION ...... - 1 -

1.1 Biology and Natural History of the Archiceroptera genus complex ...... - 2 -

1.2 History of the Archiceroptera genus complex ...... - 2 -

1.3 Relationships with other Limosininae ...... - 3 -

1.4 Thesis objectives ...... - 3 -

CHAPTER 2 - MATERIALS AND METHODS ...... - 5 -

2.1 Label Information and Distribution Maps ...... - 5 -

2.2 Depositories of Material Examined ...... - 6 -

2.3 Compound Microscopy Photography and Illustration ...... - 6 -

2.4 Morphology ...... - 7 -

2.5 Figures ...... - 8 -

CHAPTER 3 – Relationships OF the Archiceroptera genus complex ...... - 16 -

3.1 Materials and Methods ...... - 16 -

3.2 RESULTS ...... - 20 -

3.3 DISCUSSION ...... - 21 -

3.4 Tables and Figures ...... - 25 -

CHAPTER 4 - PECTINOSINA, a new neotropical genus of Limosininae (Diptera: Sphaeroceridae) ...... - 31 -

4.1 Abstract ...... - 31 -

4.2 Introduction ...... - 31 -

4.3 Materials and Methods ...... - 31 -

4.4 Pectinosina gen. nov...... - 33 -

4.4.1 Species Descriptions ...... - 36 - iv

4.5 Chapter References ...... - 46 -

4.6 Pectinosina Figures ...... - 48 -

CHAPTER 5 – A revision of the genus Rudolfina Roháček (Sphaeroceridae: Limosininae) ...... - 54 -

5.1 Abstract ...... - 54 -

5.2 Introduction ...... - 54 -

5.3 Materials and Methods ...... - 57 -

5.4 Analysis ...... - 59 -

5.5 Results of Phylogenetic analysis ...... - 61 -

5.6 Rudolfina Roháček 1987 ...... - 63 -

5.7 Key to the New World Rudolfina...... - 65 -

5.8 Species descriptions (alphabetically organized) ...... - 69 -

5.9 Described species in other genera previously treated as Rudolfina ...... - 90 -

5.10 Discussion ...... - 91 -

5.11 References ...... - 92 -

5.12 Table List ...... - 94 -

5.13 Figure List ...... - 94 -

5.14 Figures ...... - 98 -

CHAPTER 6 – A Revision of Archiceroptera Papp 1977 ...... - 119 -

6.1 Abstract ...... - 119 -

6.2 Introduction ...... - 119 -

6.2.1 Biology ...... - 120 -

6.2.2 Related genera ...... - 120 -

6.3 Materials and Methods ...... - 120 -

6.4 Archiceroptera Papp 1977 ...... - 122 -

6.4.1 Diagnosis ...... - 122 -

6.4.2 Redescription ...... - 123 -

6.5 Phylogeny ...... - 125 -

6.5.1 Morphological ...... - 125 - v

6.5.2 Morphological characters and character states used in the phylogenetic analysis of Archiceroptera . - 126 -

6.5.3 Molecular sequences ...... - 129 -

6.5.2 Discussion ...... - 130 -

6.6 Key to the species of Archiceroptera Papp ...... - 132 -

6.7 Species Descriptions ...... - 141 -

6.7.1 Archiceroptera addenda species group ...... - 141 -

6.7.2 Archiceroptera mahukani species group ...... - 147 -

6.7.3 Archiceroptera ternum-species group ...... - 155 -

6.7.4 Archiceroptera brevivilla species subgroup ...... - 196 -

6.8 Discussion ...... - 207 -

6.9 Chapter References ...... - 208 -

6.10 Archiceroptera Tables and Figures ...... - 210 -

6.10.1 Archiceroptera addenda species group ...... - 218 -

6.10.2 Archiceroptera mahukani species group ...... - 224 -

6.10.3 Archiceroptera ‘ternum species group’ ...... - 229 -

6.10.4 Archiceroptera brevivilla species subgroup ...... - 262 -

6.10.5 Distribution maps ...... - 272 -

CHAPTER 7 – SUMMARY ...... - 275 -

CHAPTER 8 – References ...... - 276 -

Appendix 1 – Synopsis of Newly Described Sphaerocerdiae Species since the last Catalog Update...... - 283 -

A1.1 DESCRIBED LIMOSININAE SINCE LAST CATALOG UPDATE ...... - 284 -

A1.2 DESCRIBED COPROMYZINAE SINCE LAST CATALOG UPDATE ...... - 304 -

A1.3 DESCRIBED SPHAEROCERINAE SINCE LAST CATALOG UPDATE ...... - 305 -

A1.4 DESCRIBED ARCHIBORBORINAE SINCE LAST CATALOG UPDATE ...... - 306 -

A1.5 PENDING SPECIES DESCRIPTIONS ...... - 317 -

A1.6 References for Appendix 1 (citations for pending species descriptions are not currently included): ...... - 326 -

vi

LIST OF TABLES

Table 3.1. Morphological character states for the epandrial process group (EPG)...... - 25 -

Table 5.1. Character states used in the phylogenetic analysis of Rudolfina...... - 98 -

Table 6.1. Morphological character states for phylogenetic study of Archiceroptera...... - 210 -

LIST OF FIGURES

Figure 1.1. Anterolateral view of the dissected epandrium for four members of the Archiceroptera genus complex illustrating the epandrial process. A) Archiceroptera; B) Rudolfina; C) Archiceroptera (ternum-group); and D)

Pectinosina prominens...... - 4 -

Figure 2.1. Head chaetotaxy: lateral, Archiceroptera venezolana (Richards) (debu01077469), and dorsolateral, A. browni n. sp. (debu01077561)...... - 8 -

Figure 2.2. Thoracic chaetotaxy of Archiceroptera mahukani (debu00295088)...... - 9 -

Figure 2.3. Mid tibial chaetotaxy terminology (modified from Buck and Marshall 2009). Abbreviations as follows: ad – anterodorsal; p ad - predistal anterodorsal; d ad - distal anterodorsal; d aa - distal anteroapical; pd – posterodorsal; p pd - predistal posterodorsal; d pd - distal posterodorsal; d d- distal dorsal; d pa – distal posteroapical...... - 10 -

Figure 2.4. Wing of two members of the Archiceroptera genus complex. A) Archiceroptera venezolana

(debu00378968); B) Rudolfina exuberata (debu00276674)...... - 11 -

Figure 2.5. Wings of the Archiceroptera genus complex. A) Archiceroptera addenda (Extension) group

(debu00190199); B) Archiceroptera brevivilla group (debu00228460); C) enigmata group (slide mount, no number);

D) sororcula group (debu00258519); E) sororcula group (Hull, QC); F) Rudolfina digitata group (debu01086069). ..

...... - 12 -

Figure 2.6. Archiceroptera venezolana male terminalia: A) abdomen, ventral view; B) sternite 5–7, ventral view; C) epandrium, cercus and surstylus, caudal view; D) same, lateral view; E) surstylus, close up lateral; F) postgonite; lateral view; G) phallus, dorsal view; H) same, dorsolateral view; I) same, lateral view. A-I) from debu00373840 ......

...... - 13 -

Figure 2.7. Archiceroptera venezolana phallus. A) dorsal view; B) dorsolateral view; and C) lateral view. A-C) from debu00373840...... - 14 - vii

Figure 2.8. Rudolfina megepandria , female terminalia: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. (A-D from debu01086086)...... - 15 -

Figure 3.1. Maximum likelihood analysis of cytochrome c oxidase subunit I (COI) 5P (Part 2). Numbers at nodes are aBayes values. The leading codes given for each specimen are the unique identifiers within the BOLD database. The colour codes are as follows: pink = non-Limosininae; black = Limosininae excluding epandrial process group; blue

= epandrial process group excluding the Archiceroptera genus group; and red = Archiceroptera genus complex.

Vertical lines denote different parts of the Archiceroptera genus complex...... - 26 -

Figure 3.2. Maximum likelihood analysis of cytochrome c oxidase subunit I (COI) 5P (Part 1). Numbers at nodes are aBayes values. The leading codes given for each specimen are the unique identifiers within the BOLD database. The colour codes are as follows: pink = non-Limosininae; black = Limosininae excluding epandrial process group; and blue = epandrial process group excluding the Archiceroptera genus complex...... - 27 -

Figure 3.3. Maximum likelihood analysis of cytochrome c oxidase subunit I (COI) 5P for the genera belonging to the epandrial process group with aBayes support values for each node. The leading specimen codes are the unique identifiers within the BOLD database. Lines denote specimens belonging to the Archiceroptera genus complex: 1 =

Rudolfina; 2 = prominens group; 3 = enigmata and sororcula (in part) groups; 4 = Archiceroptera mahukani, ternum and brevivilla groups; 5 = sororcula group (in part); 6 = R. exuberata and Extension groups; and 7 = sororcula group (in part)...... - 28 -

Figure 3.5. Strict Consensus Tree (A) and Majority Rules Tree (B) from the morphological analysis of the epandrial process group...... - 29 -

Figure 3.6. Morphological phylogeny of the epandrial process group, including species groups of the Archiceroptera genus complex. One of 11 equal length trees...... - 30 -

Figure 4.1. Pectinosina prominens: A) head and thorax, lateral view; B) wing; C) left mid tibia, dorsal view, showing distinctive chaetotaxy (arrows indicating setae placement that separate Pectinosina from related genera)......

...... - 48 -

Figure 4.2. Male terminalia of P. carro: A) abdomen, ventral view; B) sternite 4–5 and anterior part of synsternite

6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, close up

viii lateral; F) postgonite; lateral view; G) distiphallus and basiphallus, dorsal view; H) same, dorsolateral view; I) same, lateral view. A-I) from debu01082094 ...... - 49 -

Figure 4.3. Pectinosina carro, female terminalia and spermathecae: A) terminal abdominal segments, dorsal view;

B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-

C) from debu01082100 and D) from debu01082102...... - 50 -

Figure 4.4. Pectinosina prominens, male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse portion of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, close up lateral; F) postgonite; lateral view; G) phallus, dorsal view; H) phallus, postgonite and phallapodeme, dorsolateral view; I) same, lateral view. A-I) from debu00287406...... - 51 -

Figure 4.5. Pectinosina prominens, female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-C) from debu00190483, D) from debu01082507...... - 52 -

Figure 4.6. Distribution of Pectinosina prominens and P. carro...... - 53 -

Figure 5.1. Rudolfina head and wing morphology: A) R. howdeni head (debu01086104); and B) R. exuberata wing

(debu00276674)...... - 98 -

Figure 5.2. Rudolfina tumida. A) male epandrium, cercus and surstylus, posterior view; B) same, lateral view.

Illustration and photograph from debu01086083...... - 99 -

Figure 5.3. Male morphology. A) sternite 5 (Rudolfina tumida, debu01086083); B) hypandrium (R. exuberata, debu00242299); C) postgonite, lateral view (R. cavernicola, debu01086077); D) phallus (including the basiphallus and distiphallus), postgonite and phallapodeme, lateral view (R. tumida, debu01086083)...... - 100 -

Figure 5.4. Rudolfina megepandria , female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086086)...... - 101 -

Figure 5.5. Strict consensus tree for the six trees obtained from traditional search (TNT)...... - 102 -

Figure 5.6. Majority Rules consensus tree from the 6 optimized trees retained from Traditional Search (TNT). - 102 -

Figure 5.7. Phylogeny of Rudolfina species. Character and character states refer to table 1. One of six equal length trees. Length=72, Ci=56. Ri=63...... - 103 -

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Figure 5.8. Rudolfina bucki, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus, dorsal view; E) phallus, postgonite and phallapodeme, dorsolateral view; F) same, lateral view. (A–F from debu01086239)...... - 104 -

Figure 5.9. Rudolfina exuberata, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral;

C) sternite 5; D) postgonite, lateral; E) phallus, dorsal view; F) same, dorsolateral view; G) same, lateral view. (A–G from debu00242299)...... - 105 -

Figure 5.10. Rudolfina exuberata, female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. A–C) from debu00242299; D) from debu00242286)...... - 106 -

Figure 5.11. Rudolfina howdeni, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus and postgonites; dorsal view; E) phallus and postgonite, lateral view; F) same, dorsolateral view. (Photos A–F from debu01086163)...... - 107 -

Figure 5.12. Rudolfina howdeni, female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu1086100)...... - 108 -

Figure 5.13. Rudolfina megepandria, male terminalia: A) epandrium, surstylus and cercus, lateral view; B) same, caudal view; C) sternites 4-7, ventral view; D) phallus and postgonite, dorsal view; E) phallus, postgonite and phallapodeme, dorsolateral view; F) same, dorsolateral view. (A–G from debu01086085)...... - 109 -

Figure 5.14. Rudolfina newtoni, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternites 4 and 5, ventral view; D) phallus, dorsal view; E) phallus, dorsolateral view; F) phallus and postgonite, lateral view. (A–E from debu01086234, F from debu01086234)...... - 110 -

Figure 5.15. Rudolfina newtoni , female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086226)...... - 111 -

Figure 5.16. Rudolfina pauca, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus, postgonites and phallapodeme, dorsal view; E) phallus, postgonites, hypandrium and phallapodeme, dorsolateral view; F) same, lateral view. (A–F from debu1086247)...... - 112 -

Figure 5.17. Rudolfina pauca, female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086258)...... - 113 -

x

Figure 5.18. Rudolfina pilosa, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus and ejaculatory apodeme, dorsal view; E) same, dorsolateral view; F) same, lateral view; G) postgonite, lateral view. (A–G from debu01086241)...... - 114 -

Figure 5.19. Rudolfina remiforma, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus and postgonites, dorsal view; E) phallus, postgonites and phallapodeme, dorsolateral view; F) same, lateral view. (A–F from debu01086286)...... - 115 -

Figure 5.20. Rudolfina remiforma, female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086288)...... - 116 -

Figure 5.21. Rudolfina tumida: A) sternite 5, close up ventral; B) phallus and postgonites, dorsal view; C) same, dorsolateral view. (A–C from debu01086083)...... - 117 -

Figure 5.22. Distribution of New World Rudolfina species: A) R. cavernicola, R. digitata, and R. tumida; B) R. bucki, R. megepandria, and R. howdeni; C) R. pauca, R. pilosa, R. newtoni, and R. remiforma; and D) R. exuberata. -

118 -

Figure 6.1. Strict consensus tree for Archiceroptera of 70 retained trees from Traditional Search (TNT). Bootstrap and Jackknife values > 50 are given above and below (respectively)...... - 211 -

Figure 6.2. Majority Rules tree for Archiceroptera from 70 trees from Traditional Search (TNT)...... - 212 -

Figure 6.3. Phylogeny of Archiceroptera. Tree selected from 70 equally parsimonious trees (Length = 225, Ci = 34,

Ri = 62)...... - 213 -

Figure 6.4. Maximum likelihood analysis of cytochrome c oxidase subunit I (COI) 5P for Archiceroptera with 3rd codon included (A) and excluded (B). Numbers at nodes are aBayes values. The leading codes for each specimen are the unique identifiers within the BOLD database...... - 214 -

Figure 6.5. Archiceroptera venezolana male terminalia: A) abdomen, ventral view; B) sternite 5, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, close up lateral; F) postgonite; lateral view; G) phallus, dorsal view; H) same, dorsolateral view; I) same, lateral view. A-I) from debu00373840 ......

...... - 215 -

Figure 6.6. Archiceroptera venezolana phallus. A) dorsal view; B) dorsolateral view; and C) lateral view. A-C) from debu00373840...... - 216 -

xi

Figure 6.7. Archiceroptera venezolana female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00373801...... - 217 -

Figure 6.8. Archiceroptera addenda male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior half of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) distiphallus, postgonite and phallapodeme, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00260830...... - 218 -

Figure 6.9. Archiceroptera addenda female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084477...... - 219 -

Figure 6.10. Archiceroptera crenulata male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) left surstylus; lateral view; F) distiphallus and postgonite, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-E) from debu01084494; F-H) from debu01084488...... - 220 -

Figure 6.11. Archiceroptera crenulata female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084493...... - 221 -

Figure 6.12. Archiceroptera triclavus male terminalia: A) abdomen, ventral view; B) posterior margin of sternite 5, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus; lateral view; F) distiphallus and postgonite, dorsal view; G) distiphallus, phallapodeme (in part) and postgonite, dorsolateral view;

H) same, lateral view. A-H) from debu01084483...... - 222 -

Figure 6.13. Archiceroptera triclavus female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00378580...... - 223 -

Figure 6.14. Archiceroptera browni (holotype; debu01077561). A) habitus, lateral view; B) habitus, dorsal view; C) head, dorsolateral view...... - 224 -

Figure 6.15. Archiceroptera browni male terminalia: A) abdomen, ventral view; B) sternites 5–8, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, close up lateral; F) postgonite; xii lateral view; G) phallus, postgonite and phallapodeme, dorsal view; H) phallus, dorsolateral view; I) same, lateral view. A-I) from holotype (debu01077561)...... - 225 -

Figure 6.16. Archiceroptera mahukani (holotype). A) habitus, dorsal view; B) habitus, lateral view; C) head, dorsolateral view...... - 226 -

Figure 6.17. Archiceroptera mahukani female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00295088...... - 227 -

Figure 6.18. Archiceroptera venezolana (holotype). A) head, anterior view; B) habitus, lateral view; C) habitus, dorsal view...... - 228 -

Figure 6.19. Archiceroptera adamas male terminalia: A) abdomen, ventral view; B) sternite 5 (with fungal thalli) and synsternite 6+7 ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00275292...... - 229 -

Figure 6.20. Archiceroptera adamas female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00379812...... - 230 -

Figure 6.21. Archiceroptera barberi male terminalia: A) abdomen, ventral view; B) posterior part of sternite 4, sternite 5, and transverse portion of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00107565...... - 231 -

Figure 6.22. Archiceroptera barberi female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00107520...... - 232 -

Figure 6.23. Archiceroptera basilia male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior parts of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) phallus, postgonite and phallapodeme, dorsal view; F) same, dorsolateral view; G) same, lateral view. A-G) from debu00140620...... - 233 -

xiii

Figure 6.24. Archiceroptera basilia female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00140638...... - 234 -

Figure 6.25. Archiceroptera bilobata male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior portion of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H from debu01084831...... - 235 -

Figure 6.26. Archiceroptera bilobata female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084833...... - 236 -

Figure 6.27. Archiceroptera bisetosus male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-D) from debu00189533;

E-H) from debu00189536...... - 237 -

Figure 6.28. Archiceroptera bisetosus female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00189555...... - 238 -

Figure 6.29. Archiceroptera caliga male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01085358; ...... - 239 -

Figure 6.30. Archiceroptera caliga female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085364...... - 240 -

Figure 6.31. Archiceroptera calligraphia male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite

6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01085530. .. - 241 -

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Figure 6.32. Archiceroptera calligraphia female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085517...... - 242 -

Figure 6.33. Archiceroptera cobolorum male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01086488. ..

...... - 243 -

Figure 6.34. Archiceroptera cobolorum female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01086522...... - 244 -

Figure 6.35. Archiceroptera dolabra male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00385443...... - 245 -

Figure 6.36. Archiceroptera maniba male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00378585. ..

...... - 246 -

Figure 6.37. Archiceroptera maniba female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00378572...... - 247 -

Figure 6.38. Archiceroptera masoni male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01085698. ..

...... - 248 -

Figure 6.39. Archiceroptera masoni female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085701...... - 249 -

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Figure 6.40. Archiceroptera megacercus male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01084910...... - 250 -

Figure 6.41. Archiceroptera megacercus female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084919...... - 251 -

Figure 6.42. Archiceroptera mexicorona: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5 and anterior part of synsternite 6+7, ventral view; D) left surstylus, lateral view; E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01085710......

...... - 252 -

Figure 6.43. Archiceroptera mexicorona female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085713...... - 253 -

Figure 6.44. Archiceroptera mitarakai male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) distiphallus and postgonite, dorsal view; F) same, dorsolateral view; G) same, lateral view. A-G) from debu00394580...... - 254 -

Figure 6.45. Archiceroptera paracercus, male terminalia male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, posterolateral view; F) postgonite; lateral view; G) distiphallus, dorsal view; H) same, dorsolateral view; I) same, lateral view. A-I) from debu00260795...... - 255 -

Figure 6.46. Archiceroptera pussula male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A) from debu00196122, B-H) from debu01085674...... - 256 -

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Figure 6.47. Archiceroptera pussula female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085659...... - 257 -

Figure 6.48. Archiceroptera ternum male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00132380...... - 258 -

Figure 6.49. Archiceroptera ternum female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-C) from debu00260557; D) from debu00132367...... - 259 -

Figure 6.50. Archiceroptera uncinata male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00382406...... - 260 -

Figure 6.51. Archiceroptera uncinata female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084712...... - 261 -

Figure 6.52. Archiceroptera braziliensis male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite

6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite, close up lateral; F) phallus,dorsal view; G) phallus, dorsolateral view; H) phallus, lateral view. A-H) from debu01088400......

...... - 262 -

Figure 6.53. Archiceroptera braziliensis female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01088403...... - 263 -

Figure 6.54. Archiceroptera brevivilla male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior portion of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite, close up lateral; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01088231...... - 264 -

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Figure 6.55. Archiceroptera brevivilla female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01086593...... - 265 -

Figure 6.56. Archiceroptera curvivilla male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite, close up lateral; F) phallus,dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01088418...... - 266 -

Figure 6.57. Archiceroptera curvivilla female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01088446...... - 267 -

Figure 6.58. Archiceroptera llama male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) phallus and postgonite, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-G) from debu00386692...... - 268 -

Figure 6.59. Archiceroptera llama female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00386665...... - 269 -

Figure 6.60. Archiceroptera megavilla male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite, close up lateral; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. (A and C-H from debu01088807, B from debu01086594)...... - 270 -

Figure 6.61. Archiceroptera megavilla female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. (A and C from debu01086596; B and D from debu01086595)...... - 271 -

Figure 6.62. Distribution of Archiceroptera species: A) the addenda species group (A. crenulata, A. addenda, and A. triclavus); B) Archiceroptera s. str. (A. browni, A. mahukani, and A. venezolana); C) A. maniba, A. adamas, and A. calligraphia; and D) A. masoni, A. mexicorona, A. uncinata, and A. cobolorum...... - 272 -

Figure 6.63. Distribution of Archiceroptera species: A) A. megacercus and A. paracercus; B) A. caliga, A. pussula,

A. mitarakai, and A. dolabra; C) A. basilia, A. bisetosus, and A. bilobata; and D) A. ternum...... - 273 - xviii

Figure 6.64. Distribution of Archiceroptera species: A) A. barberi; B) A. brevivilla, A. curvivilla, and A. llama; C) A. braziliensis and A. megavilla; and D) all Archiceroptera specimens...... - 274 -

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CHAPTER 1 - INTRODUCTION

The Sphaeroceridae is currently divided into six subfamilies, of which the Limosininae is the most diverse with about 150 described genera and 1650 described species worldwide [based on Marshall et al. 2011 plus publications since 2011 (see Appendix 1)]. Species of Limosininae can be found on every continent with the exception of Antarctica and are associated with a variety of decaying organic substances, such as carrion, dung, and decaying vegetation. Limosinine diversity is highest in the tropics, especially in the Neotropics, and many groups remain to be adequately studied. One such group is currently treated as the Archiceroptera genus complex (Marshall and Buck 2010). This complex includes two described genera, Archiceroptera Papp and Rudolfina Roháček, several unplaced species groups, seven described species (two Archiceroptera, and five Rudolfina) and an estimated 50-90 undescribed species.

The Archiceroptera genus complex belongs to a group of largely Neotropical genera characterized by a unique finger-like process that extends medially from the right margin of the epandrium (Marshall and Cui 2005,

Luk and Marshall 2014). Species within this epandrial process group all have the following combination of characters: 3 or more equal interfrontal setae; scutellum with only 2 pairs of setae; cell cup absent; alula narrow with straight hind margin; mid tibia with at least 1 anterodorsal seta on basal 1/3 (secondarily lost in Aptilotella Duda), 1 anterodorsal seta at midlength, and 1 anterodorsal and 1 posterodorsal seta on apical 3/4; and mid tibia with apical ventral seta. This combination of characters will separate the "epandrial process" group from almost all other

Limosininae genera with the exception of species that fall into the “boliviensis” group, an unplaced species group in which males lack an epandrial process. Marshall and Buck (2010) divided the complex into several species groups, noting that some probably deserve generic rank. These groups are the enigmata group, the brevivilla-group, the sororcula group (including the discalis group), the addenda group, the ternum-group, the prominens group, and the exuberata group.

- 1 -

1.1 Biology and Natural History of the Archiceroptera genus complex

Sphaerocerids are associated with a variety of decaying organic material, such as dung, carrion, fruits, and leaf litter, where the larvae are microbial grazers (Marshall and Richards 1987, Roháček et al. 2001). Members of the Archiceroptera genus complex have been collected in a wide variety of habitats, mostly using dung and carrion baits, pan traps, and flight intercept traps, and little is known beyond what can be extrapolated from data labels.

Rudolfina cavernicola Marshall & Fitzgerald is thus far known only from caves in the southwestern United States

(Marshall & Fitzgerald 1997), while R. digitata is associated with high alpine meadows (Marshall 1991).

Not much is known about the behaviour of these or most other Sphaeroceridae, but label data for one species of the prominens species group provides notes indicating the flies were riding or following dung beetles

(Coleoptera: Scarabaeidae, Scarabaeinae; see discussion for Pectinosina prominens). Kleptoparasitism of scarab dung balls by sphaerocerids is documented in other genera, and was reviewed by Sivinski et al. (1999), but this behaviour has not been recorded previously in the Archiceroptera genus complex.

1.2 History of the Archiceroptera genus complex

Papp (1977) described Archiceroptera based on two species: the newly described A. mahunkai Papp and A. venezolana (Richards), which Papp transferred from Ceroptera. Papp separated Archiceroptera from other

Limosininae on the basis of a single large costagial seta (as found in Ceroptera), unique leg chaetotaxy, and several small setulae near the basal pair of scutellar setae. Roháček (1982, 1983) transferred Limosina rozkosnyi Roháček to a new genus, Rudolfia, based on a number of autapomorphic characters (a single enlarged costagial seta, weakly sclerotized distiphallus, presence of small sclerites below gonopore [phallotrema], epandrium with lateral row of long setae and strikingly developed cerci, additional small sclerites below gonopore [phallotrema], reduced female sternite 8, female cerci of unusual “thornlike shape” and fused with the epiproct). Roháček (1987) later renamed

Rudolfia as Rudolfina, as Rudolfia was preoccupied by a copepod crustacean. Since then, two new North American

Rudolfina species have been described (Marshall 1991, Marshall & Fitzgerald 1997) and two species have been transferred into the genus (Pitkin 1989, Roháček et al. 2001). A chapter in an unpublished thesis by S.A. Marshall

(1982) considered the material available at the time, but was not published because further material, especially from

- 2 - the Neotropics, was required to clarify generic limits of Rudolfina and Archiceroptera. Marshall and Buck (2010) summarized ongoing work on the complex, continued by the present author as a doctoral project.

1.3 Relationships with other Limosininae

Marshall and Roháček (1986) considered Rudolfina as a possible outgroup for Trachyopella but it is now considered that the Archiceroptera genus complex belongs to a larger clade of mostly Neotropical genera diagnosed by the presence of a small finger-like process that extends from the right side of the epandrium to the hypandrium

(Fig. 1.1; see also Marshall and Cui 2005, Fig. 1), which is considered a synapomorphy for the group. This group

(referred to here as the epandrial process group, or EPG) also includes Robustagramma, Bromeloecia,

Pterogramma, Aptilotella, and Bitheca, but excludes Trachyopella. The relationships between this clade and other groups within the Limosininae are presently unclear.

1.4 Thesis objectives

The major focus of this thesis was to study the species diversity of the species groups previously included within the Archiceroptera genus complex and place them in a broader phylogenetic context within the subfamily

Limosininae. The generic limits of Archiceroptera and Rudolfina were both redefined and revised, and their relationships with each other and related genera were examined using both molecular and morphological character sets. The remaining species groups of the Archiceroptera genus complex were also placed into a phylogenetic framework within the broader EPG clade and one new genus was described for a previously described species that was excluded from Rudolfina and Archiceroptera.

- 3 -

Figure 1.1. Anterolateral view of the dissected epandrium for four members of the Archiceroptera genus complex illustrating the epandrial process. A) Archiceroptera; B) Rudolfina; C) Archiceroptera (ternum- group); and D) Pectinosina prominens.

- 4 -

CHAPTER 2 - MATERIALS AND METHODS

Over 8000 specimens of the Archiceroptera genus complex were studied, with the major portion (> 95%) of that material currently deposited at the University of Guelph Insect Collection, the largest collection of New

World Sphaeroceridae. Some material had already been sorted to the Archiceroptera genus group (“Rudolfina s.l.”) prior to this study, but another 20+ drawers of unidentified New World Limosininae were also examined for additional material. Further material came from two inquiries for bulk identifications of Sphaeroceridae. The first inquiry was from the Zurqui All Diptera Biodiversity Inventory (ZADBI) project, which documented the fly diversity of a single plot in Costa Rica that was later expanded to include two other sites. Of the ~7000 sphaerocerid specimens identified to genus or species for this project, only ~20-30 belonged to the Archiceroptera genus complex but the quality of the specimens allowed sequencing of several species. The other request was from a smaller collection effort in Mitaraka, French Guiana by Dr. M. Pollet. Approximately 2000 specimens were made available and provided access to material of previously unknown species along with new distributional data for several other species.

2.1 Label Information and Distribution Maps

Label data was presented in a consistent manner, not verbatim from the labels; in a few cases obvious spelling errors were corrected. Shortforms or abbreviations used on the labels were normally interpreted and given in full.

Geographical coordinates were given if present on the original label. Specimens were given unique identifiers and their collection data was captured within the BIOTA database (Colwell 2012); the specimen data will ultimately be hosted on the Canadensys website, but unique identifiers were not repeated in the text except for holotypes or for imaged specimens. Collection data for paratypes and other specimens examined were organized alphabetically by country, state/province, and locality name. Species distribution maps were generated using SimpleMappr

(Shorthouse 2010). Geographic relief maps were used to help identify regions of occurrence and potential barriers to species limits, such as mountain ranges.

- 5 -

2.2 Depositories of Material Examined

Depository abbreviations are as follows: AMNH (American Museum of Natural History, New York, New York);

DEBU (School of Environmental Sciences, University of Guelph, Guelph, Ontario, Canada); CNCI (Canadian

National Collection, Ottawa, Ontario, Canada), FMNH (Field Museum of Natural History, Chicago, Illinois); INBC

(Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica); MHNM (Museum National d’Histoire Naturelle, Paris, France); MIZA (Museo del Instituto de Zoología Agrícola Francisco Fernández Yépez;

Universidad Central de Venezuela, Maracay, Venezuela); MUSM (Museo de Historia Natural, Universidad

Nacional Mayor de San Marcos, Lima, Peru); MZSP (Museu de Zoologia, Universidade de São Paulo, São Paulo,

São Paulo, Brazil); NHMW (Vienna Museum of Natural History, Vienna, Austria), QCAZ (Departamento de

Biología, Pontífica Universidad Católica del Ecuador, Quito, Ecuador); ROME (Royal Ontario Museum, Toronto,

Ontario), UASC (Museo de Historia Natural Noel Kempff Mercado, Santa Cruz de la Sierra, Bolivia); UNAM

(Universidad Nacional Autónoma de México, Mexico City, Mexico); USNM (United States National Museum of

Natural History, Smithsonian Institution, Washington, D.C., U.S.A.); UVGC (Colleción de Artrópodos, Universidad de Valle de Guatemala, Guatemala City, Guatemala). Deposition of type material was determined by the source of the material and/or repatriation agreements with the country of origin. Material is deposited in DEBU unless otherwise noted.

2.3 Compound Microscopy Photography and Illustration

A Canon PowerShot S5IS with an ocular mount paired with a Leitz Laborlux 11 compound microscope was used to capture images of male and female genitalia. Series of images were combined using Zerene Stacker version 1.04 (Zerene Systems LLC, Richland, WA, U.S.A.) with the DMax algorithm and manually edited.

Additional editing with Adobe Photoshop CS5 (Adobe, San Jose, California, U.S.A.) was done to generate standardized plates. A standard complement of genitalic images is given for all species, except in cases where structures were broken or otherwise unavailable. For males the following standardized photos were provided: a ventral view of the terminalia distal to sternite 4; a close up of sternite 5 (usually including the anterior portion of synsternite 6+7); posterior and lateral views of the cercus, surstylus and epandrium; a close up of either the surstylus or postgonite (if not visible in the other images); and dorsal, dorsolateral and lateral views of the internal genitalia to

- 6 - show the various sclerites and membranes. Photographs of females included a dorsal, lateral and ventral view of the terminalia and a composite image of the spermatheca. In a few cases, the photographs were accompanied by previously drawn illustrations to provide an atlas of the various structures. Descriptions refer to structures in the standard orientation shown in the figures.

2.4 Morphology

External morphological terminology follows Cumming and Wood (2010), and internal morphological terminology follows Smith and Marshall (2004), with the following modifications. Seta(e) and setula(e) are the large and small socketed macrotrichia, respectively. The M1 and CuA1 stubveins refer to the portion of each vein that extends beyond cell dm-cu. M1 is considered traceable to the wing margin when the melanistic distal remnants of M1

(referred to as “pseudoveins” by some authors) continue to the wing margin (best observed with pale background).

The spermathecal stem is used to describe the individual duct of each of the paired spermatheca. Figures 2.1–2.2 illustrate the head and thoracic setae and setulae. Figure 2.3 illustrates the mid tibial chaetotaxy. Figures 2.4–2.5 illustrate the wings of various species groups in the Archiceroptera genus group; Figure 2.6-2.7 illustrates the male terminalia. Figure 2.8 illustrates the female terminalia.

Measurements and ratios

The measurements and ratios are based on a minimum of 10 specimens (if available), including a variety of collections and localities. Body length should be treated as an approximation as different preservation and drying methods can alter the length of membranous tissue between sclerotized plates. The following measurements and ratios are used.

Body Length: distance from the front of the head (excluding antennae) to the tip of the abdomen.

Eye height: the largest vertical distance from the bottom of the compound eye, as observed in lateral view.

Genal height: distance from the most ventral portion of the compound eye to the margin of the gena (usually the narrowest vertical distance), as observed in lateral view.

Costal sectors 2 and 3: sectors of the costal vein between radial vein apices.

- 7 -

2.5 Figures

Figure 2.1. Head chaetotaxy: lateral, Archiceroptera venezolana (Richards) (debu01077469), and dorsolateral,

A. browni n. sp. (debu01077561).

- 8 -

Figure 2.2. Thoracic chaetotaxy of Archiceroptera mahukani (debu00295088).

- 9 -

Figure 2.3. Mid tibial chaetotaxy terminology (modified from Buck and Marshall 2009). Abbreviations as follows: ad – anterodorsal; p ad - predistal anterodorsal; d ad - distal anterodorsal; d aa - distal anteroapical; pd – posterodorsal; p pd - predistal posterodorsal; d pd - distal posterodorsal; d d- distal dorsal; d pa – distal posteroapical.

- 10 -

Figure 2.4. Wing of two members of the Archiceroptera genus complex. A) Archiceroptera venezolana

(debu00378968); B) Rudolfina exuberata (debu00276674).

- 11 -

Figure 2.5. Wings of the Archiceroptera genus complex. A) Archiceroptera addenda (Extension) group

(debu00190199); B) Archiceroptera brevivilla group (debu00228460); C) enigmata group (slide mount, no number); D) sororcula group (debu00258519); E) sororcula group (Hull, QC); F) Rudolfina digitata group

(debu01086069).

- 12 -

Figure 2.6. Archiceroptera venezolana male terminalia: A) abdomen, ventral view; B) sternite 5–7, ventral view; C) epandrium, cercus and surstylus, caudal view; D) same, lateral view; E) surstylus, close up lateral;

F) postgonite; lateral view; G) phallus, dorsal view; H) same, dorsolateral view; I) same, lateral view. A-I) from debu00373840.

- 13 -

Figure 2.7. Archiceroptera venezolana phallus. A) dorsal view; B) dorsolateral view; and C) lateral view. A-C) from debu00373840.

- 14 -

Figure 2.8. Rudolfina megepandria , female terminalia: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae.

(A-D from debu01086086).

- 15 -

CHAPTER 3 – RELATIONSHIPS OF THE ARCHICEROPTERA GENUS COMPLEX

The epandrial process group (EPG), the group of genera including the Archiceroptera genus complex, is currently diagnosed by the finger-like process extending medially from the right anteroventral angle of the epandrium along the posterior margin of sternite 8 (Marshall and Cui 2005, Smith and Marshall 2004). This process is otherwise unknown in the Limosininae and this unique structure is considered a synapomorphy for the EPG.

Although this process is usually finger like, Yau and Marshall (2017) found that the epandrial process was variable within the genus Bromeloecia; in some species it forms a bridge from the right to the left side of the epandrium while in other species it is almost completely absent. The structural variability of the epandrial process within

Bromeloecia, a genus strongly supported by independent characters, suggests that it may be subject to rapid change.

We treat it here as uniquely derived in the EPG group as this structure is known only from New World taxa and is correlated with both mid tibial chaetotaxy and wing venation development.

Molecular and morphological character sets were examined to provide the first phylogeny of the

EPG. A preliminary molecular analysis of the Sphaeroceridae using five prime region of cytochrome c oxidase I

(COI 5P) was done to examine the support for a monophyletic EPG and what, if any, relationships there were with other limosinine genera. A narrower analysis of COI 5P for species within the EPG was done to examine the relationships within the group. Finally, a morphological analysis of the EPG was done to establish the relationships between the described genera and unplaced species groups. The entire EPG was considered as the relationships between the groups within the Archiceroptera genus complex with the rest of the EPG was unclear.

3.1 Materials and Methods

3.1.1 Molecular Analysis

Available sequences from all genera of the Limosininae were gathered from BOLD systems

(http://www.boldsystems.org), including public data. Exemplar specimens for each group within the Archiceroptera genus complex were submitted, if adequate material was available, to the Biodiversity Institute of Ontario for extraction, amplification and sequencing. Recently collected material from within the past 10 years was preferentially used, but generally the material available was of limited value for sequencing because it was taken in pan traps, and often deteriorated in pan trap fluid prior to storage in alcohol. Sampling methods that collected - 16 - specimens directly into alcohol (e.g., Malaise traps) generally yield better material for sequencing but relatively few specimens in the EPG are taken by these methods. Additional identified sequence data was obtained through examination and the identification of digital images of sequenced but unidentified Limosininae. A dataset DOI of the sequences used in these analyses has been requested and will be provided if future publication of this work; a

BOLD dataset of all the sequences is available online at www.boldsystems.org as dataset DH-SPHSMP.

Sequence data was imported from BOLD into Mesquite (Maddison and Maddison 2017) and aligned by hand. In total, 300 sequences, including representatives from the Archiborborinae, Copromyzinae, Sphaerocerinae,

Homalomitrinae and Limosininae were included in the broader analysis. In the narrower EPG, a total of 133 sequences were included, including all described genera except Aptilotella, which did not have sequence data available. Sequences less than 600bp were not included to maximize comparable base pair data, with the exception of four Archiceroptera genus group sequences in the EPG analysis. The sequences were exported into PhyML 3.0

(Guindon et al. 2010) for maximum likelihood analysis (GTR+G+I substitution model and SPR tree improvement; node support used aBayes fast likelihood-based support).

3.1.2 Morphological Analysis

A character matrix (Table 1) of 42 characters was generated for EPG genera using Mesquite (version 3.10;

Maddison and Maddison 2011), and exported for analysis in TNT (Goloboff et al. 2008). A Traditional Search with

10 random seeds and 5000 replications using the tree bisection reconnection (TBR) swapping algorithm was used for the analysis. Bootstrap and Jackknife values were calculated with 100 replications and reported for values larger than 50. Trees were optimized in WINCLADA (Nixon 2002).

There is no established outgroup to the EPG. Papp (1977) considered Archiceroptera as part of the

Ceroptera genus group (sensu Papp 2008). Archiceroptera females do not have the telescopic terminalia, nor does either sex have reduced abdominal sclerotization, or the enlarged puvilli and claws used by Papp (1977) to define the group. The remaining characters Papp (1977) used are either ground plan for the Limosininae (3 spermathecae), derived but widely occurring in the Limosininae (e.g., cup absent, tibial spur), or are unclear in his diagnosis (leg chaetotaxy). In the absence of any putative synapomorphies between Archiceroptera and Ceroptera + Ceroptella, we do not consider Ceroptera or Ceroptella as closely related to the EPG. Trachyopella was also suggested as a possible sister group to Bromeloecia or Rudolfina before the EPG group was recognized (Marshall 1983b, Roháček - 17 - and Marshall 1986) and is now considered a sister group to Thoracochaeta (Marshall and Roháček 2000). In the absence of an established outgroup, New World limosinine genera were compared with a generalized EPG ancestor and two possible outgroups, Thoracochaeta and Archicollinella. Archicollinella shares with a generalized EPG ancestor in having three strong anterodorsal setae (one basal, one at the midlength and one apically) and a strong posterodorsal seta on the mid tibia, the male mid tibia has an apical setal comb, and the male cercus is basally free from the epandrium. The setal orientation found in Archicollinella and the EPG is not commonly found in other

New World Limosininae (many genera have lost the anterodorsal seta near the midlength). Thoracochaeta shares with the hypothetical ancestor a weakly bilobed surstylus and a distally broadened distiphallus; the mid tibial chaetotaxy of Thoracochaeta is consistent with the EPG but it is much more variable compared with Archicollinella.

Both Thoracochaeta and Archicollinella also have a row of inclinate orbital setulae, which occurs in several but not all EPG (apparently reduced or secondarily lost in these groups).

List of characters used in the morphological analysis of the EPG

The following 42 morphological characters are organized by body region and sex. Character states were polarized using Thoracochaeta and Archicollinella as outgroups. Multistate characters that had clear linear transformation series were treated as ordered (characters 12, 42) and are noted with an asterisk (*).

CHARACTER STATES

Head

1. Frons - orbital setae: (0) 2 present, (1) 1 present.

2. Frons - inclinate orbital setulae: (0) 2 or more present, (1) absent, or only 1 small anterior pair present.

Thorax

3. Dorsocentral setae: (0) 2 or more pairs of dorsocentral setae, (1) 1 pair of dorsocentral seta.

4. Scutellum – (0) no extra scutellar setae present; (1) extra scutellar seta present

Wing

5. Costagial seta development: (0) pair of equal or subequal setae present; (1) one seta distinctly enlarged.

6. R4+5 apical morphology: (0) apically curved towards costa; (1) apically straight.

7. Costal termination: (0) ending at apex of R4+5; (1) extending short distance beyond R4+5. - 18 -

8. M1 - Distance between r-m and dm-cu compared to the length of dm-cu: (0) > 3.0X, (1) 2.0-3.0X, (2) <

2.0X.

9. M1 termination: (0) traceable to costal margin; (1) terminating on wing disk.

Legs

10. Mid tibia – basal seta placement: (0) anterodorsal, (1) dorsal.

11. Mid tibia – Medial anterodorsal seta cluster: (0) 2 or more present; (1) one present.

12. *Mid tibia – Medial posterodorsal seta cluster: (0) no setae present; (1) 1 seta present; (2) 2 or more setae

present.

13. Mid tibia – Predistal anterodorsal seta: (0) present, (1) absent.

14. Mid tibia – Predistal dorsal seta: (0) present, (1) absent.

15. Mid tibia – Apical posterodorsal seta: (0) present, (1) absent.

16. Mid basitarsus – basal setae: (0) present, (1) absent.

17. Mid tibia ventral seta present in male: (0) present, (1) absent.

18. Mid tibia with midventral seta present in female: (0) present, (1) absent.

19. Male mid tibia with apical ventral comb: (0) absent; (1) present

20. Male mid femur with basoventral setal cluster: (0) present, (1) absent.

21. Hind femur with preapical seta: (0) present, (1) absent

Male Abdomen

22. Sternite 5 – additional posterior sclerite: (0) absent; (1) present.

23. Epandrial process: (0) absent; (1) present.

24. Epandrial-cercus articulation: (0) free, articulating; (1) fused or rigidly abutted to epandrium.

25. Cercus – projecting process: (0) absent; (1) present.

26. Surstylus – anterior lobe: (0) absent or extremely reduced; (1) present, well developed.

27. Surstylus – anterior lobe morphology: (0) regular, not laminate; (1) laminate.

28. Epiphallus: (0) absent; (1) present.

Female Abdomen

29. Terminalia telescopic: (0) no; (1) yes.

30. Tergite 7 posterolaterally: (0) free, (1) fused. - 19 -

31. Tergite 8 – posterolateral corner: (0) rounded, (1) acute, projecting.

32. Tergite 8 – medial part: (0) absent; (1) present.

33. Epiproct – sclerotization: (0) evenly sclerotized; (1) medially weakened

34. Epiproct - development: (0) well developed, (1) reduced.

35. Cercus – dorsal chaetotaxy: (0) setulose, pad-like, (1) elongate, glabrous.

36. Cercus – apical seta morphology: (0) regular, (1) flattened.

37. Cercus – apical chaetotaxy: (0) apical and preapical setae equal in length; (1) apical seta distinctly longer.

38. Cercus – fusion with epiproct: (0) not fused; (1) fused.

39. Sternite 8: (0) present, entire; (1) medially weakened or divided.

40. Sternite 8 – spinulose plates: (0) absent; (1) present.

41. Spermatheca – morphology: (0) ovoid or barrel-shaped, (1) disc-shaped, (2) bilobed, (3) cup-shaped.

42. *Spermathecal duct – length of stems relative to length of spermatheca: (0) elongate, > length of

spermatheca; (1) duct shorter than spermathecal length, but longer than ½ spermathecal length; (2) duct <

1/2 spermathecal length.

3.2 RESULTS

3.2.1 Specimen Sequencing

Of the 48 specimens submitted for sequencing, only 20 (41.7%) returned sequences, and only 12 (25%) were greater than 500 base pairs. Specimens less than 5 years old returned proportionally more sequences, but no substantial differences were noticed with the age of specimens for sequences of greater than 500 base pairs.

Collection method appeared to have an impact on the sequence success and quality, with Malaise traps providing the largest bp/sequence, and pan traps having the lowest bp/sequence. The difference in the quality and number of returned sequences between the collection methods is likely a result of the medium that the flies were collected into.

Most Malaise traps samples were collected directly into alcohol, while most pan trap samples were collected using soapy water and later transferred to alcohol.

- 20 -

3.2.2 Maximum-likelihood Analyses

Figures 3.1–3.2 and 3.3 show the maximum likelihood trees for the broad (sphaerocerid) and narrow (EPG) analysis, respectively. In the broad analysis, there was poor resolution of the subfamilies. In the broad analysis, the

EPG, the Archiceroptera genus complex and most limosinine genera were not recovered as monophyletic suggesting a limitation for the use of COI in Sphaeroceridae higher phylogeny. A subsequent analysis that excluded the third codons provided a similar tree. As such, no insight into a possible sister group for the EPG was available. Within the narrower analysis, Pterogramma sequences provided the most well-sampled taxon.

Within the Archiceroptera genus complex, Archiceroptera ternum-group and the A. brevivilla-group were recovered together with strong support in both analyses. Rudolfina was not recovered as monophyletic in either analysis; R. rozkosnyi and R. digitata were recovered together, while R. exuberata was consistently recovered with

[ternum-group + brevivilla-group] in the broad analysis and with the Extension group in the narrower (EPG) analysis. The remaining groups were recovered with both EPG and other limosinine genera in the broader analyses.

The prominens group was recovered as sister to [Archiceroptera Extension-group + Robustagramma] +

Bromeloecia in the EPG (narrower) study, or Pterogramma (in part) in the broader analysis. The enigmata and sororcula groups were largely recovered with the prominens group in the narrower (EPG) analysis.

3.2.3 Morphological Analysis

A Traditional Search (TNT) with 5000 replications and 10 random seeds using TBR found 11 trees (length

87; Ci = 50 Ri = 66) out of 5,553,331 arrangements, summarized here as a strict consensus tree and majority rules tree (Fig. 3.4). Bootstrap and Jackknife values > 50 are given on the strict consensus tree. Characters were optimized on one tree selected from the eleven equally parsimonious trees (Fig. 3.5). This tree was selected from the other ten trees by excluding both outgroups (i.e. Thoracochaeta and Archicollinella) to provide a monophyletic EPG and with the sororcula and enigmata groups as sister groups based on shared male genitalic characters.

3.3 DISCUSSION

Both molecular and morphological datasets did not recover the Archiceroptera genus complex as a monophyletic group. The EPG was not recovered as monophyletic in the broader limosinine analysis but this apparent paraphyly may have resulted from the weak phylogenetic signal within COI. Both Trunz et al. (2016) and - 21 -

Ekrem et al. (2016) found poor phylogenetic signal within COI compared to other genes, and Trunz et al. (2016) suggested that COI is not informative for groups that have diverged more than 4 mya. The oldest known fossil

Limosininae are from Dominican amber (see Marshall et al. 1999), dated between 13-20 mya and belong to extant genera, so it is very possible that many limosinine genera or generic groups will not be recovered as monophyletic using COI data. The use of COI for understanding species relationships within a single genus will be examined further with Archiceroptera (Chapter 6). The relatively small taxon sampling size within both the broader

Limosininae and within the EPG combined with the low quality of the EPG material for molecular study may have further limited the recovery of a monophyletic EPG and it is important that future studies try to include taxa that are not represented in the current dataset. However, Rokas and Carroll (2005) suggest that an increase in the number of genes is more helpful in resolving phylogenies than the addition of further taxa and future studies will need to obtain suitably preserved material to test this within the Limosininae.

Within the Archiceroptera genus group, Archiceroptera mahukani group and Rudolfina were recovered in the morphological analysis (along with the molecular analysis) as distinct genera that were more closely related to other EPG genera than they were to each other. The modified spinose female cercus that originally confused the two genera is a convergence that likely reflects a common ovipositing behaviour in the females. The recovery of

Archiceroptera mahukani group + [ternum-group + brevivilla-group] was consistent in both analyses, but in the molecular data it was somewhat confounded by the recovery of R. exuberata and Pterogramma sequences between

Archiceroptera mahukani group and [ternum-group + brevivilla-group]. Rudolfina exuberata is considered to be a derived species within Rudolfina with strong morphological support for its treatment there (see Chapter 5). The recovery of these two taxa within Archiceroptera mahukani group + ternum-group + brevivilla-group may be due to long branch attraction. The treatment of the ternum-group and brevivilla-group together was supported by both molecular and morphological analyses, although the molecular analysis recovered the brevivilla-group as a subgroup within the ternum-group. The posterolateral fusion of female tergite 7 to tergite 8 and the presence of dorsal basal seta on the mid tibia are considered here as synapomorphies for Archiceroptera + ternum-group + brevivilla-group.

The Extension group is treated here as a basal lineage of Archiceroptera, supported by four female abdominal characters: the teardrop-shaped female cercus, the apically flattened seta on the cercus, the medially weakened/divided epiproct, and the medial division of sternite 8. This is also consistent with the EPG analysis, with the Extension group recovered as sister to Archiceroptera mahukani group. (excluding the R. exuberata and - 22 -

Pterogramma sequences discussed previously). Although there are morphological differences that can separate the

Extension group from the rest of Archiceroptera, they are considered here as either plesiomorphic characters for the clade or as synapomorphic characters for the Extension species group.

The three remaining species groups are here treated as two clades that warrant treatment as genera. The prominens group was previously treated as part of Rudolfina, and includes only two species. This group was recovered as a sister group to either Robustagramma or Archiceroptera in the 11 morphological trees. Both molecular analyses recovered it outside and distant to Robustagramma, Archiceroptera and Rudolfina, which suggests that it is not a derived clade within any of those genera. We treat it here as a separate genus (see chapter 4).

The sororcula and enigmata groups were retained as sister groups in the morphological analysis, and are unique from other members of the Archiceroptera genus complex as the male mid femur has lost the ventrobasal setal cluster and the female terminalia are telescopic, unlike all EPG genera except Bitheca. In the EPG molecular analysis, these two groups were recovered together with the prominens group. The EPG major diversity is centered in the Neotropics although several clades appear to have dispersed into the Nearctic (and one into the

Palaearctic). Archiceroptera (including the ternum-group and brevivilla-group) and the prominens group appear to be the only EPG genera that have remained largely Neotropical (two Archiceroptera species ranges extend into several highland sites in Mexico), while Rudolfina is the only EPG genus that is largely Nearctic. There is relatively little geographic overlap of Rudolfina and Archiceroptera, with the exception of the widespread R. exuberata. Both

Archiceroptera, to the north, and. Rudolfina, to the south, have distributions that are apparently limited by the

Isthmus of Tehuantepec (see Barrier et al. 1998). The relatively low elevation gap that occurs in the Isthmus appears to be a significant barrier to the primarily high elevation Rudolfina, while the mountains to the north have limited the primarily low elevation Archiceroptera.

There are still gains to be made in further examining the relationships both within the EPG and within the

Limosininae. Future studies should look at obtaining suitable material, collected and preserved appropriately, to expand on the molecular study to include of nuclear genes. Further resolution of the EPG phylogeny may be assisted by inclusion of character sets not considered here, but that have been utilized in other studies. The use of wing interference patterns (WIP) (Shevtsova et al. 2011) was shown by Yau (2017) to provide some phylogenetic signal in Bromeloecia and could be of some value in support of relationships between genera. They are known to also occur in Pterogramma, and Robustagramma, but no distinct patterns have been observed in Archiceroptera, the - 23 - prominens group, or Rudolfina. Some members of the sororcula and enigmata groups appear to have consistent WIP patterns but the homology of this character set with the other genera is not clear.

- 24 -

1

2 3.4 Tables and Figures

3 Table 3.1. Morphological character states for the epandrial process group (EPG).

Character 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42

Thoracochaeta 0 0 0 0 0 0 1 1 0 0 1 1 0 0 1 1 1 1 0 1 0 01 0 1 0 0 0 01 1 0 0 01 0 0 0 0 0 0 0 0 0 2 Archicollinella 0 0 0 0 0 0 0 1 0 0 1 1 1 1 1 1 0 1 1 0 0 0 0 1 1 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 2 Archiceroptera 0 0 0 1 0 0 0 0 0 1 0 2 0 0 0 1 01 1 1 0 0 0 1 0 1 0 0 0 0 01 0 1 1 0 1 1 1 0 1 0 0 2 mahukani group Archiceroptera 0 0 1 01 0 0 0 0 0 1 0 12 0 0 0 1 01 1 1 0 0 0 1 0 1 0 0 0 0 1 01 1 1 0 1 1 1 0 1 0 0 2 (ternum-group) Archiceroptera 0 0 1 0 1 0 0 0 0 0 0 2 0 0 0 1 01 1 1 01 0 1 1 0 1 0 0 0 0 0 0 01 0 0 0 1 1 0 1 0 0 2 (Extension group)

Archiceroptera 0 0 1 0 0 0 0 0 0 1 1 0 01 01 01 1 0 1 1 0 0 0 1 0 1 0 0 0 0 1 01 1 1 0 1 1 1 0 1 0 0 2 brevivilla-group Prominens group 0 0 1 0 0 0 0 0 1 0 0 12 0 0 0 1 1 1 1 0 0 0 1 0 1 1 0 0 0 0 1 0 0 0 0 0 1 0 0 0 1 0 Rudolfina 0 1 1 0 1 0 1 1 1 0 1 0 1 1 1 1 0 0 0 0 1 0 1 1 0 1 01 0 0 0 0 1 1 0 1 1 0 01 0 1 23 0 Sororcula group 0 1 1 0 1 01 1 2 1 0 1 0 1 1 1 0 0 0 0 0 0 0 1 0 1 1 1 1 1 0 0 0 0 0 0 0 0 0 0 0 23 01 Enigmata group 0 1 01 0 1 1 1 2 1 0 1 0 1 1 1 0 0 0 0 0 0 0 1 1 1 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 2 Pterogramma 1 1 1 0 1 01 1 2 1 0 1 0 1 1 1 0 0 0 1 0 1 0 1 0 1 0 0 01 0 0 0 0 0 0 0 0 0 0 0 01 01 0 Aptilotella 1 1 1 0 ? ? ? ? ? 0 1 0 1 1 1 0 0 0 1 0 1 01 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 1 0 0 Bitheca 0 1 1 0 0 01 0 1 1 0 01 12 1 1 1 1 1 0 0 1 0 0 1 1 0 0 0 1 1 0 0 1 0 0 0 0 0 0 1 0 02 ? Robustagramma 01 0 01 0 0 0 0 2 1 0 1 12 0 0 0 0 0 1 1 0 1 1 1 0 1 1 01 0 0 0 0 0 0 0 0 0 1 0 1 0 2 0 Bromeloecia 0 1 1 0 01 0 0 01 01 0 1 01 1 01 1 0 0 0 0 0 1 0 1 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 0 4

- 25 -

Figure 3.1. Maximum likelihood analysis of cytochrome c oxidase subunit I (COI) 5P (Part 2).

Numbers at nodes are aBayes values. The leading codes given for each specimen are the unique identifiers within the BOLD database. The colour codes are as follows: pink = non-

Limosininae; black = Limosininae excluding epandrial process group; blue = epandrial process group excluding the Archiceroptera genus group; and red = Archiceroptera genus complex. Vertical lines denote different parts of the Archiceroptera genus complex.

- 26 -

Figure 3.2. Maximum likelihood analysis of cytochrome c oxidase subunit I (COI) 5P (Part 1).

Numbers at nodes are aBayes values. The leading codes given for each specimen are the unique identifiers within the BOLD database. The colour codes are as follows: pink = non-

Limosininae; black = Limosininae excluding epandrial process group; and blue = epandrial process group excluding the Archiceroptera genus complex.

- 27 -

Figure 3.3. Maximum likelihood analysis of cytochrome c oxidase subunit I (COI) 5P for the genera belonging to the epandrial process group with aBayes support values for each node.

The leading specimen codes are the unique identifiers within the BOLD database. Lines denote specimens belonging to the Archiceroptera genus complex: 1 = Rudolfina; 2 = prominens group; 3 = enigmata and sororcula (in part) groups; 4 = Archiceroptera mahukani, ternum and brevivilla groups; 5 = sororcula group (in part); 6 = R. exuberata and Extension groups; and 7 = sororcula group (in part).

- 28 -

Figure 3.4. Strict Consensus Tree (A) and Majority Rules Tree (B) from the morphological analysis of the epandrial process group.

- 29 -

Figure 3.5. Morphological phylogeny of the epandrial process group, including species groups of the Archiceroptera genus complex. One of 11 equal length trees.

- 30 -

CHAPTER 4 - PECTINOSINA, A NEW NEOTROPICAL GENUS OF LIMOSININAE (DIPTERA: SPHAEROCERIDAE)

4.1 Abstract

Pectinosina nov. gen. is described to include P. prominens (Duda) and P. carro n. sp., two

Neotropical species with an unusual comb-like posterolateral margin of female tergite 8 and a mid tibial chaetotaxy that separates it from related genera.

4.2 Introduction

The genus Pectinosina is described to include P. prominens (Duda), a species frequently collected in dung and carrion traps from Central and South America, and P. carro n. sp. from South America.

These species will key out to the “Archiceroptera genus complex” in couplet 78 in Marshall and

Buck (2010). Pectinosina is defined by the pectinate female tergite 8 (Fig. 3B and C), and the mid tibial chaetotaxy, which has 4–6 setae anterodorsally on the basal 1/3, 3–4 smaller setae posterodorsally near the midlength and five dorsal setae on the apical 1/3 (Fig. 1C). The relationships between Pectinosina and other related genera are discussed.

4.3 Materials and Methods

All specimens examined were dried and most were point-mounted with white glue. Abdomens of selected specimens were removed and cleared by immersion into hot 10% potassium hydroxide for

6-10 minutes before being neutralized with 10% acetic acid for 10 minutes, rinsed in deionized water, and then placed into glycerin for examination. Dissected genitalia were stored in glycerine in microvials pinned below the specimen.

Label Data and Distribution Maps

Label data are presented in a standardised form. Short-forms or abbreviations used on the labels are, where possible, given in full. Geographical coordinates are given if present on the original label.

Specimens were given unique identifiers and their collection data was captured within the BIOTA - 31 - database at the University of Guelph Insect Collection (Guelph, Ontario, Canada); this data will ultimately be hosted on Canadensys but is not repeated in the text except for holotypes or as image reference. Collection data for paratypes and other specimens examined were organized alphabetically by country, state/province, and locality name. Species distribution maps (Fig. 4.6) were generated using SimpleMappr (Shorthouse 2010).

Depositories of Material Examined

Depository abbreviations are as follows: DEBU (University of Guelph Insect Collection, University of Guelph, Guelph, Ontario, Canada); CNCI (Canadian National Collection, Ottawa, Ontario,

Canada), INBC (Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica);

MHNM (Museum National d’Histoire Naturelle, Paris, France); MIZA (Museo del Instituto de

Zoología Agrícola Francisco Fernández Yépez; Universidad Central de Venezuela, Maracay,

Venezuela); MUSM (Museo de Historia Natural, Universidad Nacional Mayor de San Marcos,

Lima, Peru); MZSP (Museu de Zoologia, Universidade de São Paulo, São Paulo, São Paulo, Brazil);

NHMW (Vienna Museum of Natural History, Vienna, Austria), QCAZ (Departamento de Biología,

Pontífica Universidad Católica del Ecuador, Quito, Ecuador); ROME (Royal Ontario Museum,

Toronto, Ontario, Canada); UASC (Museo de Historia Natural Noel Kempff Mercado, Santa Cruz de la Sierra, Bolivia); UNAM (Universidad Nacional Autónoma de México, Mexico City, Mexico);

USNM (United States National Museum of Natural History, Smithsonian Institute, Washington,

D.C., U.S.A.). Material is deposited in DEBU unless otherwise noted.

Terminology

External morphological terminology follows Cumming and Wood (2010), Marshall and Buck

(2010) and Smith and Marshall (2004) with some modifications as follows. Seta(e) and setula(e) are the large and small socketed macrotrichia, respectively. The CuA1 stubvein refers to the portion of

CuA1 that extends beyond cell dm-cu. Internal morphological terminology follows Smith and

Marshall (2004) and Paiero and Marshall (in prep.). The spermathecal stem is used to describe the individual ducts of each of the paired spermatheca. - 32 -

4.4 Pectinosina gen. nov.

Type Species: Pectinosina prominens (Duda) comb. nov., by present designation.

Diagnosis: Relatively stout and hirsute limosinine. R4+5 straight or weakly curved just before meeting costa. M1 ending before wing margin. Mid tibia with series of 4-6 anterodorsal setae extending from basal 1/3 to midlength, series of 2-4 posterodorsal setae at midlength and 5 strong distal dorsal setae (see Fig. 4.1C; see also Fig. 26 in Marshall and Buck 2010); midventral seta present in both sexes; male with ventral comb of 20–25 robust dark seta on distal half. Male sternite

5 with deep medial emargination on posterior margin flanked by strong seta. Female tergite 8 bipartite, with lateral sclerites extending posteroventrally and ending in 3–4 teeth

Description: Length 1.4–3.1 mm. Brown with scape, notopleuron, postpronotal lobe (usually), and lower half of frons yellow to orange-yellow; tibiae and tarsi usually pale brown or yellow; haltere variable from completely white to almost completely black, but usually dark with apex pale.

Head: Eye large, ovoid with slight anterodorsal emargination; height 2.0–2.4× genal height. Frons matt, with 2 strong exclinate orbital setae, 4–5 interfrontal setae, 2–5 inclinate orbital setulae, and 5-

10 small setulae along orbital plate; orbital setae equal in length; interfrontal setae, except for shorter anterior seta, equal in length; inclinate orbital setulae ~1/2 length of interfrontal setae. Outer vertical seta strong, exclinate. Inner vertical strong and inclinate. Occipital seta weak, inclinate.

Paravertical seta weak, inclinate. Postvertical weak, inclinate. Ocellar triangle with pair of ocellar setae and 6–7 additional setulae between ocelli. Face with upper half tuberculate between antennae, with broad, poorly-defined medial carina extending from tubercle onto lower half; face weakly excavated below tubercle to make lower margin appear weakly projecting. Vibrissa strong. Gena with 2 strong subvibrissal setae; subvibrissal setae subtended by cluster of 5–20 weaker setae.

Clypeus short and wide. Maxillary palp with weak apical seta and 2–3 strong preapical setae on ventral surface. First flagellomere apically rounded with weak dorsal angle. Arista arising preapically on dorsal surface of first flagellomere; pubescence short and uniform.

- 33 -

Thorax: Scutum pruinose. Postpronotal lobe with 1 strong outer seta and 1 weaker inner humeral seta. Notopleurual seta, presutural supra-alar seta, postalar seta and intrapostalar seta strong.

Dorsocentral seta in 1 strong prescutellar pair. Acrostichal setulae in 10–12 rows, with pair of strong prescutellar acrostichal seta. Scutellum wider than long, with basal and apical pair of strong setae.

Katepisternum with 2 setae; weak anterior setae and stronger posterior seta (~3.0× anterior setae) near posterodorsal corner. Prosternum narrow and bare.

Legs: Fore femur with 4–6 dorsal and ventral setae. Mid femur with row of 9–12 short setae on anterior face, distally with 5-6 long setae on anteroventral edge; male ventrally with basal cluster of

15–30 dark robust setae. Mid tibia with unique dorsal chaetotaxy (see diagnosis); midventral seta present in both sexes; male with ventral comb of 20-25 small robust setae on distal half. Mid- basitarsus with weak ventral seta. Hind femur evenly setose. Hind tibia with 2-7 enlarged posterodorsal setae and a weak ventral spur (rarely a midventral hind tibial seta present; see debu01082072). Hind tarsus with only basal tarsomere swollen.

Wings (Fig. 4.1B): Completely hyaline. Costa basally with 2 equal costagial setae. Costa ending before wing apex with R4+5; R2+3 curved towards costa near apex; R4+5 weakly curved anteriorly.

Second costal sector 1.25× third costal sector. Cell dm with small CuA1 stub vein present beyond cell dm. Allula narrow, with posterior margin straight.

Abdomen: Abdominal sclerites well sclerotized and hirsute; setae longer along posterior and lateral margins.

Male abdomen: Posterior margin of sternite 5 medially with some degree of desclerotization extending anteriorly from medial emargination on posterior margin; posterior margin usually with dense setae adjacent to medial emargination. Transverse portion of sternite 6 broadly arcuate medially, expanded on right side to quadrate sclerite that is closely associated with large ring sclerite. Epandrium asymmetrical, with narrow process on right side extending to hypandrium, subanal plate incomplete. Cercus elongate and acuminate with tip bent posteriorly. Surstylus variable between species; generally weakly trilobed with 2 or more robust setae present on inner surface of posterior lobe. Hypandrium Y-shaped, with posterior pair of arms extending to postgonite; right arm laterally notched for reception of tip of epandrial process. Pregonite small, - 34 - triangular. Postgonite with apical ½ narrow. Basiphallus simple (epiphallus absent). Distiphallus: basal sclerite present; U-shaped sclerite present; dorsal sclerite whip-like and projecting distally beyond the tip of the acrophallus; acrophallus well-developed, with mid-lateral projections; lateral flanking sclerite extending from base of dorsal sclerite lateroventrally at base of acrophallus; acrophallus with distinct lateral spinose projections; distal ventral sclerite present but poorly developed.

Female Abdomen: Terminalia short and broad, not telescoping. Tergite 8 dorsally desclerotized and divided into 2 lateral sclerites; lateral sclerites with 3–4 teeth on posteroventral margin; ventral surface, near anterolateral corner, with small process. Epiproct evenly sclerotized, hirsute with pair of seta near midlength. Cercus ovoid and relatively short, not projecting posteriorly beyond the tip of the abdomen; surface setose with 2–4 larger setae near tip. Sternite 7 with posterior margin evenly rounded. Sternite 8 transverse, entire and poorly sclerotized with cluster of 2-3 setulae on membrane posterior to posterolateral corners of sclerite. Hypoproct narrow, transverse, horseshoe shaped. Spermathecae (2+1) biconcave discs with elongate sclerotized ducts.

Etymology: The genus name is derived from the Latin word for comb (pectine) and used in combination with “-sina”, a common ending used in a number of Limosinine genera. Gender feminine.

Biology: Most material of this group was collected in dung traps, or in pan traps associated with dung or carrion baits. Specimens have also been collected in Malaise and flight intercept traps.

Label data for some specimens of Pectinosina prominens (Duda) indicate an association with dung rolling scarabs (see species discussion).

Relationships: Pectinosina prominens Duda was originally treated in Acuminiseta (as a subgenus of

Leptocera; Duda 1925) but was placed in Rudolfina Roháček by Roháček et al. (2001) because it was more similar to, and more closely related to, Rudolfina than to Acuminiseta (which does not occur in the New World). Pectinosina prominens does not, however, fit within the genus Rudolfina as recently redefined by Paiero & Marshall (in prep.). A morphological phylogeny of the epandrial process group (Chapter 3), a primarily New World clade recognized by Marshall and Cui (2004) to include Rudolfina, Robustagramma Marshall & Cui, Archiceroptera Papp, Pterogramma Spuler, - 35 -

Aptilotella Duda and Bitheca Marshall, recovered Pectinosina with Robustagramma. However,

Pectinosina does not fit with Robustagramma as it is currently defined (Marshall and Cui 2004), and while Robustagramma could be redefined to include Pectinosina as a basal clade, to do so would make Robustagramma less diagnosable, and we consider the apomorphic mid tibial chaetotaxy and female terminalia enough support to warrant its treatment as a unique genus outside of Robustagramma.

The habitus and mid tibial chaetotaxy of Pectinosina is superficially similar to Leptocera, and the female tergite 8 of some Leptocera species are margined posteriorly with numerous modified setae to make it appear pectinate. Despite some general similarities between the two genera, Leptocera does not belong within the EPG as it lacks the epandrial process. The similarity of tergite 8 between the genera is certainly homoplastic as the combs in Pectinosina are a modification of the sclerite and not the setae as in Leptocera. Pectinosina may also be confused with the closely related

Archiceroptera, which can be easily separated by M1 extending to the wing margin, the mid tibial chaetotaxy and several differences in the female terminalia, including the spinose cercus.

4.4.1 Species Descriptions

Pectinosina species are externally indistinguishable and species identification requires close examination of the male or female abdomen. The following descriptions are therefore limited to abdominal characters.

Pectinosina prominens (Duda), n. comb

Synonyms: Rudolfina prominens (Duda, 1925); Leptocera (Acuminiseta) prominens Duda 1925

Description: Length 1.4-3.1 mm. Eye height ~2.0–2.4× genal height.

Male Abdomen (Fig. 4.4): Sternite 5 posteriorly with pair of acute teeth on each side of medial desclerotization; posterior margin sinuate lateral to teeth; medial desclerotization “T” shaped with transverse desclerotization 1/5th from anterior margin, and with elongate desclerotization extending to posterior margin. Epandrium with numerous setae. Surstylus (in lateral view) hirsute with

- 36 - anterior and posterior lobes (may appear trilobed in lateroventral view); anterior lobe boot-shaped with square “toe” and sinuate distal edge; posterior lobe simple but pronounced, with two short robust setae on inner surface. Cercus acute, as long as surstylus, with apex abruptly posteriorly recurved. Distiphallus (Fig.4. 4G-I): first dorsal sclerites elongate, projecting beyond tip of acrophallus by approximately half the length of sclerite before tip; acrophallus with rounded mid- lateral projections.

Female Abdomen (Fig. 4.5): Tergite 7 with posterior margin usually with a small V-shaped emargination (sometimes reduced to small concavity or absent entirely). Lateral sclerites of tergite 8 posteriorly with 3 (rarely) or 4 (usually) ventral teeth Paired spermathecae each with stems slightly less than width of spermatheca and with sclerotized portion of common duct ~1/3 length of stems; single spermatheca with sclerotized duct length slightly less than width of spermatheca.

Distribution: Widespread in the Neotropics.

Specimens Examined: Syntypes (1 male, 1 female): PARAGUAY: S. Bernardino, (K.A.G.)

Fiebrig (no date given) (NHMW). Non-type Material (2,375 specimens): BELIZE: Cayo: 1 male,

1 female, Caves Branch, forest, dung, 23-29 Aug 1972, S. & J. Peck; Toledo: 1 male, BARC, near

San Pedro Columbia, 16°17'N, 88°58'W, malaise & pans, 10-12 Mar 2002, J. Skevington.

BOLIVIA: La Paz: 16 males, 3 females, Arroyo Tuhiri W Mapiri, 15°17'27”S, 68°15'29”W, 10

Apr 2001, S.A. Marshall; 1 male, Coroico, 1700 m, grassy slope, dung traps, 5 Apr 2001, S.A.

Marshall; 1 male, Heath River Wildlife Centre, ~21 km SSW Puerto Heath, 12°40'S, 68°42'W, rainforest, malaise, 1-11 May 2007, S.M. Paiero; 10 males, 4 females, Heath River Wildlife Centre,

~21 km SSW Puerto Heath, 12°40'S, 68°42'W, tree fall, yellow pans, 5-9 May 2007, Paiero & Kits

(DEBU and UASC); 12 males, 1 female, Heath River Wildlife Research Centre, 12°40'S, 68°42'W, treefall, yellow pans, 5-9 May 2007, Paiero & Kits; 2 males, San Antonio, ca. 8 km S Mapiri,

15°20'56”S, 68°13'31”W, secondary forest, dung pans, 11 Apr 2001, S.A. Marshall; Santa

Barbara: 1 male, N. Coroico, Yungas, 1100m, 4-5 Jan 1976, L.E. Peña; Santa Cruz: 2 females,

Potrerillos de Guenda, 17°40'29”S, 63°27'22”W, 4-7 Apr 1998, H. & A. Howden (CNCI); 1 male,

Refugio Los Volcanes, 4 km N Bermejo, 18°6'15”S, 63°35'55”W, 1058m, trail along river, Malaise,

3-7 Oct 2014, Norrbom et al. (USNM); 2 females, Refugio Los Volcanes, 4 km N Bermejo, - 37 -

18°6'15”S, 63°35'55”W, 1058m, Malaise trap along river, 1-4 Mar 2014, Norrbom et al.. BRAZIL:

Bahia: 2 females, B.A., 15km NE Porta Sequro (Ecological Reserva “Pau-Brasil”), primary

Atlantic forest, Shannon trap, 19-27 Feb 1986, D.S. & V.C.S. Amorim; Espirito Santo: 3 males, 2 females, Linhares, malaise trap, Nov 1967, F.M. Oliveira; Mato Grosso: 1 female, Tiradentes, pan traps along creek, 15-17 Feb 1990, S.A. Marshall; 1 male, Tiradentes, Serra de Tiradentes, human dung, 16 Feb 1990, S.A. Marshall; Mina Gerais: 2 females, Lavras, 1km E, dung traps in ditch, 18-

20 Feb 1990, S.A. Marshall; Paraná: 2 males, 2 females, Londrina, Mata dos Godoy, 28-31 Jan

1990, S.A. Marshall (MZSP); 1 male, Curitiba, Curitiba survey, 19 Jan 1990; 1 female, Curitiba,

FIT in woods behind Nat. Hist. Museum, 5-9 Feb 1990, S.A. Marshall; 10 males, 4 females,

Curitiba, 30 km SE, BR 277, dung traps, 6-9 Feb 1990, S.A. Marshall (DEBU, MZSP); Rio de

Janeiro: 1 female, Grajau, 25 Jul 1964, H.S. Lopes; 1 male, Grajau, 6 Sep 1964, H.S. Lopes; São

Paulo: 2 males, 3 females, USP Biology Station, human dung, 5-6 Feb 1979, R. Woodruff & J.

Runnacles (MZSP). COLOMBIA: Norte de Santander: 3 males, 5 females, Chinacota, 3mi. N,

3000ft, 8 Jun 1974, S. Peck; 2 males, Santiago, 2000ft, dung trap, 11-13 May 1974, S. Peck.

COSTA RICA: Alajuela: 1 male, 3 females, Florencia Forest, dung tp., 28 Feb 1980, H. Howden;

1 male, 6 females, Río Peñas Blancas, 700 m, 18 Aug 1986, L. Masner; 1 male, 2 females, Volcán

Tenorio, N slope near Bijagua Biological Station, 700 m, rainforest, RET over Atta mound, 16-20

Jun 2000, S.A. Marshall; 2 males, Volcán Tenorio, N slope near Bijagua Biological Station, 700 m, sweeping over Atta mound, 18 Jun 2000, S.A. Marshall; Cartago: 6 males, 5 females, Turrialba

Catie, 600 m, 26 Feb 1980, H. & A. Howden (INBC); 10 males, 4 females, Turrialba Catie, 600 m,

28 Feb 1980, H. & A. Howden; Guanacaste: 2 males, Cacao Field Station, 1000m, carrion traps,

18-20 Feb 1996, S.A. Marshall; 1 male, Cacao Field Station, 1250m, dung trap, 12-15 Feb 1996,

S.A. Marshall; 1 male, Guanacaste Cons. Area, Pitilla Field Station, Malaise, 29 Jan 1996, J. Noyes;

17 males, 15 females, Guanacaste Cons. Area, Ricon de la Vieja, Las Pailas, 1400 m, Clusea rosea forest litter, 18-20 Feb 1996, R. Anderson (DEBU, INBC); 2 males, Maritza Field Station, malaise,

3-9 Feb 1996, J. Noyes; Heredia: 1 male, 5 females, 10km N Puerto Viejo, La Selva Verde, FIT, 3

Mar 1991, H. & A. Howden (CNCI); 3 males, 3 females, 10km W Puerto Viejo, La Selva Verde, 2-

4 Mar 1991, H. & A. Howden; 1 male, La Selva, 50-100m, carrion trap, 18 Feb 1980, H.F. Howden; - 38 -

1 male, La Selva, around dung ball rolled by Canthon moniliatus, 2 Feb 1990, N. Grieg; 1 female,

La Selva, black light trap, 9 May 1989, B.V. Brown; 1 male, 1 female, La Selva, malaise trap, 15-21

May 1989, B. Brown & D. Feener; 1 female, La Selva, 1* rainforest, malaise, CES 200, 23-26 May

1988, B.V. Brown; 1 female, La Selva Biological Station, Malaise trap, SSO 1500, 1-8 May 1989,

B. Brown & D. Feener; 2 males, Puerto Viejo, La Selva Verde, FIT & dung traps, 3 Mar 1991, S.

Peck (INBC); 1 female, Puerto Vieja, La Selva Biological Station, black light, 23 Apr 1989, B.V.

Brown; Limon: 2 males, 3 females, Estrella Valley, Pandora, carrion trap, 20 Feb 1984, H.

Howden; Puntarenas: 2 females, Las Alturas, 1700m, dung, 12 Aug 1995, S.A. Marshall; 1 male, 2 females, Las Alturas, 1700m, dung trap, 12-13 Aug 1995, S.A. Marshall; 1 male, 1 female, Coto

Brus, Z.P. Las Tablas, Estacion Biologica Las Alturas, 8°57'7”N, 82°50'4”W, 1500-1600 m, malaise trap, 26 Nov-3 Dec 2012, ZADBI, (INBC); 1 male, Las Alturas, 8°57'N, 82°58'W, 1600 m, malaise trap, 11-14 Aug 1995, S.A. Marshall; 1 male, Monteverde, 10°18'N, 84°49'W, Pension Queteal, on human dung, 24 May 1987, A. Norrbom; 1 female, Monteverde, 1500 m, cloud forest, 29 Feb 1980,

W.R.M. Mason; 1 female, Monteverde, 1500m, cloud forest, dung traps, 19-25 Aug 1993, E.R.

Barr; 1 male, Monteverde, 1520 m, FIT, 11-18 Jun 1983, D.H. Lindeman; 2 males, Monteverde,

1520 m, FIT, 15-23 Jul 1983, D.H. Lindeman; 1 female, Monteverde, 1520 m, FIT, 9-13 Jul 1983,

D.H. Lindeman; 3 males, 3 females, Monteverde, 1560 m, dung trap, 11-18 Jun 1983, D.H.

Lindeman; 7 males, 7 females, Monteverde, 3 dung traps, 27 Feb 1991, H. & A. Howden; 1 female,

Monteverde, near biology station, sweep, 25 May 1998, S.A. Marshall; 1 female, Monteverde, San

Luis, 1000-1350 m, malaise trap, Jan 1993, Z. Fuentes; 5 males, 1 female, Osa Peninsula, Rincón,

2.5 km S, 8°42'1”N, 83°30'50”W, ~50 m, secondary forest, dung pans, 11 Aug 2001, M. Buck; 2 males, Osa Peninsula, Rincón, 2.5 km S, 8°42'1”N, 83°30'50”W, ~50 m, prim. forest, dung pitfalls,

11 Aug 2001; 1 male, Osa Peninsula, Rincón, 2.5 km S, 8°42'1”N, 83°30'50”W, ~50 m, secondary forest, fish pitfalls, 10-11 Aug 2001; 1 male, Osa Peninsula, Rincón, 2.5 km S, 8°42'1”N,

83°30'50”W, ~50 m, rainforest, sweeping, 10 Aug 2001, M. Buck; 1 male, Parque Nacional

Amistad, Estacion Las Mellizas, Fca. Cafrosa, 1300m, L-S-316100, 596100, Apr 1991, G. Mora,

(INBC); 1 female, Parque Nacional Amistad, Estacion Las Mellizas, Finca Cafrosa, 1300m, L-S-

316100-596100, Oct 1989, M. Ramirez & G. Mora (INBC); 1 male, 1 female, San Vito, Las Cruces, - 39 -

1200m, on elytra of Sulcophanaeus velutinus, 1 Mar 1983, B. Gill. ECUADOR: Guayas: 1 male,

78 km N Santa Elena, 27 km S Puerto López, 500ft, dung trap, 25-27 Jul 1976, S. Peck; Manabi: 1 male, Chone, 20km N, 300m, cacao plantation, 2 dung traps, 6-9 Jun 1976, S. Peck; Napo: 1 male,

Baeza, 5 Mar 1979, S.A. Marshall; 1 male, 2 females, Coca, Río Napo, 250 m, May 1965, L.E.

Peña; 6 males, 5 females, Jatun Sacha Reserve, 6 km E Misahuallí, 1°4'S, 77°37'W, 450 m, varzea, dung pans, 1-2 May 2002, Buck & Lonsdale (QCAZ); 61 males, 67 females, Jatun Sacha Reserve, 6 km E Misahuallí, 1°4'S, 77°37'W, 450 m, varzea, dung pans, 2-7 May 2002, M. Buck (DEBU,

QCAZ); 5 males, 2 females, Jatun Sacha Reserve, 6 km E Misahuallí, 1°4'S, 77°37'W, 450 m, by stream, dung pans, 5-7 May 2002, S.A. Marshall; 2 males, 2 females, Jatun Sacha Reserve, 6 km E

Misahuallí, 1°4'S, 77°37'W, 450 m, varzea, sweeping, 2 May 2002, M. Buck; 1 male, Pompeya, Río

Napo, 14-22 May 1965, L.E. Peña; 1 male, Rio Piocullin, S side, SW Puerto Napo, S. Limonchicta,

600m, 1*lowland rainforest, malaise head, ROM 870020, 23-27 May 1987; 1 female, Tena, 500m, secondary rainforest, malaise head, 22-27 May 1987, Brown & Coote; 23 males, 28 females,

Tiputini Biodiversity Station, vicinity Yasuní National Park, 0°38'S, 76°0'W, human dung pitfalls,

14-19 Feb 1998, D.C. Darling (DEBU, ROME); 2 females, Tiputini Biodiversity Stn., 0°36'50”S,

76°9'1”W, May 2011, S.A. Marshall; 26 males, 11 females, Tiputini Biodiversity Stn., vicinity

Yasuni National Park, 0°38'S, 76°10'W, pitfall trap (human dung), 14-19 Feb 1998, D.C. Darling; 2 males, 3 females, Yasuní National Park, Yasuní Research Station, 0°38'S, 76°36'W, rainforest, malaise trap, 3-20 Nov 1998, Pape & Viklund; Pichincha: 1 male, Alluriquin, 23km E, Chiriboyo

Ret., 4600ft, dung, 19-27 Jun 1975, S. Peck; 1 female, Maquipucuna Biological Reserve, main trail,

0°7'34”N, 78°37'57”W, 1400-1600 m, 27 Apr 2002, S.A. Marshall; 20 males, 9 females,

Maquipucuna Biological Reserve, river trail, 0°7'34”N, 78°37'57”W, 1200 m, near stream, pans/ dung, 26-28 Apr 2002, S.A. Marshall (DEBU, QCAZ); 2 males, 1 female, Nanegalito, 7 km SE, trout farm 'San José', 0°3'54”S, 78°40'36”W, 1500 m, river edge, pan traps, 30-31 Oct 1999, S.A.

Marshall; 1 male, Nanegalito, 7 km SE, trout farm 'San José', 0°3'54”S, 78°40'36”W, 1500 m, riverine forest, sweeping tree falls, 27-30 Oct 1999, S.A. Marshall; 3 males, 1 female, Nanegalito, 7 km SE, trout farm 'San José', 0°3'54”S, 78°40'36”W, 1500 m, 30 Oct 1999, S.A. Marshall; 31 males,

10 females, Palenque, day 3 trap, 24-25 Mar 1976, S. Peck; 1 male, 1 female, Quito, 46km E, - 40 -

4000m, elfin forest, dung traps, 2-6 Mar 1976, S. Peck; 6 males, 5 females, Rio Palenque, carrion,

27 Feb 1979, S.A. Marshall; 242 males, 226 females, Rio Palenque, dung, 25 Feb 1979, S.A.

Marshall; 124 males, 123 females, Rio Palenque, dung, 27 Feb 1979, S.A. Marshall; 55 males, 51 females, Rio Palenque, dung trap, 22-23 Feb 1976, S. Peck (DEBU, QCAZ); 24 females, Rio

Palenque, dung trap, 25-26 Feb 1976, S. Peck; 19 males, 8 females, Rio Palenque, J. Glasser trap,

26 Feb 1976, S. Peck; 1 male, 1 female, Rio Palenque, 22 Feb 1976, S. Peck; 59 males, 27 females,

Rio Palenque, 25-26 Jan 1976, S. Peck (CNCI, DEBU); 17 males, 5 females, Rio Palenque, 26 Feb

1976, J. Glaser; 1 female, Rio Palenque Reserve Station, Malaise trap, Feb 1983, M. Sharkey & L.

Masner; 1 male, Rio Palenque Science Center, 47km S Santo Domingo, 180m, 1*lowland rainforest, malaise head, 29 Apr-5 May 1987, Coote & Brown; 1 female, Rio Palenque Science

Center, 47km S Santo Domingo, 180m, rotting fruit, 1-5 May 1987, Brown & Coote; 1 male, 1 female, Río Palenque Station, 47 km S Santo Domingo, 26-27 May 1975, S. Peck; 3 males, Río

Palenque Station, 47 km S Santo Domingo, 28 May 1975, S. Peck; 4 males, Río Palenque Station,

47 km S Santo Domingo, carrion, 27-28 May 1975, S. Peck; 2 males, 1 female, Río Palenque

Station, 47 km S Santo Domingo, carrion traps, day 3, 26-27 May 1975, S. Peck; 27 males, 18 females, Río Palenque Station, 47 km S Santo Domingo, traps 3-5, day 1, 22-23 Feb 1976, S. Peck;

1 male, Rio Palenque Station, 47km S Santo Domingo, 250m, carrion trap, day 4, 27-28 May 1975,

S. Peck; 31 males, 22 females, Rio Palenque Station, 47km S Santo Domingo, 250m, dung, 17-25

Feb 1979, S.A. Marshall; 3 females, Rio Palenque Station, 47km S Santo Domingo, 250m, forest, dung traps, 22-27 Feb 1976, S. Peck; 1 male, 1 female, Rio Palenque Station, 47km S Santo

Domingo, 250m, rainforest, malaise-FIT, 5 May-25 Jul 1985, S. & J. Peck; 26 males, 21 females,

Rio Palenque Station, 47km S Santo Domingo, 250m, 17-25 Feb 1979, S.A. Marshall; 1 male, 1 female, Santo Domingo, 4km SE, 500m, 3 forest dung pans, 8-11 Jun 1976, S. Peck; 2 males,

Tinalandia, 1120m, wet lower montane rainforest, Malaise head, ROM870007, 9-13 May 1987,

L.D. Coote & B.V. Brown; 8 males, 4 females, Tinalandia, 16 km SE Santo Domingo, 680 m, dung trap, 21-22 Jun 1975, S. Peck (CNCI); 4 males, 9 females, Tinalandia, 16 km SE Santo Domingo,

680 m, forest, dung traps 32, 16-28 Jun 1975, S. Peck; 19 males, 15 females, Tinalandia, 16 km SE

Santo Domingo, 680 m, rainforest, malaise-FIT, 4 May-25 Jul 1985, S. & J. Peck. FRENCH - 41 -

GUIANA: St. Laurent du Maroni: 1 male, Mitaraka, MIT-A-SL, 2°14'18”N, 54°27'8”W, 352m, tropical moist forest (slope), yellow pan traps, 3-8 Mar 2015, M. Pollet (MHNM); 2 females,

Mitaraka, MIT-C-TOP, 2°13'59”N, 54°26'38”W, 433m, tropical moist forest (plateau), blue pan traps, 2-8 Mar 2015, M. Pollet (MHNM); 1 female, Mitaraka, MIT-C-TOP, 2°13'59”N,

54°26'38”W, 433m, tropical moist forest (plateau), yellow pan traps, 2-8 Mar 2015, M. Pollet; 1 female, Mitaraka, MIT-DZ, 2°14'2”N, 54°27'1”W, 306m, tropical moist forest (plateau-slope- cleared), FIT, 1 Mar 2015, Touroult & Poirier; 1 male, Mitaraka, MIT-C-RBF1, 2°14'11”N,

54°26'50”W, 258m, tropical moist forest (bas fond), yellow pan traps, 27 Feb-8 Mar 2015, M.

Pollet; 1 male 1 female, Mitaraka, MIT-C-SL, 2°14'8”N, 54°26'42”W, 373m, tropical moist forest

(slope), yellow pan traps, 2-8 Mar 2015, M. Pollet (MHNM). GUATEMALA: Izabal: 1 male,

Izabal, 350 m, malaise trap, 14 Dec 1986, M.J. Sharkey; Sacatepéquez: 4 males, 4 females, Volcán

Atitlán, Ref. Quetzal, 14°33'2”N, 91°11'32”W, 1670m, cloud forest, FIT, 13-16 Jun 2015, Falin &

Carrillo (UVGC); 13 males, 8 females, Volcán Atitlán, Ref. Quetzal, 14°33'2”N, 91°11'32”W,

1670m, cloud forest, FIT, 3-6 Jun 2015, Falin & Carrillo; 1 male, La Unión, 3.5 km SE, 1500 m,

FIT tp#1, 23-25 Jun 1993, Ashe & Brooks; 1 male, La Unión, 3.5 km SE, 1500 m, FIT, #102, 23-25

Jun 1993, Ashe & Brooks; 1 female, La Unión, 3.5 km SE, 1500 m, FIT, #127, 25-27 Jun 1993,

Ashe & Brooks; 1 male, 2 females, La Unión, 3.5 km SE, 1500 m, FIT, #128, 25-27 Jun 1993, Ashe

& Brooks; Peten: 1 female, Tikal, dung trap, 28-30 Jul 1978, Helava & Kukal. GUYANA:

Mazaruni-Potaro: 1 male, Takutu Mountains, 6°15'N, 58°55'W, window trap in montane rainforest near logging area, 8-10 Dec 1983, Perkins & Steiner; Potaro-Siparuni: 1 male, Mount Wokomung,

5°7'53”N, 59°48'31”W, 698m, 1° forest, pitfall trap (human dung), 21-26 Oct 2004, B. Hubley;

Rupununi: 16 males, 11 females, Kurupukari, Essequibo River, 200ft, 1° forest, dung traps, 9 Oct

1990, B. Hubley (DEBU, ROME); 1 male, Kurupukari, W side Essequibo R., 200ft, 1° rainforest/cattle trail, screen sweep ROM 905047, 10 Oct 1990, L.D. Coote (ROME); 5 males,

Kabocalli, Iwokrama Forest Reserve, 100 m, FIT, 22-25 May 2001, Brooks & Falin; 6 males,

Kabocalli, Iwokrama Forest Reserve, 60 m, FIT, 3-5 Jun 2001, Brooks & Falin. HONDURAS:

Olancho: 1 male, La Muralla, FIT, 12 Jan 1995, R. Cordire; 1 female, Parque Nacional La

Murilla/La Union, FIT, Jan 1995, R. Cordire; Cortez: 9 males, Lago de Yojoa, 2600ft, dung trap, 1- - 42 -

2 Jun 1994, Howden. MEXICO: Chiapas: 2 males, 4 females, Laguna Belgica, 16 km NW

Ocozocoaulta, 970m, FIT, 13 Jun 1990, H. & A. Howden & Gill; 1 male, 2 females, Laguna

Belgica, 16 km NW Ocozocoaulta, 970m, 31 May 1990, H. & A. Howden; 2 females, Nahá,

16°56'57”N, 91°35'41”W, 960 m, mesophil forest, malaise trap, 29 May 2008; 11 males, 9 females,

Ocozocoautla, 11mi NW, 3400ft, oak-evergreen forest, human dung, 19-25 Aug 1971, A. Newton

(FMNH); 2 females, Palenque, 4mi S, 700ft, rainforest, human dung, 7-15 Aug 1971, A. Newton; 1 female, Trinitaria, 2mi S, 5100ft, oak-trop. decid., human dung, 21-24 Aug 1971, A. Newton;

Hidalgo: 1 male, 2 females, Tlanchinol, 2.5mi N, 5200ft, cloud forest, dung, 6-11 Jul 1973, A.

Newton; 3 males, 2 females, Tlanchinol, 3.5mi N, 5100ft, cloud forest, human dung, 6-11 Jul 1973,

A. Newton (UNAM); Oaxaca: 5 males, 1 female, Valle Nacional, 5mi S, 1600ft, trop.oak.evgn., dung, 20 Jul-1 Aug 1971, A. Newton; Puebla: 1 female, Huanchinango, 5mi W, 6000ft, hardwood- pine, human dung, 3-7 Jul 1971, A. Newton; 1 female, Teziutlan, 4.5mi E, 5000ft, cloud forest, human dung, 10-14 Jul 1971, A. Newton; Veracruz: 2 males, Fortin, SW of Rio Metlas, 3250ft, human dung, 13-18 Jul 1971, A. Newton; 7 males, 3 females, Huatusco, 4mi N, 4100ft, cloud forest, dung, 11-16 Jul 1971, A. Newton; 2 males, 1 female, Teocelo, 10mi SW, 4400ft, oak, wet, human dung, 11 Jul 1971, A. Newton. PANAMA: Chiriquí: 1 female, Cerro Punta, 2 km E, 1760 m,

Baldwin forest, dung traps, 30 May-8 Jun 1977, S. Peck; 8 males, 3 females, Hartmann's Finca,

1550 m, dung trap, 31 May 1977, S. Peck; 1 male, 3 females, Hartmann's Finca, 1700 m, 28 Jun-3

Jul 1981, B. Gill; 6 males, Hartmann's Finca, 15 km NW Hato de Volcán, 1200 m, dung trap, 20-25

May 1977, S. Peck; 19 males, 26 females, Hartmann's Finca, 15 km NW Hato de Volcán, 1200 m, dung trap, 20-31 May 1977, S. Peck (CNCI, USNM); 12 males, 9 females, Hartmann's Finca, 15 km

NW Hato de Volcán, 1500m, dung, 20-25 May 1977, S. Peck; 3 females, La Fortuna Dam, 1000 m,

5-6 Jul 1981, B. Gill; 35 males, 11 females, Lagunas, 5km SW Hato del Volcan, 1360m, dung, 22-

26 May 1977, S. Peck; 10 males, 3 females, Lagunas, 5km SW Hato del Volcan, 1360m, dung, 22-

27 May 1977, S. Peck; 17 males, 3 females, Lagunas, 5km SW Hato del Volcan, 1360m, 22 May

1977, S. Peck; 1 male, Las Lagunas, 4.5 km WSW Hato del Volcán, 8360 ft, sweeping, 1-8 Jun

1977, S. & J. Peck; Colón: 1 male, 2 females, Santa Rita Ridge, 10 mi SE Colón, 270 m, dung trap,

10-12 Jun 1977, S. Peck. PARAGUAY: Caazapá: 1 male, Hermosa, San Rafael Reserve, prop. - 43 -

Lopez family, 26°18'29”S, 55°45'3”W, 80 m, FIT, 1-3 Dec 2000, Z.H. Falin; 1 male, Hermosa, San

Rafael Reserve, prop. Lopez family, 26°19'15”S, 55°45'3”W, 90 m, FIT, 3-6 Dec 2000, Z.H. Falin.

PERU: Loreto: 1 female, Campamento San Jacinto, 175-215 m, FIT, 5 Jul 1993, R. Leschen; 3 females, Teniente López, riverine forest, FIT, #199, 24 Jul 1994, R. Leschen; 1 female, Teniente

López, Riv. forest, FIT, #211, 26 Jul 1993, R. Leschen; Madre de Dios, 3 males, 12 females,

Amazonas Lodge, N of Atalaya, 12°52'12”S, 71°22'36”W, 480 m, FIT, 10-13 Nov 2007, D.

Brzoska (MUSM); 1 female, CICRA Field Station, 12°34'10”S, 70°6'4”W, 260m, garden, Malaise trap, 19-26 Aug 2010, M.J. Endara; 1 male, 1 female, CICRA, trail 2, 12°33'40”S, 70°6'23”W,

267m, Malaise, 10-16 Nov 2013, J. Caballero (MUSM); 1 female, Los Amigos Biological Station, palm swamp, yellow pans, 6-10 Jun 2006, Paiero & Klymko (MUSM); 1 male, 1 female, Pakitza, malaise trap & blacklight trap, 7 Mar 1992, Brown & Feener; 7 males, 10 females, Pantiacolla

Lodge, Alto Madre de Dios River, 12°39'18”S, 71°13'54”W, 420 m, FIT, 14-19 Nov 2007, D.

Brzoska; 1 female, Rio Tambopata Reserve, 30 km SW Puerto Maldonado, 12°12'S, 069°16'W, tropical moist forest, 19 Sep-10 Oct 1984, D.A. Grimaldi (AMNH); 2 females, Zona Reserva Manu,

Pakitza, 11°57'S, 71°17'W, 400 m, malaise trap, 13-18 Feb 1992, B. Brown & D. Feener; 1 male,

Zona Reserva Manu, Pakitza, 11°57'S, 71°17'W, 400 m, malaise trap, 18-23 Feb 1992, B. Brown &

D. Feener; 1 male, 1 female, Zona Reserva Manu, Pakitza, 11°57'S, 71°17'W, 400 m, malaise trap,

23-28 Feb 1992, B. Brown & D. Feener. U.S.A.: Florida: 1 male, No Name Key, seaweed drift, 16

Feb 1983, S.A. Marshall. VENEZUELA: Aragua: 6 males, 10 females, Henri Pittier National

Park, Rancho Grande Biological Station, 10°21'N, 67°41'W, 1250 m, May 1998, Ashe, Brooks &

Hanley; 1 male, 3 females, Maracay, Rancho Grande, 1200m, dung traps, 27-28 Feb 1995, S.A.

Marshall; 4 males, 8 females, Maracay, Rancho Grande, 1200m, cloud forest, FIT, 1-10 Aug 1987,

Bordon & Peck; 5 males, 13 females, Rancho Grande Biological Station, 10°21'N, 67°41'W, 1200 m, dung traps along near trail, 1 Mar-5 Aug 1995, S.A. Marshall (DEBU, MIZA); 1 male, 1 female,

Rancho Grande Biological Station, 10°21'N, 67°41'W, 1200 m, flight intercept trap, 14 May 1998,

Ashe, Brooks & Hanley; 1 female, Rancho Grande, La Cumbre cloud forest, 1500m, FIT, 1-10 Aug

1987, Borden & Peck; 1 female, Rancho Grande, Parque Nacional Henry Pittier, 1100 m, flight trap,

3 Apr 1967, M.E. Irwin; Bolivar: 37 males, 36 females, 10km S El Dorado, 200m, 17 Jul-7 Aug - 44 -

1986, B.D. Gill (CNCI, DEBU, MIZA); 5 males, 3 females, 20 km S El Dorado, 220 m, 20-23 Jul

1986, B. Gill; 1 male, 1 female, 22km S El Dorado, lowland rainforest, FIT, 25 Jun-12 Jul 1987, S.

& J. Peck; 13 males, 5 females, 33km S El Dorado, 220m, 2-7 Aug 1986, B.D. Gill (CNCI); 3 males, km40 Sta. Elena Icabaru Rd., 220m, 4-6 Aug 1986, B.D. Gill; 2 males, Quebrada de Jaspe,

19-20 Jul 1986, B. Gill; Lara: 2 males, Yacambu, 1200m, cloud forest, 7 May 1981, H.K. Townes;

Táchira: 1 male, 2 females, El Pinal, 57km SE San Cristobal, 1500ft, dung trap, 19-21 May 1974,

S. Peck.

Comments: Pectinosina prominens is the most frequently collected and widely distributed species in the Archiceroptera genus group, with a largely Neotropical distribution across a wide range of altitudes and latitudes. Two labels for P. prominens indicate the flies were found associated with dung rolling scarabs in Costa Rica: Grieg found a male around a dung ball rolled by Canthon moniliatus Bates in La Selva, and Gill found a male and female sitting on the elytra of

Sulcophanaeus velutinus (Murray) in San Vito. Other sphaerocerid genera are well known kleptoparasites of scarab beetles (see Sivinski et al. 1999 and references therein), but the observations of P. prominens with the scarabs may simply be of insects attracted to the same bait.

Pectinosina carro Paiero & Marshall, new species

Description: Length 1.8-2.8 mm. Eye height ~2.0-2.2× genal height.

Male Abdomen (Fig. 4.2): Sternite 5 with large discal area desclerotized; desclerotized area approximately circular, extending posteromedially to divide posterior margin; posterior margin on each side of break with numerous robust blunt setae. Epandrium hirsute with setae equal in length.

Surstylus bilobed; anterior lobe boot-shaped; posterior lobe simple, not as prominent as in P. prominens, with two robust setae on inner surface. Cercus acute, as long as surstylus, with apex abruptly posteriorly recurved. Distiphallus: first dorsal sclerites elongate, projecting beyond tip of acrophallus by more than length of sclerite before tip; acrophallus with mid-lateral projections spinose.

- 45 -

Female Abdomen (Fig. 4.3): Tergite 7 with posterior margin entire. Lateral sclerites of tergite 8 posteriorly with 3 ventral teeth. Paired spermathecae each with stems ~0.5× width of spermatheca and with sclerotized portion of common duct equal in length to stems; single spermatheca with sclerotized duct length ~2/3 width of spermatheca.

Type Material: Holotype (male, debu01082117, MIZA) and 5 Paratypes (4 males, 1 female): VENEZUELA: Bolivar: Quebrada de Jaspe, 19-20.vii.1986, B.Gill. Additional

Paratypes: COLOMBIA: Amazonas: 3 males, Leticia, 1 Mar 1974, pepper farm, dung, V. Nealis;

1 female, Leticia, 28 Feb 1974, V. Nealis. GUYANA: Rupununi: 1 male, 2 females, Kurupukari,

Essequibo River, 200’, 1° rainforest, dung traps, 9.x.1990, ROM 905042, B. Hubley (ROME).

VENEZUELA: Bolivar: 18 males, 3 females, km 40 Sta. Elena Icabaru Rd., 100 m, 4-6.viii.1986,

B. Gill (CNCI, DEBU, MIZA); 7 males, 3 females, same as previous except 220 m; 11 males, 5 females, same as previous except 1000m (MIZA, DEBU).

Comments: Male P. carro can readily be separated from P. prominens by the shape of sternite 5 and the surstylus, and by the relative length of the dorsal sclerite of the distiphallus. Females are more difficult to separate. Although there are usually only three teeth on each half of tergite 8 in P. carro females and four in P. prominens, some P. prominens also have only three teeth. Furthermore, the available material of P. carro is from a small number of collections and may not reflect the extent of intraspecific variation that occurs. The most consistent difference between females of these two species seems to be the relative length of spermathecal stems: the stems in P. carro are equal in length to the sclerotized portion of the common duct, while the stems in P. prominens are longer than the sclerotized portion of the common duct.

Etymology: The specific epithet is the Latin for “comb”, referring to the dense cluster of setae along the posterior margin of male sternite 5.

4.5 Chapter References

Cumming, J.M., and D.M. Wood. 2010. Adult morphology and terminology. Pp.9–63 In B.V.

Brown (ed.) Manual of Central American Diptera Volume 1. NRC Research Press.

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Duda, O. 1925. Die außereuropäischen Arten der Gattung Leptocera Olivier - Limosina Macquart

(Dipteren) mit Berücksichtigung der europäischen Arten. Archiv für Naturgeschichte,

Berlin, Abteilung A, 90(11)(1924): 5–215.

Marshall, S.A. and M. Buck. 2010. Sphaeroceridae (Small dung flies). Pp1165–1187 in Manual of

Central American Diptera. Eds. B.V. Brown, A. Borkent, J.M. Cumming, D.M. Wood,

N.E. Woodley and M.A. Zumbado. NRC Research Press, Ottawa, Ontario. 1442 p.

Marshall, S.A. and Y. Cui. 2005. Systematics of Robustagramma, a new genus of New World

Sphaeroceridae (Diptera). Zootaxa, 1026: 1–122.

Paiero, S.M. and S.A. Marshall. In prep. A revision of the genus Rudolfina Roháček

(Sphaeroceridae: Limosininae).

Roháček, J., S.A. Marshall, A.L. Norrbom, M. Buck, D.I. Quiros and I. Smith. 2001. World catalog

of Sphaeroceridae (Diptera). Slezské zemské muzeum, Opava, 414 pp. (also online at

http://www.uoguelph.ca/debu/catalog.htm).

Sivinski, J., S.A. Marshall and E. Petersson. 1999. Kleptoparasitism and phoresy in the Diptera. The

Florida Entomologist, 82(2): 79–197.

Shorthouse, D.P. 2010. SimpleMappr, an online tool to produce publication-quality point maps.

[Retrieved from http://www.simplemappr.net. Accessed January 29, 2015].

Smith, I.P., and S.A. Marshall. 2004. A review of the New World genus Pterogramma Spuler and a

revision of the Pterogramma sublugubrinum group (Diptera: Sphaeroceridae:

Limosininae). Contributions in Science, 499: 1–163.

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4.6 Pectinosina Figures

Figure 4.1. Pectinosina prominens: A) head and thorax, lateral view; B) wing; C) left mid tibia, dorsal view, showing distinctive chaetotaxy (arrows indicating setae placement that separate

Pectinosina from related genera).

- 48 -

Figure 4.2. Male terminalia of P. carro: A) abdomen, ventral view; B) sternite 4–5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, close up lateral; F) postgonite; lateral view; G) distiphallus and basiphallus, dorsal view; H) same, dorsolateral view; I) same, lateral view. A-I) from debu01082094

- 49 -

Figure 4.3. Pectinosina carro, female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-C) from debu01082100 and D) from debu01082102.

- 50 -

Figure 4.4. Pectinosina prominens, male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse portion of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, close up lateral; F) postgonite; lateral view; G) phallus, dorsal view;

H) phallus, postgonite and phallapodeme, dorsolateral view; I) same, lateral view. A-I) from debu00287406.

- 51 -

Figure 4.5. Pectinosina prominens, female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-C) from debu00190483, D) from debu01082507.

- 52 -

Figure 4.6. Distribution of Pectinosina prominens and P. carro.

- 53 -

CHAPTER 5 – A REVISION OF THE GENUS RUDOLFINA ROHÁČEK

(SPHAEROCERIDAE: LIMOSININAE)

This chapter is currently in the process of being submitted for publication and may not reflect all recent changes made previous to submission.

A revision of the genus Rudolfina Roháček (Sphaeroceridae: Limosininae)

STEVEN MARK PAIERO & STEPHEN A. MARSHALL

University of Guelph Insect Collection and Insect Systematics Laboratory, School of Environmental

Sciences, University of Guelph, Guelph, Ontario, Canada, N1G 2W1. Email: [email protected] and [email protected]

5.1 Abstract

The genus Rudolfina is revised and redefined with the description of the following new species: R. bucki, R. exuberata, R. howdeni, R. megepandria, R. newtoni, R. pauca, R. pilosa, R. remiforma, and R. tumida. Rudolfina is compared to closely related genera in the Archiceroptera genus complex, which in turn is recognized as part of a large, mostly Neotropical clade including

Robustagramma Marshall & Cui, Pterogramma Spuler, Aptilotella Duda, Bitheca Marshall,

Bromeloecia Spuler and Archiceroptera Papp.

5.2 Introduction

The genus Rudolfina Roháček 1987 was described (as Rudolfia Roháček 1982) for a single

Palaearctic species, Limosina rozkosnyi Roháček. Three species have since been described: R. digitata Marshall and R. cavernicola Marshall & Fitzgerald from North America (Marshall 1991,

Marshall and Fitzgerald 1997) and R. zhangi Su from China (Su et al. 2017). Marshall (1982) also - 54 - recognized several undescribed species of Rudolfina s. str. from Mexico, but deferred publishing descriptions of those species until adequate material was available to properly treat the Neotropical fauna and to determine the limits between Rudolfina and superficially similar Neotropical species in the Archiceroptera genus complex.

We here redefine the genus Rudolfina and describe nine new species following the examination of approximately 2,000 specimens of Rudolfina and over 6,000 specimens of other species in the

Archiceroptera genus complex.

Rudolfina is diagnosed by the following characters: mid tibia with 1 proximal anterodorsal seta, 1 medial anterodorsal seta, 1 distal anterodorsal seta and 1 distal posterodorsal seta; costa with 1 well- developed costagial seta; male sternite 5 posteromedially emarginate with darkened lobe or process on each side of the emargination; female tergite 8 tripartite; female epiproct medially weakened; female cercus strap-like with strong, flattened apical seta; and female abdomen with paired bisetose sclerites posterior to sternite 8. All Rudolfina except R. cavernicola have the female cercus partially fused with the epiproct. Most of these diagnostic characters are also defining characters that support the genus Rudolfina, as defined here, as a monophyletic group. The strongest synapomorphies for the genus are characters of the female terminalia, including a stout and generally upturned strap-like cercus with a strong flattened apical seta, an epiproct that is medially desclerotized except near the anterior margin, a middle sclerite of tergite 8 articulating with the anterior margin of the epiproct, and a pair of small bisetose sclerites posterior to the weakly sclerotized sternite 8. The strongly developed costagial seta is also considered a synapomorphy, as is the male sternite 5 with its characteristic pair of posteromedial lobes separated by a medial emargination.

Related and similar genera

All Rudolfina species will key out to “Rudolfia” in the key to Nearctic Sphaeroceridae by Marshall and Richards (1987) but they will key out as “Archiceroptera genus complex, in part” in Marshall

- 55 - and Buck’s (2010) key to Neotropical Sphaeroceridae. This treatment reflected uncertainty about the limits between Rudolfina and the many undescribed Neotropical species in the Archiceroptera genus complex. Like other members of the complex, Rudolfina species have two orbital setae, strong interfrontal setae, the costa ending at or slightly beyond R4+5, R4+5 straight or weakly curved to costa, and the mid tibia with an apical ventral seta. The Archiceroptera complex is part of larger group of Limosininae (including Aptilotella Duda, Archiceroptera Papp, Bitheca Marshall,

Bromeloecia Spuler, Pterogramma Spuler, and Robustagramma Marshall & Cui) characterized by an unusual process extending medially from the lower right margin of the epandrium. Within this group, Rudolfina resembles Archiceroptera, due to the general appearance of the highly modified female terminalia. Most Archiceroptera differ from Rudolfina species in having more than four strong dorsal mid tibia setae, but some have the pattern of four dorsal mid tibial setae typical of

Rudolfina. These species, however, differ markedly from Rudolfina in characters synapomorphic for

Archiceroptera (row of inclinate orbital setulae, M1 traceable to wing margin, CuA1 either not projecting beyond the apex of cell dm or with only short (< 1/2 dm-cu) stub vein present, male cercus either triangular or with prominent ventral process, female tergites 7 and 8 partially fused, female sternite 8 divided into lateral triangular sclerites, epiproct completely divided) and in the lack of other characters synapomorphic for Rudolfina species.

Biology

Roháček (1987) recorded R. rozkosnyi from dung, mud, and decaying vegetation, but most of the new species considered here were collected in dung traps. Larvae remain unknown.

Distribution

Rudolfina has a mostly western Nearctic montane distribution, with high endemism in the southwest and into the mountains of Mexico (Sierra Madre del Sur, Sierra Madre Oriental, and Sierra Madre de Chiapas). Two widely separated species occur in the Palaearctic region (R. rozkosnyi and R. zhangi) and one species (R. exuberata) is widespread at low elevations from the southern United

- 56 -

States to South America. Other than R. exuberata, no true Rudolfina are known from south of

Guatemala. Other Neotropical species previously treated as Rudolfina are discussed below.

5.3 Materials and Methods

Most specimens were collected into fluid, stored in alcohol, and later dried and point-mounted.

Abdomens of some specimens were removed and cleared in hot 10% potassium hydroxide for 6–10 minutes before being neutralized with 10% acetic acid for 10 minutes, rinsed in deionized water, and then placed into glycerin for examination. Dissected genitalia were stored in glycerin-filled microvials, and pinned below the specimen.

Species descriptions. All label data were presented in a consistent manner, not verbatim from the labels; in a few cases, obvious spelling errors were corrected. Short-forms or abbreviations used on specimen labels are normally interpreted and given in full. Geographical coordinates are given only if present on the original label. All specimens were given unique identifiers and their collection data were captured within the University of Guelph Insect Collection database; these are not repeated in the text except for holotypes or for imaged specimens. The specimen data will ultimately be hosted on Canadensys. Collection data for paratypes and other specimens examined were organized alphabetically by country, state/province, and locality name. Distribution maps are given for all

New World species (Fig. 5.22) and were generated using SimpleMappr (Shorthouse 2010).

Terminology. The terminology for external morphology largely follows Cumming and Wood (2010) with a few modifications given below; terminology for male and female genitalia follows Smith and

Marshall (2004) with modifications from Cumming and Wood (2010). Figures 1–4 illustrate head chaetotaxy, wing venation, male and female genitalia. Seta(e) and setula(e) are large and small

(respectively) socketed macrotrichia. The CuA1 and M1 stub veins are the short portions of these veins that project distally beyond cell dm. The subanal plate is the portion of the epandrium below the anal opening. Female tergite 8 is tripartite, with the medial part free, and posteriorly articulating

- 57 - with the epiproct; in some species, this medial part is poorly sclerotized (e.g., R. pauca). The female abdomen has a pair of small bisetose sclerites posterior to sternite 8. These bisetose sclerites are homologous with the posterior portion of sternite 8 in related taxa (e.g., Pterogramma and

Robustagramma; see Smith & Marshall (2004) and Marshall & Cui (2006)) and reflect a general desclerotization of sternite 8, leaving a distinct transverse anterior portion and the paired posterior sclerites. Body length was measured from the anterior portion of the frons to the tip of the abdomen.

Depositories. Material examined for this study is deposited in the following institutions: CASC

(California Academy of Sciences, San Francisco, California, U.S.A.), CNCI (Canadian National

Collection, Ottawa, Ontario, Canada), DEBU (School of Environmental Sciences, University of

Guelph, Guelph, Ontario, Canada); FMNH (Field Museum of Natural History, Chicago, Illinois,

U.S.A.); INBC (Instituto Nacional de Biodiversidad, Santo Domingo de Heredia, Costa Rica; now part of the Museo Nacional de Costa Rica); MIZA (Museo del Instituto de Zoología Agrícola

Francisco Fernández Yépez, Universidad Central de Venezuela, Maracay, Venezuela); MNNC

(Museo Nacional de Historia Natural, Santiago, Chile); MZSP (Museu de Zoologia, Universidade de São Paulo, São Paulo, São Paulo, Brazil); NMNH (National Museum of Natural History,

Smithsonian Institute, Washington, D.C., U.S.A.); QCAZ (Departamento de Biología, Pontífica

Universidad Católica del Ecuador, Quito, Ecuador); ROME (Royal Ontario Museum, Toronto,

Ontario, Canada); UASC (Museo de Historia Natural Noel Kempff Mercado, Santa Cruz de la

Sierra, Bolivia); UNAM (Universidad Nacional Autonoma de Mexico, Mexico City, Mexico).

Specimens are deposited at DEBU unless otherwise noted.

Illustrations and photography. Microphotographs of male and female genitalia were obtained using a Canon PowerShot S5IS mounted on a Leitz Laborlux 11 compound microscope. Series of images were aligned and combined using Zerene Stacker version 1.04 (Zerene Systems LLC, Richland,

WA, U.S.A.) with the DMax algorithm. Additional editing with Adobe Photoshop CS5 (Adobe, San

Jose, California, U.S.A.) was done to enhance clarity and visibility of the genitalic characters. - 58 -

Photographic plates were supplemented with drawings previously prepared for an earlier manuscript. All plates list the unique specimen identifier for the specimen(s) photographed.

5.4 Analysis

A character matrix (Table 1) was generated using Mesquite (version 3.10; Maddison and Maddison

2011), and exported for analysis in TNT (Goloboff et al. 2008) using Traditional Search, with 10 random seeds and 5000 replications with the tree bisection re-connection (TBR) swapping algorithm. Trees were optimized in WINCLADA (Nixon 2002).

The following 36 morphological characters used in the phylogenetic analysis are organized by body region and sex. Character states were polarized using the hypothetical groundplan of the

Archiceroptera genus-complex as an outgroup. The three multistate characters (1, 4 and 9) were treated as ordered. Rudolfina zhangi is only recently described and no material was available for study. This species therefore was excluded from the phylogenetic analysis although the description suggests that it is closely related to R. rozkosnyi, the only other Palaearctic species in the genus.

Table 1 gives the character states for each species.

Character states

Head

0) Eye height:genal height ratio: (0) 1:1; (1) 1.5:1; (2) 2:1; (3) 2.5:1. The ratios of eye height

to genal height are considered here to be a linear transformation series, reflecting the

relative size of the eye.

Thorax

1) Acrostichal setulae – number of rows anterior to suture: (0) 6 rows; (1) 8 rows.

Wings

2) Costagial seta: (0) regular, comparable to nearby seta; (1) enlarged, distinctly longer than

nearby seta.

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3) Costagial seta length: (0) short, apex not surpassing humeral break; (1) long, apex

surpassing humeral break but not reaching subcostal break; (2) very long, apex reaching or

surpassing subcostal break. The length of the seta is treated as linear transformation series.

Legs

4) Mid tibia (male) with apical ventral setal comb: (0) well developed; (1) reduced.

5) Mid femur (male) – ventral cluster of setae at base: (0) well developed, with 5 or more

robust setae present; (1) reduced, with < 5 robust setae present

Male Abdomen

6) Sternite 5 – posterior lobes: (0) absent; (1) present.

7) Sternite 5 – medial process: (0) absent; (1) present.

8) Sternite 5 – depth of posteromedial emargination: (0) < 1/4 length of sclerite or more; (1)

1/3–1/4; (2) > 1/4. The length (depth) of the emargination is treated here as a linear

transformation series.

9) Sternite 5 –clusters of setae flanking posteromedial emargination: (0) absent; (1) present.

10) Epandrium – prominence: (0) regular, equal in width to tergite 5; (1) swollen, wider than

tergite 5.

11) Epandrium – subanal plate: (0) incomplete; (1) complete.

12) Subepandrial sclerite: (0) transverse; (1) arched.

13) Cercus – development: (0) well developed, prominent; (1) reduced.

14) Cercus – general shape: (0) ovoid; (1) elongate conical.

15) Cercus – projecting posteriorly: (0) no; (1) yes.

16) Cercus – distal seta: (0) absent; (1) present.

17) Surstylus – posterior lobe: (0) rounded, not projecting; (1) elongate, extending posteriorly.

18) Surstylus – posterior lobe with flattened/modified seta on posterior surface: (0) absent; (1)

present

19) Surstylus – anterobasally distinctly rounded: (0) no; (1) yes.

20) Surstylus – anterior laminate lobe present: (0) no, (1) yes.

21) Surstylus – anterior laminate lobe modified: (0) no; (1) yes. - 60 -

22) Postgonite – shape at mid length: (0) parallel sided; (1) narrowed.

23) Postgonite – apical morphology: (0) simple; (1) with distinct apical setula.

24) Distiphallus – dorsal sclerite with distinct swellings along length: (0) absent; (1) present.

Female Abdomen

25) Median part of tergite 8: (0) absent; (1) present. This character occurs in several related

genera and is coded as both present and absent in the outgroup.

26) Median part of tergite 8 – shape: (0) wider than long; (1) longer than wide.

27) Sternite 8: (0) well developed, evenly sclerotized; (1) weakly sclerotized.

28) Bisetose sclerites posterior to sternite 8: (0) absent; (1) present.

29) Epiproct: (0) entire; (1) medially desclerotized.

30) Epiproct – anterior margin: (0) rounded (1) straight.

31) Epiproct: (0) anteriorly truncate or rounded; (1) anteriorly produced.

32) Cercus – shape: (0) ovoid; (1) strap-like.

33) Cercus – development: (0) regular; (1) reduced.

34) Cercus – dorsal chaetotaxy: (0) evenly hirsute; (1) glabrous.

35) Cercus – anterolaterally fused with epiproct: (0) no; (1) yes.

36) Cercus – apical seta: (0) regular; (1) flattened.

5.5 Results of Phylogenetic analysis

Six most parsimonious trees were generated, summarized here as a strict consensus tree (Fig. 5.4) and a majority rules consensus tree (Fig. 5.6). Characters were optimized on one of the equal length trees (Fig. 5.7).

The strict consensus tree supports Rudolfina cavernicola as the sister taxon to the remaining species, which form a monophyletic group characterized by the fusion of the female cercus with the posterolateral corner of the epiproct, and by other characters of the male cercus and surstylus. This tree suggests a New World origin for Rudolfina. No specimens of the Chinese species R. zhangi were available for examination, so it was not included in the phylogenetic

- 61 - analysis. Based on the described characters of the male genitalia (sternite 5, surstylus, cercus) it is probably the sister species to R. rozkosnyi. The Palaearctic clade (R. zhangi + R. rozkosnyi) is probably the sister group to the rest of Rudolfina except R. cavernicola.

All species other than R. cavernicola appear to form a clade characterized by the elongated medial part of tergite 8, and within that clade R. rozkosnyi, R. digitata and R. tumida appear to be basal lineages predating the origin of a clade comprising the Mexican-Guatemalan species. This largely Mexican clade can be recognized by the loss of the laminate anterior lobe of the male cercus and the absence of dorsal swellings on the dorsal sclerite of the distiphallus. In the strict consensus tree this clade is largely unresolved, with only the R. exuberata group standing out as strongly supported. The R. exuberata group (including R. exuberata, R. remiforma, and R. pauca) is characterized by the small elongate male cercus, tulip-shaped epiproct, and a reduction of the female cercus. Rudolfina remiforma and R. pauca are known from only a few localities at higher altitudes, as is typical of the genus, but the widespread R. exuberata occurs at much lower altitudes than its more localized congeners.

Molecular data:

Five Rudolfina species were sequenced for COI ("barcoded") at the Biodiversity Institute of Ontario

(University of Guelph, Guelph, Ontario) but only the North American R. digitata and the European

R. rozkosnyi yielded high quality (> 600bp) sequences. Unsurprisingly, these similar species came out next to one another in a Maximum Likelihood Tree for the broader "epandrial process" group

(Chapter 3), and they were clustered with four Bitheca specimens. The miscellaneous

Sphaeroceridae on the Barcode of Life Database (http://www.boldsystems.org) with both high quality sequences and associated specimen images were examined for additional Rudolfina material; as a result, one specimen of R. exuberata was identified and included in the analysis. In the

Maximum Likelihood Tree, Rudolfina exuberata was recovered with members of the “Extension” species group, a currently unplaced species group in the Archiceroptera genus complex. Although this preliminary molecular analysis suggests that R. exuberata is closer to other members of the

- 62 -

Archiceroptera genus complex than to other Rudolfina, morphological synapomorphies strongly support our treatment of this abundant species as deeply embedded in Rudolfina.

5.6 Rudolfina Roháček 1987

Rudolfia Roháček, 1982 [junior homonym of Rudolfia Wilson, 1924]

Type species: Limosina rozkosnyi Roháček, 1975 by monotypy

Redescription:

Colour light to dark brown. Length 1.4–2.3 mm.

Head with 3–5 interfrontal setae (of equal length or the foremost shorter), 1 (rarely 2) inclinate orbital setula and 4–10 small orbital setulae inside and below the 2 strong exclinate orbital setae; ocellar triangle with pair of strong setae and 3–5 additional small setulae; outer vertical seta strong, exclinate; inner vertical seta inclinate; occipital and paravertical setae inclinate, well-developed; postocellar seta inclinate, weakly developed. Eye-to-gena height ratio variable between species

(1.5:1 to 3.5:1). Vibrissa strong. Gena with 1–2 strong subvibrissal setae and 4–9 smaller setulae.

Thorax: Surface pruinose. Postpronotal lobe with 2–3 setae, outer seta strong, inner seta(e) reduced.

Notopleural seta, 2 supra-alar setae and prescutellar dorsocentral seta strong. Acrostichal setulae in

4–8 rows, with 1 enlarged prescutellar acrostical setae (almost as long as dorsocentral).

Katepisternum with strong elongate posterior seta and reduced anterior seta.

Legs: Fore femur with 3–5 elongate setae dorsally (except R. exuberata). Fore tibia with 3–5 elongate setae ventrally. Mid femur with row of 3–10 anterodorsal setae extending from base, row of 2–5 dorsal setae on apical 1/4, and basal cluster of 4–21 small setae ventrally; males usually with additional ventral seta (often in ventrobasal cluster). Mid tibia with 4 dorsal setae (basal anterodorsal, medial anterodorsal, distal anterodorsal and distal posterodorsal); males with ventral comb of 4–13 setae on apical 1/2 or less (R. exuberata and R. remiforma with setae of ventral comb weakly developed); females usually with midventral seta (absent in R. megepandria). Hind tibia with small apical ventral spur.

- 63 -

Wing: Always fully developed, with wing tip reaching or exceeding the apex of the abdomen. Costa extending to or just beyond end of R4+5, and with single costagial seta > 2.0× length of nearby setae.

R4+5 slightly curved towards costa distally. Cell dm with short stub veins of M1 and Cu-A1 extending beyond dm-cu. Alula narrow, posterior margin straight.

Abdomen: Sternites and tergites well sclerotized and setose (posterior and lateral margins more densely setose). Male sternite 4 usually simple (rarely densely setose medially).

Male abdomen: Posterior margin of sternite 5 with lobe on each side of medial emargination (shape and size of emargination and lobes varies between species). Transverse (ventral) portion of sternite

6 narrow; straight or weakly arcuate. Ring sclerite (in the right membrane of segment 7, possibly derived from a spiracle) large and distinct. Epandrium setose, often with larger setae lateral to anal opening, and with right anteroventral corner drawn out into a finger-like extension that extends to the hypandrium. Male cercus usually distinct, fused with the epandrium (reduced and obscured beneath the epandrium in a few species; e.g. R. pilosa, R. remiforma). Hypandrium (Fig. 5.3) Y- shaped with posteromedial extension; posteromedial extension emarginate; hypandrial arms with posterior margin emarginate on distal half. Pregonite distinct, small, near anterior base of postgonite. Postgonite generally simple and slender, with 3–4 setulae on anterior margin but modified in some species; ejaculatory apodeme small and finger-like with small globular sperm pump, usually close to the basiphallus (easily lost during dissection); basiphallus simple (without an epiphallus); distiphallus with distinct elongate dorsal sclerite; acrophallus with dorsolateral lobes and a single ventral sac (often reduced).

Female abdomen: Tergite 8 apparently tripartite, with two lateral triangular sclerites and a medial sclerite (secondarily reduced in several species). Epiproct bare except for the usual pair of small setae and a few other scattered setulae; strongly sclerotized, and fused laterally with cerci (except in

R. cavernicola). Cercus with single flattened apical seta. Sternite 7 variable. Sternite 8 weak, transverse, covered in small setulae; pair of small, spinulose plates along posterior margin.

Hypoproct a very narrow, horseshoe-shaped band immediately below the cerci. Spermathecae (1 pair +1 single), generally disc-shaped or lenticular, with thin, long sclerotized ducts.

- 64 -

5.7 Key to the New World Rudolfina.

Accurate identification of Rudolfina species is largely dependent on examination of male sternite 5 and genitalic characters of both sexes; dissection may be required. Females of R. tumida, R. bucki,

R. pilosa and R. zhangi are unknown.

1. Males…2

- Females…14

2. Sternite 5 with dense clusters of setae on each side of posteromedial emargination (Fig.

5.14 and 5.18)…3

- Sternite 5 evenly setose, without distinct clusters of setae…4

3. Eye ~2.5× genal height. Sternite 4 medially with cluster of long setae (denser along

posterior margin; Fig. 5.18). Sternite 5 with triangular lobe on each side of triangular

medial emargination on posterior margin; emargination extending anteriorly ½ length of

sternite. Surstylus (in lateral view) boot-like with 4–6 long setae originating from median

knob on posterior surface; distal 1/3 evenly covered in small setulae. Postgonite with

distinct apical swelling…R. pilosa sp. n.

- Eye ~1.5× gena height. Sternite 4 evenly setose (Fig. 5.14). Sternite 5 with small nipple-

like lobe on each side of medial emargination on posterior margin; emargination extending

anteriorly almost to base of sternite. Surstylus (in lateral view) strap-like, elongate and

narrow; relatively bare except for small scattered setae. Postgonite simple apically,

uniformly narrow…R. newtoni sp. n.

4. Posterior margin of sternite 5 with elongate, parallel sided lobes (e.g., Fig. 5.9) on each

side of medial emargination; pair of long setulae on margin of desclerotized area adjacent

to base of the lobes. Length of M1 between crossveins dm-cu and r-m < 1.4× dm-cu…5

- Posterior margin of sternite 5 with an acutely angled lobe on each side of medial

emargination; emargination without long setulae. Length of M1 between crossveins dm-cu

and r-m usually > 1.5× dm-cu (if less, second costal sector < 0.35× third costal sector)…6

- 65 -

5. Second costal sector < 0.4× third costal sector. Mid tibia with ventral comb of seta weakly developed, with only 1 strong seta at midlength and 1 long preapical setae. Mid femur with single distinct ventrobasal setae…R. exuberata sp. n.

- Second costal sector~0.6× third costal sector. Mid tibia with ventral comb composed of

11–13 setae on apical ½. Mid femur with 4–5 small setae ventrobasally…R. pauca sp. n.

6. Sternite 5 with small medial triangular lobe (Marshall and Fitzgerald 1997, Fig. 5).

Surstylus strap-like, with base ~1.5× wider than distal margin; anterior margin weakly lamellate (Marshall and Fitzgerald 1997, Fig. 2)…R. cavernicola Marshall & Fitzgerald

- Sternite 5 without medial lobe. Surstylus variable but usually with long posterior lobe and small anterior lobe…7

7. Sternite 5 with broad (> 1/4 width of sternite) emargination between posterior lobes.

Subanal plate complete or incomplete…8

- Sternite 5 with small (< 1/5 width of sternite) emargination between lobes. Subanal plate complete…10

8. Second costal sector ~0.35× third costal sector. Subanal plate narrowly complete (Fig.

5.20). Sternite 5 posterior margin lateral to posterior lobes straight (Fig. 5.19). Cercus elongate, small. Surstylus with long, glabrous, oar-like posterior lobe and small rounded anterior lobe with 4–5 long setae on surface…R. remiforma sp. n.

- Second costal sector 0.8–1.0× third costal sector. Subanal plate broadly complete or incomplete. Sternite 5 posterior margin lateral to posterior lobes emarginate (e.g., Fig.

5.22). Cercus ovoid, large. Surstylus with posterior lobe variable; anterior lobe complex….9

9. Eye small, ~1.0× genal height. Length of M1 between crossveins dm–cu and r-m ~4.0× dm-cu. Epandrium swollen, distinctly wider than two preceding abdominal segments (Fig.

5.2). Subanal plate incomplete. Subepandrial sclerite distinctly arcuate….R. tumida sp. n.

- Eye larger, > 1.2× genal height. Length of M1 between crossveins dm-cu and r-m < 2.5× dm-cu. Epandrium regular, as wide as two preceding abdominal segments. Subanal plate broadly complete. Subepandrial sclerite transverse…10 - 66 -

10. Second costal sector equal to third costal sector. Surstylus with posterior lobe elongate, with irregular pectinate anterior surface (Marshall 1991, Fig. 4–5)…R. digitata Marshall

- Second costal sector < 1.0× third costal sector. Surstylus not as above, with posterior lobe either hoe-shaped or with 3 elongate processes…11

11. Eye 2.0–2.3× genal height. Mid tibia distinctly arcuate in anterior view. Length of M1 between crossveins dm-cu and r-m ~1.5× dm-cu. Sternite 5 posterior margin with tips of medial lobes not reaching level of posterior margin adjacent to medial emargination

(Roháček 1985, Fig. 1084). Surstylus with posterior lobe hoe-shaped (Roháček 1985, Fig.

1075)…R. rozkosnyi Roháček

- Eye ~1.3× genal height. Mid tibia weakly arcuate in anterior view. Length of M1 between crossveins dm-cu and r-m ~2.0× dm-cu. Sternite 5 posterior margin with medial lobes tips extending beyond level of posterior margin adjacent to emargination (Su et al. 2017, Fig.

1F). Posterior lobe of surstylus with 3 elongate projections (Su et al. 2017, Fig. 1C)…R. zhangi Su

12. Posteromedial emargination of sternite 5 with dark margin; emargination deep, extending anteriorly ~1/6 sternite length, and nearly closed posteriorly by inwardly directed lateral lobes (Fig. 5.11). Epandrium simple, not distinctly wider than preceding abdominal segments. Surstylus with anterior lobe small but well-developed, knob-like; posterior lobe elongate, clavate, with numerous long setae on distal third. Cercus clavate, almost as long as surstylus, projecting ventrally…R. howdeni sp. n.

- Posterior emargination of sternite 5 without dark sclerotized margin; emargination shallow (< 1/10 sternite length) and broadly open with short posteriorly projecting lateral lobes (Fig. 5.8 and 5.13). Epandrium swollen, distinctly wider than preceding abdominal segments. Surstylus with anterior lobe reduced and indistinct; posterior lobe either elongate and narrow (R. megepandria) or weakly clavate (R. bucki); setae more widely dispersed over apical 1/2. Cercus either elongate and projecting posteriorly, or small and not distinctly projecting …13

- 67 -

13. Sternite 5 with small nipple-like lobes on posterior margin; posterior margin lateral to lobes entire (Fig. 5.13). Epandrium (in lateral view) with dorsal surface as long as posterior surface; setae below anal opening regular, not elongate. Surstylus with posterior lobe narrow and elongate, weakly constricted on distal ¼, with small rounded swelling near midlength. Cercus elongate, projecting posteriorly. Postgonite apically acute…R. megepandria sp. n.

- Sternite 5 with lobes on posterior margin triangular, obtuse; posterior margin lateral to lobes emarginate (Fig. 5.8). Epandrium (in lateral view) with dorsal surface ~1/2 length of posterior surface; 4–6 pairs of long cruciate setae adjacent to cercus (usually obscuring cercus in caudal view). Surstylus with posterior lobe weakly clavate, with long thickened seta on posterior margin (near midlength). Cercus obscure, small, indistinct. Postgonite apically truncate…R. bucki sp. n.

14. Epiproct tulip-shaped, with narrow anterior elongation broadening near midlength into rounded posterior ‘bulb’ (Fig. 5.10, 5.17, and 5.20). Length of M1 between crossveins dm- cu and r-m < 1.5× dm-cu. Medial portion of tergite 8 small, weakly sclerotized. Cercus shorter than flattened apical seta…15

- Epiproct triangular or trapezoidal. Length of M1 between crossveins dm-cu and r-m >

1.5× dm-cu. Medial portion of tergite 8 distinct, well sclerotized. Cercus as long or longer than flattened apical seta…17

15. Sternite 7 with posterior margin entire. Spermathecae ovoid (Fig. 5.20)…R. remiforma sp. n.

- Sternite 7 with posterior margin broadly emarginate (Fig. 5.10). Spermatheca bilobed…16

16. Eye 2.0× genal height. Second costal sector < 0.5× third costal sector…R. exuberata sp. n.

- Eye 2.5× genal height. Second costal sector > 0.5× third costal sector…R. pauca sp. n.

17. Eye ~1.5× genal height. Tergite 8 posteromedially emarginate (Fig. 5.15). Epiproct triangular. Spermathecae mushroom-shaped…Rudolfina newtoni sp. n. - 68 -

- Eye ≥1.75× genal height. Tergite 8 posteriorly entire or entirely desclerotized at middle.

Epiproct either trapezoidal or anteriorly rounded. Spermathecae variable…18

18. Eye height 2.5× genal height. Length of M1 between crossveins dm-cu and r-m 3.0×

dm-cu. Median part of tergite 8 elongate, rectangular (Fig. 5.4)…Rudolfina megepandria

sp. n.

- Eye height ≤2.3× genal height. Length of M1 between crossveins dm-cu and r-m 2.0 dm-

cu. Median part of tergite 8 variable…19

19. Second costal sector shorter than third costal sector. Epiproct diamond shaped, with

anterolateral margins almost straight (Fig. 5.12)…R. howdeni sp. n.

- Second costal sector equal in length to third costal sector. Epiproct with anterolateral

margins broadly rounded…20

20. Medial part of tergite 8 wider than long, with posterior margin weakly emarginate

(Marshall and Fitzgerald 1997, Fig. 5). Epiproct triangular…R. cavernicola Marshall &

Fitzgerald

- Medial part of tergite 8 as long or longer than wide; posterior margin entire. Epiproct

diamond-shaped…21

21. Medial part of tergite 8 longer than wide (Marshall 1991, Fig. 1). Epiproct with surface

even, smooth; anterior margin broadly rounded…R. digitata Marshall

- Medial part of tergite 8 as long as wide (Roháček 1985, Fig. 1079). Epiproct with surface

wrinkled on posterior half; anterior margin weakly trilobed…R. rozkosnyi Roháček

5.8 Species descriptions (alphabetically organized)

Rudolfina bucki Paiero & Marshall n. sp.

Description: Length 1.6mm. Eye height 2.5× genal height. Head with 3–4 interfrontal setae and 4–5 small setae on inner margin of orbital plate. Gena with 2 strong setae and 7–8 smaller setulae.

Acrostichal setulae in 4–6 rows. Costagial seta extending to midpoint between humeral and

- 69 - subcostal break. Second costal sector 0.7–0.8× third costal sector; length of M1 between crossveins dm-cu and r-m 3.5× dm-cu; CuA1 stub vein ~3.0× M1 stub vein. Male mid femur with 5–7 strong ventral setae and a circular cluster of 5–10 weaker setae ventrobasally. Male mid tibia with ventral comb of 4–5 robust setae on apical 1/3, without an enlarged midventral seta.

Male abdomen (Fig. 5.8): Sternite 5 with pair of medial triangular lobes on posterior margin; distance between lobes > 1.0× basal width of single lobe; lobe length ~1/5 length of remainder of sternite. Transverse part of sternite 6 weakly arcuate. Epandrium strongly convex with irregular cluster 6–8 of long inclinate posterolateral setae (appearing fan-like in posterior view); subanal plate broad, complete. Surstylus with posterior lobe elongate, weakly clavate; apical half of posterior lobe covered in small setae and with 1 strong elongate seta on posterior margin. Cercus small, ovoid, with strong preapical seta. Postgonite apically truncate. Distiphallus (Fig. 5.8E–G): dorsal sclerite without distinct swellings.

Female: Unknown

Type Material: Holotype (male, debu01086238, FMNH) and 1 paratype (male): MEXICO:

Oaxaca: Jct. Mex. 175-Yuvila Rd., 4.1 mi. W, 9300 ft, oak-fir-pine forest, 8–19 Aug 1973, A.

Newton..

Etymology: The specific name is a patronym for Dr. Matthias Buck, a friend and previous worker on Sphaeroceridae.

Rudolfina cavernicola Marshall & Fitzgerald 1997

Distribution: Nearctic: USA (AZ, CO); Mexico (MEX, newly recorded here).

Description: See Marshall and Fitzgerald 1997.

Material Examined: The original type series from Kremmer’s Cave was re-examined along with the following material: MEXICO: San Luis Potosi: 1 male, Cueva de Cinquenta y Ocho, 5 km S

San Francisco, 40 km E San Luis Potosi, Municipio de Zaraqoza, 3000 m, 18 May 1972, Elliott,

Ralph & Lynn.

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Comments: This species is newly recorded from Mexico. The illustration of this species that accompanied the original description (Marshall and Fitzgerald 1997, Fig. 1) includes what appears to be a large seta coming off the anterior base of the surstylus; this is an unsocketed laminate lobe.

Rudolfina digitata Marshall 1991

Description: See Marshall (1991).

Distribution: Nearctic: Canada (AB, BC, YT, ON), U.S.A. (AK, CO, NH, NY, WY), Mexico

(MEX).

Specimens Examined: In addition to the original type material, the following material was examined: CANADA: AB: 1 female, Coleman, 24–26 Jul 1980, S.A. Marshall; 1 female, Hailstone

Butte, 60km W Nanton, dung cup, under cow parsnip, 21–23 Jul 1987, S.A. Marshall; BC:

Kootenay Land Distr.: 1 female, Ainsworth, Woodbury Creek, dung, 5 Jul 1980, S.A. Marshall;

Okanagan–Similkameen: 1 male, Mt. Kobau, 1760m, 29 May–3 Jun 1991, Blades & Maier; Peace

River Land Distr.: 1 male, Pink Mountain, marmot dung, dwarf willow, 16 Jul 1987, S.A. Marshall;

ON: 1 female, Agnes River, SP2, plot 18, Jul 1994, A.P. Applejohn; MEXICO: México: 2 females, Tenancingo, 3 mi SW, 2164 m, km 52 1/4, oak–juniper, human dung, 31 Aug–6 Sep 1971,

A. Newton; U.S.A.: NH: Coos Co.: 1 male, East Inlet Dam, 1 mi NE, F.I.T., 25 Jun–9 Jul 1986,

D.S. Chandler; NY: Greene Co., 1 male, Cairo, 1 Jul 1980, S.A. Marshall; WY: Sheridan Co.: 2 males, 1 female, Antelope Butte Rec. Area, 20.3 km W of Burgess Jct., along stream, pans, cow dung, 5–20 Aug 1990, J.E. Swann; 1 male, Black Mountain, off Hwy 14, pine forest, pans, cow dung, 5–20 Aug 1990, J.E. Swann; 2 males, 1 female, same as previous except: pines, lupines, pans,

5–20 Aug 1990, J.E. Swann; 1 male, same as previous except: pines, pans, cow dung, 5–20 Aug

1990, J.E. Swann.

Comments: Rudolfina digitata, the second species to be described in this monobasic originally

Palaearctic genus, is widely distributed in western North America and also occurs in a few eastern

North American sites, including Mount Washington, New Hampshire. Mt. Washington is known to have other species with disjunct western Cordilleran and eastern Appalachian distributions [e.g.,

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Oeneis meillsa (Fabricius) (Lepidoptera: Nymphalidae)], and the apparently disjunct population of

R. digitata on Mt. Washington suggests it may have once had a more extensive range. The other

New Hampshire locality is also one of two known localities for the rare monotypic sphaerocerid genus Volumosina Roháček and Marshall (Roháček and Marshall 2017), otherwise known from one old growth forest in Ontario.

Rudolfina exuberata Paiero & Marshall n. sp.

Description: Length 1.5–2.0 mm. Eye height 2.0× genal height. Head with 4 interfrontal setae and row of 5–6 small setulae on inner margin of orbital plate. Gena with 1 strong setae and 5–7 smaller setulae. Acrostichal setulae in 6–8 rows. Costagial seta extending to humeral break. Second costal sector 0.3–0.4× third costal sector. Length of M1 between crossveins dm-cu and r-m 1.2× dm-cu;

CuA1 stub vein ~3.0× M1 stub vein. Male mid femur with single ventrobasal seta. Male mid tibia with ventral comb reduced to 1 enlarged preapical seta and 1 smaller seta near midlength. Female mid tibia with midventral seta.

Male abdomen (Fig. 5.9): Sternite 5 with deep irregular posteromedial emargination flanked by blunt lobe on each side; emargination extending anteriorly 1/3 length of sternite. Transverse part of sternite 6 straight. Surstylus (in lateral view) bilobed; anterior lobe rounded and bare; posterior lobe rounded but apically flattened and with 5–10 setae. Cercus projecting posteriorly, with apical seta.

Postgonite simple, slightly sinuate, with small robust seta apically and 1–2 setulae along length.

Distiphallus (Fig. 5.9F–H): Dorsal sclerites without distinct swellings.

Female Abdomen (Fig. 5.10): Tergite 7 with posterior margin entire. Tergite 8 with middle part elongate, narrower anteriorly, posteriorly closely approximated with epiproct. Epiproct tulip-shaped, heavily sclerotized; longitudinally lightly sclerotized on apical half. Cercus flattened, with distal apical seta flattened. Sternite 7 wider than sternite 6; posterior margin with broad shallow emargination. Spermatheca bilobed, constricted near midlength; distal portion cup-shaped, basal portion ovoid and narrowed before duct junction; spermatheca stem as long as spermathecal width.

- 72 -

Type Material: Holotype (male, debu01087005) and 51 Paratypes (25 males 26 females; DEBU,

FMNH & CNCI): UNITED STATES OF AMERICA: Florida: Marion Co., Ocala National

Forest, creekside, dung, 14–18 Jun 1984, S.A. Marshall. Additional paratypes: ARGENTINA:

Misiones: 4 males, Puerto Iguazo, 5km E, FIT/pans/dung pans, 2–7 Feb 1992, S.A. Marshall.

BELIZE: Cayo: 4 males 2 females, Caves Branch, forest, dung, 23–29 Aug 1972, S. & J. Peck; 3 males, Belmopan, palm forest, carrion, 26–30 Jul 1972, S. & J. Peck; 2 males 3 females, San

Ignacio, Maya Mt. Lodge, 17°9'N, 89°4'W, Atta mound trail, 2 Jan 1991, S.A. Marshall; 1 male 2 females, San Ignacio, Maya Mt. Lodge, 17°9'N, 89°4'W, dung trap, 7–8 Jan 1991, S.A. Marshall; 1 male, San Ignacio, Maya Mt. Lodge, 17°9'N, 89°4'W, Malaise, 7 Jan 1991, S.A. Marshall; 1 male

11 females, Mountain Pine Ridge, Hidden Valley Inst., 2500 ft, broadleaf forest, dung traps, 10–15

Jan 1991, S.A. Marshall; 1 female, Mountain Pine Ridge, Hidden Valley Inst., 2500 ft, pine–jungle edge, sweep, 14 Jan 1991, S.A. Marshall; 1 male 1 female, Mountain Pine Ridge, Hidden Valley

Inst., 2500 ft, dung traps in pine, 14 Jan 1991, S.A. Marshall; 10 males 18 females, Mountain Pine

Ridge, Hidden Valley Inst., 2500 ft, grass–pine dung traps, 10–15 Jan 1991, S.A. Marshall; 2 males

1 female, Beave, 39mi. from Highway, swamp forest, dung trap, 6–10 Aug 1972, S. & J. Peck.

BOLIVIA: La Paz: 1 female, Heath River Wildlife Centre, ~21 km SSW Puerto Heath, 12°40'S,

68°42'W, treefall, yellow pans, 5–7 May 2007, Marshall & Kits; 2 males 1 female, Heath River

Wildlife Research Centre, 12°40'S, 68°42'W, treefall, yellow pans, 5–9 May 2007, Paiero & Kits

(UASC); 1 female, San Antonio, ca. 8 km S Mapiri, 15°20'56"S, 68°13'31"W, secondary forest, dung pans, 11 Apr 2001, S.A. Marshall. BRAZIL: Minas Gerais: 1 male 1 female, Lavras, 1km E, dung traps in ditch, 18–20 Feb 1990, S.A. Marshall; Paraná: 4 males 2 females, Londrina, carrion pan traps, 1–2 Feb 1990, S.A. Marshall (MZSP); 2 males 4 females, Londrina, Mata dos Godoy,

28–31 Jan 1990, S.A. Marshall (MZSP); 1 male 1 female, Curitiba, 30 km SE, BR 277, dung traps,

6–9 Feb 1990, S.A. Marshall; Rio de Janiero: 3 males 1 female, Tijuca Forest Res., Malaise, 1–28

Feb 1990, S.A. Marshall (MZSP); 1 male, Nova Friburgo, Sitio Edelweiss, Malaise, 26 Jan 1990,

S.A. Marshall; São Paulo: 1 male, Estación Biologica Boracea, dung trap, 2 Dec 2008, G.F.G.

Miranda (MZSP); 1 female, Sao Paulo, Jaragua, 8 Feb 1990, S.A. Marshall; 8 males 7 females, USP

Biology Station, human dung, 5–6 Feb 1979, R. Woodruff & J. Runnacles (MZSP). CHILE: - 73 -

Valparaíso: 2 female, La Campana National Park, 22 Nov 2006, S.A. Marshall (DEBU and

MNNC). COLOMBIA: Norte de Santander: 2 males 2 females, Cucuta Quebrada de Honda,

20mi. S, 2500 ft, carrion trap, 13–15 May 1974, S. Peck; 1 male 1 female, Chinacota, 2mi. N, 3000 ft, carrion, 12–14 May 1974, S. Peck. COSTA RICA: Alajuela: 1 female, Volcán Tenorio, N slope nr. Bijagua Biological Station, 700 m, pan traps in tree fall, 18 Jun 2000, Buck & Marshall; 1 female, Río Peñas Blancas, 700 m, 18 Aug 1986, L. Masner; 1 male, Florencia Forest, dung trap, 28

Feb 1980, H. Howden; Cartago: 13 males 14 females, Turrialba Catie, 600 m, 26 Feb 1980, H. &

A. Howden (CNCI, DEBU, and INBC; 6 males 9 females, Turrialba Catie, 600 m, 28 Feb 1980, H.

& A. Howden; 1 male, Turrialba, Catie, Florence Forest, 600m, cup traps, 28 Feb 1980, H. & A.

Howden; Guanacaste: 1 male, Guanacaste Cons. Area, Pitilla Fld. Stn., Malaise, 29 Jan 1996, J.

Noyes; Heredia: 1 female, La Selva, around dung ball rolled by Canthon moniliatus, 2 Feb 1990,

N. Grieg; Limon: 4 females, Estrella Valley, Pandora, carrion trap, 20 Feb 1984, H. Howden (CNCI and INBC); 1 male 1 female, Bribri, 4 km NE, 50 m, Dec 1989–Mar 1990, P. Hanson. CUBA:

Santiago: 1 female, Gran Piedra Met Radar, 1100 m, elfin forest, carrion traps, 6–17 Dec 1995, S.

Peck. ECUADOR: Napo: 2 males, Tiputini Biodiv. Stn., vic. Yasuní National Park, 0°38'S,

76°0'W, human dung pitfalls, 14–19 Feb 1998, D.C. Darling; 1 female, Yasuní National Park,

Yasuní Res. Stn., 0°38'S, 76°36'W, rain forest, Malaise trap, 3–20 Nov 1998, Pape & Viklund; 4 females, Jatun Sacha Res., 6 km E Misahuallí, 1°4'S, 77°37'W, 450 m, land slide in forest, Malaise trap, 30 Apr–7 May 2002, Buck et al. (QCAZ); 2 males, Tena, 500m, secondary rainforest, Malaise head, 21–27 May 1987, Brown & Coote; 3 males, Tena, 12 km SW, 500m, dung trap, 8–11 Aug

1976, S. Peck (QCAZ); 1 male, Rio Piocullin, S side, SW Puerto Napo, S. Limonchicta, 600m,

1*lowland rainforest, Malaise head, ROM 870020, 23–27 May 1987; Pichincha: 8 males 8 females,

Rio Palenque Stn., 47km S Sto. Domingo, 250m, dung, 17–25 Feb 1979, S.A. Marshall; 7 males 3 females, Rio Palenque Stn., 47km S Sto. Domingo, 250m, 17–25 Feb 1979, S.A. Marshall (QCAZ);

1 male, Rio Palenque, carrion, 25 Feb 1979, S.A. Marshall (QCAZ); 1 female, Rio Palenque, carrion, 27 Feb 1979, S.A. Marshall; 1 male, Palenque, day 3, trap 35, 24–25 Feb 1975, S. Peck; 13 males 23 females, Rio Palenque, dung, 25 Feb 1979, S.A. Marshall; 9 males 34 females, Rio

Palenque, dung, 27 Feb 1979, S.A. Marshall (DEBU and QCAZ); 1 male, Rio Palenque, dung trap, - 74 -

22–23 Feb 1976, S. Peck; 1 male, Rio Palenque, dung trap, 25–26 Feb 1976, S. Peck; 1 male, Rio

Palenque, 22 Feb 1976, S. Peck. GUATEMALA: Baja Verapaz: 1 male 2 females, Chilasco,

6.6km W, 1700m, dung, 30 May 1991, H. Howden; 1 male, Chilasco, 6.6km W, 1700m, dung, 30

May 1991, H. Howden; Guatemala: 1 male 1 female, Santa Catarina Pinula, 1840 m, dung traps,

11–13 Jun 1991, B.D. Gill; Sacatepéquez: 1 female, Antigua, 5 km SE, 14°32'14"N, 90°41'41"W,

2125 m, hardwood forest, Malaise trap, 10–13 Jun 2009; Zacapa: 1 male, La Unión, 3.5 km SE,

1500 m, cloud forest litter, 26 Jun 1993, R. Anderson; 3 females, La Unión, 3.5 km SE, 1500 m,

FIT, #128, 25–27 Jun 1993, Ashe & Brooks; Peten: 11 males 7 females, Tikal, dung trap, 28–30 Jul

1978, Helava & Kukal. GUYANA: Mazaruni–Potaro: 1 female, Takutu Mountains, 6°15'N,

58°55'W, window trap in montane rainforest near logging area, 8–10 Dec 1983, Perkins & Steiner;

Potaro–Siparuni: 1 female, Mount Wokomung, 5°6'35"N, 59°49'15"W, 1234m, 1° rainforest, human dung, pitfall trap, 27 Oct–1 Nov 2004, B. Hubley (ROME); Rupununi: 3 males 4 females,

Kurupukari, Essequibo River, 200 ft, 1° forest, dung traps, 9 Oct 1990, B. Hubley (ROME); 1 male

1 female, Kabocalli, Iwokrama Forest Res., 60 m, FIT, 3–5 Jun 2001, Brooks & Falin.

HONDURAS: Olancho: 1 female, Olancho, FIT, 23 May 1995, R. Cordire. MEXICO:

Campeche: 7 males 3 females, Chicanna, 10km W Xbuzi, 300 m, tropical seasonal forest, carrion traps, 12–14 Jul 1983, S. & J. Peck (CNCI and DEBU); 6 males 14 females, Escarcega, 53mi. E,

500 ft, tropical semi-evergreen, dung, 8–14 Aug 1971, A. Newton (DEBU and FMNH); 19 males 25 females, Escarcega, 87mi. E, 800ft., semi–evergreen, human dung, 8–14 Aug 1971, A. Newton

(DEBU and FMNH); Chiapas: 2 females, Bochil, 21mi. N, 5500 ft, pine, oak, Liquidambar, human dung, 18–24 Aug 1971, A. Newton; 1 female, Custepec, 4 km SE, 15°42'30"N, 92°55'51"W, 2100 m, cloud forest, Malaise trap, 20 May 2008; 2 females, Playón de la Gloria, 16°8'53"N,

90°53'48"W, 180 m, mature wet forest, Malaise trap, 25 May 2008 (UNAM); 1 females, Playón de la Gloria, 16°9'37"N, 90°54'7"W, 160 m, mature wet forest, Malaise trap, 25 May–24 Jun 2008

(UNAM); 2 males 4 females, Salto de Agua, 8 km SE, 17°30'45"N, 92°17'40"W, 60 m, 2° wet forest, Malaise trap, 14–17 Jun 2008 (UNAM); 2 females, Salto de Agua, 8 km SE, 17°30'58"N,

92°18'5"W, 100 m, wet forest edge, Malaise trap, 14 Jun 2008 (UNAM); 5 males 8 females,

Palenque, 100 m, rainforest, carrion trap, 6–9 Jul 1983, S. & J. Peck; 1 female, Parque Nacional - 75 -

Sumidero, 1000m, 1 Jun 1990, H. & A. Howden (CNCI); 1 male 2 females, Ocozocantha, 15mi.

NW, 2800 ft, rainforest, dung, A. Newton; 5 males 2 females, Ocozocoautla, 11mi NW, 3400 ft, oak–evergreen forest, human dung, 19–25 Aug 1971, A. Newton (FMNH); 6 males 4 females,

Palenque, 4 mi. S, 600 ft, rainforest, human dung, 7–15 Aug 1971, A. Newton; 21 males 20 females,

Palenque, 4mi S, 700 ft, rainforest, human dung, 7–15 Aug 1971, A. Newton (DEBU and FMNH);

1 male, Palenque, 80 m, 2° vegetation, FIT, 2–23 Jul 1983, S. & J. Peck; 1 male, Laguna Belgica,

16 km NW Ocozocoaulta, 970m, FIT, 13 Jun 1990, H. & A. Howden & Gill; Guerrero: 1 male 1 female, Iguala, 9 mi. NE, 4400 ft, deciduous forest, human dung, 29 Aug–4 Sep 1971, A. Newton

(FMNH); Oaxaca: 5 males 9 females, Valle Nacional, 5mi S, 1600 ft, tropical-oak-evergreen, dung,

20 Jul–1 Aug 1971, A. Newton; 3 males 2 females, Valle Nacional, 12mi S, 3200 ft, tropical montane forest, shrimp carrion, 22–31 Jul 1971, A. Newton; Pueblo: 1 male, Honey, 1mi. S, 6800 ft, Pinus, Quercus, human dung, 1–6 Aug 1971, A. Newton; Tabasco: 6 males 5 females,

Villahermosa, 46mi. SE, km74, 150 ft, 2° veg. rainforest, human dung, 8–15 Aug 1971, A. Newton

(FMNH); Tamaulipas.: 8 males 17 females, Municipio Casas, 47mi. E Cuidad Victoria, carrion pitfall trap, Nov–Dec 1986, R. Jones; Veracruz: 1 male, Catemaco, 33km NE, Los Tuxtlas

Biological Station, 160 m, ridge rainforest, FIT, 6 Jul–1 Aug 1983, S. Peck; 2 males 4 females,

Fortin, SW of Rio Metlas, 3250 ft, human dung, 13–18 Jul 1971, A. Newton (FMNH) ; 3 males 2 females, Teocelo, 10mi SW, 4400 ft, oak, wet, human dung, 11 Jul 1971, A. Newton (FMNH); 7 males 4 females, Sontecomapan, 8mi. NNW, rainforest, dung, Jul–Aug 1971, A. Newton; Yucatan:

1 female, Santo Roso, 9mi SE, km 137 1/4, 100 ft, tropical sub–deciduous forest, human dung, Aug

1971, A. Newton. PARAGUAY: Caazapá: 18 males 12 females, Hermosa, San Rafael Reserve, prop. Lopez family, 26°18'29"S, 55°45'3"W, 80 m, FIT, 1–3 Dec 2000, Z.H. Falin; 2 males 1 female, Hermosa, San Rafael Reserve, prop. Lopez family, 26°19'15"S, 55°44'55"W, 90 m, FIT, 3–

6 Dec 2000, Z.H. Falin; Itapúa: 1 female, Karonay, 17 km W San Rafael Reserve, 26°45'53"S,

55°50'37"W, 90–110 m, FIT, 18–21 Nov 2000, Z.H. Falin. PERU: Junin: 2 females, Pampa

Hermosa lodge, 22 km N San Ramon, 10°59'18"S, 75°25'30"W, 1220 m, F.I.T., 24–27 Nov 2007,

D. Brzoska; Cusco: 1 female, Cock–of–the–Rock Lodge, NE Paucartambo, 13°3'18"S,

71°32'42"W, 1120 m, FIT, 4–9 Nov 2007, D. Brzoska; Loreto: 1 male, Teniente López, FIT, 24 Jul - 76 -

1993, R. Leschen; Madre de Dios: 3 males 3 females, Zona Reserva Manu, Pakitza, 11°57'S,

71°17'W, 400 m, Malaise trap, 13–18 Feb 1992, B. Brown & D. Feener; 3 females, Pantiacolla

Lodge, Alto Madre de Dios River, 12°39'18"S, 71°13'54"W, 420 m, FIT, 14–19 Nov 2007, D.

Brzoska; 4 male 5 females, Amazonas Lodge, N of Atalaya, 12°52'12"S, 71°22'36"W, 480 m, FIT,

10–13 Nov 2007, D. Brzoska; 1 female, Los Amigos Biological Station, Malaise, 3–13 Jun 2006,

Paiero & Klymko; Huaral: 1 male, Chancay, river valley, 15 Mar 1951, Ross & Michelbacher

(CASC). TRINIDAD AND TOBAGO: Tobago: 1 male, Roxborough, 10km NE, Gilpin Trail, 450 m, rainforest, carrion traps, 26–30 Jun 1993, S. & J. Peck; Trinidad: 1 male, U. Santa Cruz,

Gasparillo, grassland/forest edge, Malaise trap, 15 Nov 1987, R. Borneo. U.S.A.: AL: Baldwin Co.:

22 males, 32 females, Bon Secour Natl. Wildlife Ref., 30°14'48"N, 87°49'45"W, dung traps in oak,

5–7 Jun 1994, S.A. Marshall; 2 females, Bon Secour Natl. Wildlife Ref., 30°14'48"N, 87°49'45"W, mushroom trap on oak, 5–7 May 1994, S.A. Marshall; Jackson Co.: 2 males, Paint Rock, dung, 7–

13 Jul 1973, S. Peck; AR: Johnson Co.: 1 male, Haw Creek Falls, 22 May 1991, B.J. Sinclair;

Logan Co.: 1 male 2 females, Ozark Natl. For., Magazine Mt., mushroom baited pans, 23 May

1991, J.E. Swann; Madison Co.: 2 females, Brashers, 3 mi S, 1600 ft, hardwood forest, dung, 19

Jun–12 Jul 1972, A. Newton; Polk Co.: 1 male, Rich Mt., 13 mi NW Mena, 2800 ft, mesic oak– hickory, 1–3 Jun 1979, S. & J. Peck; Washington Co.: 4 males 3 females, Devils Den State Park, 3 mi S, oak–hickory, 28–31 May 1979, S. & J. Peck; 1 male 3 females, Devils Den State Park, Devils

Den Cave, 28–31 May 1979, S. & J. Peck; FL: Alachua Co.: 1 female, Gainesville, hardwood forest, Malaise trap, 20–30 Nov 1986, W.R.M. Mason; 1 female, Rd. 176, carrion trap, 10 May

1983, K.W. Vick; 1 female, Gainesville, DPI, Malaise trap, 25–27 May 1983, D.S. Chandler; 1 female, High Springs, on dung, 15–16 Dec 1999, S.A. Marshall; 1 female, Gainesville, 12 Oct 1976,

Choate & Woodruff; 16 males 29 females, Gainesville, Hogtown Creek, 12 Oct 1976, Choate &

Woodruff; Clay Co.: 2 males 5 females, Gold Head Branch State Park, carrion, 4–14 Apr 1971, A.

Newton; 2 males 1 female, Gold Head Branch State Park, dung, 14 Apr 1971, A. Newton; Collier

Co.: 1 female, (no locality given), 28 Dec 1979, S.A. Marshall; 3 females, Collier Seminole State

Park, pine, palm, human dung, Apr 1971, A. Newton; Dade Co.: 1 male 6 females, Everglades

National Park, 1.5km NW Royal Palm, hardwood hammock forest, Malaise–FIT, 1 Nov 1984–3 - 77 -

Mar 1985, S. & J. Peck; 7 females, Everglades National Park, 1.5km NW Royal Palm, hardwood hammock forest, Malaise–FIT, 15 Nov 1985–24 Feb 1986, S. & J. Peck; 3 males 4 females,

Everglades National Park, 1.5km NW Royal Palm, hardwood hammock forest, Malaise–FIT, 2

May–2 Aug 1985, S. & J. Peck; 5 females, Chekika State Recreation Area, 50km SW Miami,

Grossman Hammock Forest, Malaise–FIT, 1 May–2 Aug 1985, S. & J. Peck; De Land Co.: 1 female, Hwy. 40, nr. Barberville, mushroom trap, 18–20 Jun 1984, S.A. Marshall; Hendry Co.: 1 male 1 female, La Belle, Capt. Hendry Rd., pine, hardwood nr. river, human dung, Apr 1971, A.

Newton; Hernando Co.: 1 male 3 females, Withlacoochee State Forest, 1mi. W Croom, pine & hardwood, human dung, Apr 1971, A. Newton; 1 male, Croom, 1mi. W, mixed forest, Apr 1971, A.

Newton; Highlands Co.: 1 male 1 female, Lake Placid, north shore, palmetto, dung trap, 14–16 Dec

1985, S.A. Marshall; 2 males 6 females, Archbold Biological Station, scrub, dung, 17–21 Dec 1985,

S.A. Marshall; 2 females, Archbold Biological Station, scrub, dung pitfall, 12–16 Dec 1985, S.A.

Marshall; 1 male, Archbold Biological Station, scrub–sand, dung pan, 12–16 Dec 1985, S.A.

Marshall; 6 males 12 females, Archbold Biological Station, dung pan, 15–19 Dec 1985, S.A.

Marshall; 3 males 8 females, Highlands Hammock State Park, Cypress Swamp, dung pan, 15–21

Dec 1985, S.A. Marshall; 1 female, Archbold Biological Station, mushroom bait on sand, 12–18

Dec 1985, S.A. Marshall; 1 male, Highland Hammock State Park, pans/maggot bait, 13–17 Apr

1989, S.A. Marshall; 1 male 6 females, Highlands Hammock State Park, orange grove, pig dung,

14–18 Jun 1982, Woodruff & Rench; Lake Co.: 30 males 41 females, Howey–in–the–Hills, 1mi. W, mixed hardwood forest, human dung, Apr 1971, A. Newton; 15 males 15 females, Howey–in–the–

Hills, dung, Apr 1971, A. Newton (CASC, CNCI, NMNH & DEBU); Liberty Co.: 1 male 8 females, Torreya State Park, pig dung trap, 6 Jun 1982, R.E. Woodruff; Marion Co.: 3 males 6 females, Ocala National Forest, hardwood swamp, dung trap, 11–15 Jun 1984, S.A. Marshall; 2 males 2 females, Ocala National Forest, oak grove, mushroom trap, 8–11 Jun 1984, S.A. Marshall; 6 males 7 females, Ocala National Forest, dung, Apr 1971, A. Newton (CNCI, NMNH & DEBU); 3 males, Ocala National Forest, nr. Lynne, palm–oak, dung trap, 20–24 Jun 1984, S.A. Marshall; 3 males 12 females, Ocala National Forest, Oklawaha Swamp, dung trap, 11–12 Jun 1984, S.A.

Marshall; 2 females, Ocala National Forest, Rd. 65, sand–pine, human dung, 18–20 Jun 1984, S.A. - 78 -

Marshall; 1 male 6 females, Ocala National Forest, Rd.65, 1.5mi. W St. Rd. 19, dung, 15–16 Mar

1984, R.E. Woodruff; 1 male, Ocala National Forest, Silver Springs Woods, oak, FIT, 6–23 Jun

1984, S.A. Marshall; 1 female, Ocala National Forest, Silver Springs Woods, sand–pine, human dung, 12–14 Jun 1984, S.A. Marshall; 2 males, Ocala National Forest, Silver Springs Woods, carrion FIT, 11–15 Jun 1984, S.A. Marshall; 1 male 1 female, Ocala National Forest, Silver Springs

Woods, FIT, 6–22 Jun 1984, S.A. Marshall; 2 females, Ocala National Forest, Silver Springs

Woods, 10–15 Jun 1984, S.A. Marshall; 1 male, Ocala National Forest, Zay Prairie, carrion trap,

14–18 Jun 1984, S.A. Marshall; 7 males 17 females, Rte. 316, nr. Eureka, pig dung trap, 15–16 Mar

1984, R.E. Woodruff; Monroe Co.: 1 male, Everglades National Park, Royal Palm Hammock, hammock forest, Malaise–FIT, 1 Nov 1984–3 Mar 1985, S. & J. Peck; Nassau Co.: 8 males 16 females, golf course nr. Amelia, oak forest, dung, 3–15 Apr 1971, A. Newton (NMNH & DEBU); 6 males 10 females, Jacksonville, Cary State Forest, dead squirrel, pan trap, 10–18 Apr 1989,

Marshall & Swann; 19 males 16 females, Amelia, nr., dung, 3–15 Apr 1971, A. Newton (NMNH &

DEBU); Okaloosa Co.: 1 male 1 female, Blackwater River National Forest, 1mi. N Holt, Turkey

Oak, human dung trap, 23 Oct 1978, L. Stange; Polk Co.: 1 male, Rte. 27, 7mi. N Rte. 14, pig dung trap, 2–3 Nov 1983, R.E. Woodruff; 3 males 1 female, Lake Wales, 2mi. N, pig dung trap, 25–29

Apr 1983, R.E. Woodruff; Putnam Co.: 4 males 23 females, Ocala National Forest, Johnson Field

Campground, mixed hardwood, dung trap, Apr 1972, A. Newton; Sarasota Co.: 5 males 3 females,

Myakka River State Park, pig dung trap, 21 May 1982, R.E. Woodruff; 3 males 3 females, Myakka

River State Park, pig dung trap, 22–23 May 1982, R.E. Woodruff; Volusia Co.: 3 males 1 female,

Tomoka State Park, dung trap, 20 Jun 1984, S.A. Marshall; 9 males 13 females, Tomoka State Park, mushroom trap, 20 Jun 1984, S.A. Marshall; Hernando/Sumter Cos.: 1 female, Withlacoochee St.

Forest, Croom Reserve, dung, 12 Apr 1989, K.N. Barber; 2 males 1 female, Withlacoochee St.

Forest, Croom Reserve, dung vacuum, 12 Apr 1989, K.N. Barber; GA: Charlton Co: 15 males 20 females, Folkston, 14mi. N, pan trap, human dung, 10–18 Apr 1989, J.E. Swann; Clinch Co.: 1 male

1 female, US 441, 8mi. S Fargo, dung trap, 5–25 Jun 1984, S.A. Marshall; Rabun Co.: 2 males,

Chatahoochee State Forest, US 441 N of Turnerville, mushroom trap, 5–25 Jun 1984, S.A.

Marshall; Wilkinson Co.: 8 males 20 females, Big Sandy Creek, 4mi. S Irwinton, dung trap, 5–25 - 79 -

Jun 1984, S.A. Marshall; Wilkinson Co.: 13 males 10 females, Big Sandy Creek, 4mi. S Irwinton,

FIT nr. dung, 5–25 Jun 1984, S.A. Marshall; LA: East Baton Rouge Par.: 1 male, 1 female, swine carrion pitfall, 9 Apr 1999, E.J. Watson; Grant Parish: 3 males 1 female, Stuart L. Campground,

18km N Alexandria, forest, FIT, 19 May–17 Aug 1983, S. & J. Peck; Jackson Co.: 1 male 2 females, Paint Rock National Cave, 1.5mi. SE, rat dung, Berlese #77, 9 Jul 1967, S. Peck; MO:

Texas Co.: 1 female, Licking, 10.5mi. NW, unnamed cave #1, 14 May 1980, J.E. Gardner; MS:

Claiborne Co.: 5 males 8 females, Owen's Creek, mi. 52,19km NE Port Gibson, forest, FIT, 18

May–16 Aug 1983, S. & J. Peck; Forrest Co.: 37 males 35 females, Sweet Bay Bog, 6mi. W

Wiggins, sphagnum, dung trap, 5–8 May 1994, S.A. Marshall; Scott Co.: 1 male, Forest, pans in prairie edge, 5–9 May 1994, S.A. Marshall; NC: Jackson Co.: 2 females, Cullowhee, Cane Creek,

2300ft, riparian dung trap, 5–25 Jun 1984, S.A. Marshall; 1 male, Cullowhee, FIT, 5–28 Jun 1984,

S.A. Marshall; Samson Co.: 1 male 5 females, Falcon at I(%, pig dung, 28 Sep–4 Oct 1983, R.

Woodruff; NM: Eddy Co.: 1 female, Carlsbad, 30mi. WSW, Sitting Bull Falls, 4600 ft, dung, 23–27

Jul 1975, S. Peck; OK: Lattimer Co.: 1 male, Red Oak, 5 mi W, dung trap, 5–11 Jun 1977, K.

Stephan; SC: Barnwell Co.: 1 male 1 female, Barnwell State Park, near lake, dung trap, 10–18 Apr

1989, T.A. Wheeler; 6 males 8 females, Barnwell State Park, mushroom traps in oak forest, 10–18

Apr 1989, S.A. Marshall; Colleton Co.: 1 male 3 females, Colleton State Park, pig dung trap, 27

Sep–5 Oct 1983, R.E. Woodruff; Georgetown Co.: 1 male 4 females, Hobcaw Barony, Belle Baruch

Marine Field Lab, Crabhaul Rd. nr. Picnic Rd., 33°21'22"N, 79°12'45"W, cypress swamp, yellow pans, 28–29 Apr 2004, S.A. Marshall; 21 males 36 females, Hobcaw Barony, Belle Baruch Marine

Field Lab, Crabhaul Rd. nr. Picnic Rd., 33°21'22"N, 79°12'45"W, on dung, 29–30 Apr 2004,

Cheung & Macleod; Neeses Co.: 1 male, Orangeburg, 13mi. W, 27 Mar 1980, S.A. Marshall; TN:

Henderson Co.: 1 male, Natchez Trace State Park, 1000ft., dung, 18 Jun–13 Jul 1972, A. Newton;

TX: Angelina Co.: 3 females, Angelina Natl. For., Boykin Cemetery, spring fen, dung, 13–17 Jun

1993, S.A. Marshall; 1 male 6 females, Angelina Nat. Forest, Bouton Lake Campground, dung/vac traps, 12–14 Jun 1993, S.A. Marshall; Brazos Co., 1 male, College Station, Lick Creek Pk.,

30°33'44"N, 96°12'53"W, bottomland forest nr. creek, Malaise trap, 5–9 Apr 2000, M. Buck; 1 male, College Station, Lick Creek Pk., 30°33'44"N, 96°12'53"W, post oak savanna by creek, - 80 -

Malaise trap, 5–9 Apr 2000, M. Buck; Montgomery Co.: 1 female, Montgomery, 4.5mi. N, pine/black walnut forest, FIT, 2 May–17 Jun 1987, R.S. Anderson; San Jacinto Co.: 4 males 3 females, Coldspring, Double Lake Campground, FIT, 22 May–16 Aug 1983, S. & J. Peck; 3 females, Double Lake Campground, 5km S Coldspring, forest, carrion trap, 22–24 May 1983, S.

Peck; 4 males 14 females, Double Lake Campground, 5km S Coldspring, forest, FIT, 22 May–16

Aug 1983, S. & J. Peck; Tyler Co.: 11 females, Kirby State Forest, dung traps, 12–17 Jun 1993,

S.A. Marshall; 4 females, Bouton Lake, dung/vac traps, 13–17 Jun 1993, S.A. Marshall.

VENEZUELA: Aragua: 1 female, Henri Pittier National Park, Rancho Grande, La Toma, 1150 m,

9 Apr 1994, L. Masner; 3 females, Maracay, Rancho Grande, 1200m, cloud forest, FIT, 1–10 Aug

1987, Bordon & Peck (MIZA); 2 males 4 females, San Cristobal, 20km NE, 4000 ft, dung trap, 18–

22 May 1974, S. Peck (MIZA); Bolivar: 3 males, km40 Sta. Elena Icabaru Road, 1000m, 4–6 Aug

1986, B.D. Gill; 1 male, km40 Sta. Elena Icabaru Road, 100m, 4–6 Aug 1986, B.D. Gill; 3 females,

125km S El Dorado, 1100m, 18 Jul–7 Aug 1986, B.D. Gill; 1 male 2 females, El Dorado, 135km S,

1400m, dung traps, 20 Jul–7 Aug 1986, B. Gill (CNCI); 1 male 1 female, 10km S El Dorado, 200m,

17 Jul–7 Aug 1986, B.D. Gill; 1 male, 33km S El Dorado, 220m, 2–7 Aug 1986, B.D. Gill; 1 female, km40 Sta. Elena Icabaru Rd., 220m, 4–6 Aug 1986, B.D. Gill (MIZA); 1 female, 22km S El

Dorado, lowland rainforest, FIT, 25 Jun–12 Jul 1987, S. & J. Peck.

Comments: Rudolfina exuberata, the only Rudolfina that regularly occurs at low elevations, is widespread throughout South and Central America as well as the southern United States.

Etymology: The specific name, which has been a manuscript name since 1982, is from the Latin for

"abundant”. Of the 1933 specimens examined for this study, 1649 were R. exuberata.

Rudolfina howdeni Paiero & Marshall n. sp.

Description: Length: 1.6–2.0 mm. Eye height 2.0× genal height. Head with 4–5 interfrontal setae, and row of 6–8 small setulae on inner margin of orbital plate. Gena with 2 setae and 6–8 setulae.

Acrostichal setulae in 4–6 rows. Costa with costagial seta extending to subcostal break. Second costal sector ~0.8× third costal sector. Length of M1 between crossveins dm-cu and r-m ~2.0× dm-

- 81 - cu; CuA1 stub vein ~7.0× M1 stub vein. Male mid femur with 10–15 setae in ovoid ventrobasal cluster. Male mid tibia with ventral comb of 6–7 robust, short setae on apical ½.

Male abdomen (Fig. 5.11): Posterior margin of sternite 5 with pair of large medially–projecting blunt teeth on each side of emargination; emargination extending ~1/4 length and ~1/8 length of sternite 5; emargination lined with sclerotized strip. Transverse part of sternite 6 weakly arcuate.

Epandrium with pair of long lateral setae, 3–4 smaller pairs of setae along lower margin of anal opening, and several other small setae scattered over the surface; subanal plate narrow but complete.

Surstylus (in lateral view) with small hirsute anterior lobe, and elongate clavate posterior lobe

(~4.0× length of anterior lobe); posterior lobe with 4–5 large setae and numerous smaller setae apically, and 2–3 bifurcating setae on inner surface. Cercus elongate, clavate, projecting ventrally.

Postgonite elongate, narrowly rounded apically. Distiphallus (Fig. 5.11F–H): dorsal sclerites not swollen along length and not extending beyond apex of acrophallus.

Female abdomen (Fig. 5.12): Posterior margin of tergite 7 entire or slightly emarginate medially.

Median part of tergite 8 elongate (length ~2–3× width) with anterior margin rounded. Epiproct diamond shaped. Cercus as long as epiproct. Sternite 7 entire. Spermatheca cup–like and short, with sclerotized duct ~3.5× length of spermatheca.

Type Material: Holotype (male, debu01086096, FMNH) and 8 paratypes (4 males, 4 females,

DEBU & FMNH): MEXICO: Oaxaca: Ixtlan de Juarez, 6.6mi. N, 8300 ft, oak woodland, human dung, 10–18 Aug 1973, A. Newton. Additional paratypes: MEXICO: Hidalgo: 1 male, Tenango de Doria, 7mi SW, 7000 ft, cloud oak forest, human dung, 2–6 Jul 1971, A. Newton; Jalisco: 24 males, Atenquique, 14 mi. W, 7900 ft, hardwood forest, dung, A. Newton (CASC, FMNH &

DEBU); 17 males, 8 females, Atenquique, 18 mi. W, 9300 ft, fir forest, dung, A. Newton (CNCI,

DEBU & FMNH); Mexico: 3 males, Ixtapan de la Sal, 1mi. E, km78, 6200 ft, Trop/Dec/Jun., human dung, 31 Aug–6 Sep 1971, A. Newton; 5 males, 1 female, Santa Marta, 5 mi E, km 8.5,

10100 ft, fir forest, human dung, 29 Aug–4 Sep 1971, A. Newton; Morelos: 2 females, Tres cumbres, 4mi W, km 6, 8900 ft, oak, human dung, 29 Aug–4 Sep 1971, A. Newton; 6 males, 4 females, Tres Cumbres, 4mi. W, km6 3/4, 9000 ft, oak, human dung, 29 Aug–4 Sep 1971, A.

Newton; Oaxaca: 1 male, Valle Nacional, 29.7 mi S, 6800 ft, cloud forest, dung, 11–17 Aug 1973, - 82 -

A. Newton; 1 male, Jct. Mex. 175–Yuvila Rd., 2.0 mi W, 9500 ft, oak–pine, dung, 8–19 Aug 1973,

A. Newton; 1 male, Jct. Mex. 175–Yuvila Rd., 4.1mi. W, 9300 ft, oak–fir–pine forest, 8–19 Aug

1973, A. Newton; 1 male, Jct. Mex. 175–Yuvila Rd., 5mi. E, 7600 ft, oak, pine, dung, 9–19 Aug

1973, A. Newton; Veracruz: 1 male, Ciudad Mendoza, 8.2mi. W on Mex. 150D, 6200 ft, oak woodland, carrion, 27 Jul–3 Aug 1973, A. Newton.

Comments: The male sternite 5, epandrium, surstylus and cercus are unique within Rudolfina.

Etymology: This species name, which has been a manuscript name since 1982, is a patronym for

Dr. Henry Howden. Henry was a great entomologist and friend who advised SAM during his M.Sc. at Carleton University; we regret that he did not live to see this beautiful species published.

Rudolfina megepandria Paiero & Marshall n. sp.

Description: Length: 1.9–2.2 mm. Eye height 2.5× genal height. Head with 3 interfrontal setae and row of 4–5 small setulae on inner margin of orbital plate. Gena with 2 strong setae and 7–8 smaller setulae. Acrostichal setulae in 4–6 rows. Costagial seta extending to subcostal break. Second costal sector 0.75–0.8× third costal sector. Length of M1 between crossveins dm-cu and r-m– 3.0× dm-cu;

CuA1 stub vein~10.0× M1 stub vein (M1 stub vein extremely reduced). Male mid femur with 14–15 strong setae in ventrobasal cluster. Male mid tibia with ventral comb of 4–5 robust, short setae on apical ½. Mid tibia without midventral seta (both sexes).

Male abdomen (Fig. 5.13): Sternite 5 posterior margin with 2 small nipple–like teeth on each side of small emargination; emargination ~1/15 width and ~1/9 length of sternite 5. Transverse part of sternite 6 arcuate. Epandrium swollen, broader than preceding abdominal segments; subanal plate broad, complete. Surstylus with posterior lobe elongate, length ~4× basal width, with a hirsute medial lateral lobe; apex truncate. Cercus elongate-conical, projecting posteriorly. Postgonite elongate, narrowly rounded distally and narrow basally. Distiphallus (Fig. 5.13E–G): dorsal sclerites not extending beyond apex of acrophallus.

Female abdomen (Fig. 5.4): Tergite 7 with posterior margin entire. Tergite 8 with middle part elongate, length ~2–3× width, with posterior margin emarginate. Epiproct broadly rounded

- 83 - anteriorly. Cercus ~3/4 length of epiproct; fused anterolaterally to epiproct. Sternite 7 with posterior margin broadly rounded. Spermathecae cup–shaped; paired spermathecae with stems ~1.5× width of spermatheca.

Type Material: Holotype (male, debu01086084, FMNH) and 2 paratypes (1 male, 1 female):

MEXICO: Jalisco: Atenquique, 18 mi. W, 9300 ft, fir forest, dung, A. Newton. Additional paratypes: MEXICO: Oaxaca: 1 females, Yuvila Rd., 9400 ft, mesic oak, carrion, 9–19 Aug

1973, A. Newton (FMNH).

Etymology: The specific name, which has been a manuscript name since 1982, refers to the unusually large epandrium.

Rudolfina newtoni Paiero & Marshall n. sp.

Description: Length: 1.6–2.0 mm. Eye height 1.5× genal height. Head with 4 interfrontal setae, and row of 6–7 small setulae on inner margin of orbital plate. Gena with 2 strong setae and 6–8 smaller setulae. Acrostichal setulae in 6–8 rows. Costagial seta extending slightly beyond humeral break.

Second costal sector ~1.2× as long as third. Length of M1 between crossveins dm-cu and r-m –2.0× dm-cu; CuA1 stub vein ~4.0× M1 stub vein. Male mid femur with 20–21 strong setae in ventrobasal cluster. Male mid tibia with ventral comb of 8 robust setae on apical ½.

Male abdomen (Fig. 5.14): Sternite 5 posterior margin with two distinct teeth on each side of large emargination; emargination forming a broad circular desclerotized area, ~1/3 width of sclerite and extending anteriorly ~3/4 length; margin of emargination with numerous setae on posterior 2/3.

Transverse part of sternite 6 arcuate. Epandrium with pair of long mediolateral setae present with 2–

3 smaller pairs lateral to large pair; subanal plate narrowly continuous below anal opening.

Surstylus (in lateral view) elongate, length ~4× basal width, tapering to narrowly rounded apex; bent posteriorly near basal ¼, with 4–5 setae on both ventral and dorsal surfaces. Cercus quadrate, weakly differentiated from each other and from epandrium, with pair of ventromedial setae.

Postgonite narrow, slightly sinuate with 4 small dorsal setulae on apical 1/2. Distiphallus (Fig.

5.14E–G): dorsal sclerites not swollen along length and not extending beyond apex of acrophallus.

- 84 -

Female abdomen (Fig. 5.15): Posterior margin of tergite 7 entire. Median part of tergite 8 short and kidney–shaped with posterior margin emarginate. Epiproct triangular, elongate, with mid–line desclerotized. Cercus half as long as epiproct, with flattened apical seta; narrowly fused anterolaterally to epiproct. Posterior margin of sternite 7 entire or slightly arcuate. Spermathecae inverted cup–shaped (with duct, appearing mushroom–shaped) and wrinkled; sclerotized portion of ducts ~2.0× length of spermatheca; common duct of paired spermathecae poorly sclerotized.

Type Material: Holotype (male, debu01086218): MEXICO: Oaxaca: Jct. Mex. 175–Yuvila Rd.,

4.1mi. W, 9300 ft, oak–fir–pine forest, 8–19 Aug 1973, A. Newton (FMNH). Paratypes:

MEXICO: Oaxaca: 11 males, 8 females, same as holotype; 1 male, 1 female, Ixtlan, 25 mi N,

17°33'0"N, 96°31'12"W, 9100 ft, oak–pine, human dung, 23–29 Jul 1971, A. Newton (DEBU &

FMNH); 1 female, Ixtlan, 28 mi N, 9500 ft, pine–heath, human dung, 23–29 Jul 1971, A. Newton; 8 males, 9 females, Jct. Mex. 175–Yuvila Rd., 2.0 mi W, 9500 ft, oak–pine, dung, 8–19 Aug 1973, A.

Newton; 2 females, Jct. Mex. 175–Yuvila Rd., 1.7 mi W, 2865 m, mesic oak, fish trap, 9–19 Aug

1973, A. Newton; 7 males, 6 females, Jct. Mex. 175–Yuvila Road, 1.4 mi W, 9300 ft, mesic oak forest, 9–19 Aug 1973, A. Newton; 3 males, 2 females, Jct. Mex. 175–Yuvila Road, 1.4 mi W, 9300 ft, dung, 9–19 Aug 1973, A. Newton

Etymology: The species name is a patronym for Dr. Alfred Newton, the collector of all known specimens of this species.

Rudolfina pauca Paiero & Marshall n. sp.

Description: Length: 1.4–1.9 mm. Eye height 1.3× genal height. Head with 4 interfrontal setae, and row of 8–10 small setulae on inner margin of orbital plate. Gena with 1 strong setae and 8 smaller setulae. Acrostichal setulae in 6–8 rows. Costagial seta extending to humeral break; second costal sector shorter than third (20:35); Length of M1 between crossveins dm-cu and r-m 1.2× dm-cu;

CuA1 stub vein ~2.0× M1 stub vein. Legs: Male mid femur with 4–5 weak setae in ventrobasal cluster. Male mid tibia with ventral comb of 11–13 robust setae on apical ½.

- 85 -

Male abdomen (Fig. 5.16): Sternites 3–4 evenly haired. Posterior margin of sternite 5 with lobes elongate, projecting posteriorly; emargination extending anteriorly ~1/4 length of sternite, ~1/4 width of sternite at posterior margin. Transverse part of sternite 6 weakly arcuate. Epandrium with 2 long ventrolateral setae and scattered setulae; subanal plate narrowly continuous below anal opening. Surstylus (in lateral view) trilobed, with anterior lobe rounded and bare, middle lobe small, rounded and with 4–6 distal setae apically, and posterior lobe posteriorly projecting and rounded with 1 large flattened seta and 2–3 regular setae apically. Cercus small and conical, projecting posterolaterally with 2 apical setae. Postgonite slightly capitate or –oar–shaped, with apex ventrally emarginate; 3–4 setulate present along dorsal surface. Distiphallus (Fig. 5.16D–F): dorsal sclerites present but not extending beyond apex of acrophallus.

Female Abdomen (Fig. 5.17): Posterior margin of tergite 7 medially emarginate. Median part of tergite 8 elongate, rectangular (length ~3× width) and poorly sclerotized; overlapping basal projection of epiproct. Epiproct tulip–shaped, with elongate narrow anterior extension. Cercus reduced, shorter than apical flattened seta; fused anterolaterally to epiproct. Spermathecae acorn– shaped, with a small smooth distal bulb, a wrinkled medial section and a smooth basal section; paired spermathecae with stems as long as spermatheca; common duct of pair short, ~1/6 length of stems.

Type Material: Holotype (male, debu01086244, FMNH) and 33 paratypes (13 males, 20 females; DEBU & FMNH): MEXICO: Mexico: Tenancingo, 1mi NE, 7100 ft, oak–pine–madrono, human dung, 31 Aug–6 Sep 1971, A. Newton. Additional paratypes: GUATEMALA: San

Marcos: 1 female, San Antonio Sacatepéquez, 14°58'N, 91°44'W, 8000 ft, 29 Sep 1986, M.J.

Sharkey. MEXICO: Hidalgo: 1 male, 1 female, Tulancingo, 4 mi W, 7600 ft, human dung, 1–6 Jul

1971, A. Newton; Mexico: 3 males, 1 female, Tenancingo, 5 mi SW, km55 1/3, 7200 ft, oak–pine, human dung, 31 Aug–6 Sep 1971, A. Newton; Morelos: 1 female, Tres Cumbres, 8 mi S, 7400 ft, oak/pine forest, dung, 29 Aug–4 Sep 1971, A. Newton.

Etymology: The species name is the Latin for "few”. Compared with the closely related R. exuberata, which occurs throughout the Neotropical region and has been collected in over a hundred

- 86 - different sites, R. pauca has been infrequently collected and remains known from only five sites, in

Mexico and Guatemala.

Rudolfina pilosa Paiero & Marshall n. sp.

Description: Length: 1.5–1.8 mm. Eye height 2.5× genal height. Head with 3 interfrontal setae and

5–6 small setae along inner margin of orbital plate. Gena with 2 strong setae and 7–9 smaller setulae. Acrostichal setulae in 4–6 rows. Costagial seta extending slightly beyond humeral break; second costal sector equal to third sector; Length of M1 between crossveins dm-cu and r-m– 2.0× dm-cu; CuA1 stub vein ~5.0× M1 stub vein. Legs: Male mid femur with 4–5 small setae present in ventrobasal cluster. Male mid tibia with ventral comb of 7–8 robust setae on apical ½, midventral seta absent (female unknown).

Male abdomen (Fig. 5.18): Sternite 3 and 4 with dense cluster of setae posteromedially. Posterior margin of sternite 5 with round teeth on each side of a medial emargination; emargination less than

1/6 width of sternite and extending ~1/2 length. Transverse part of sternite 6 weakly arcuate.

Epandrium with one large pair of setae ventrolaterally, one smaller pair of setae lateral to larger pair, and several setulae scattered over surface; subanal plate broadly continuous below anal opening. Surstylus boot–shaped, posteriorly bent near base, hirsute with 3–5 setae on posterior lobe

(heel) and numerous smaller setae on anterior lobe (toe). Cercus fused with epandrium, with distinct seta on ventral margin. Postgonite clavate, with several setulae on dorsal surface. Distiphallus (Fig.

5.18E–G): dorsal sclerite without distinct swellings.

Female: Unknown

Type Material: Holotype (male, debu01086240): MEXICO: Oaxaca: Jct. Mex. 175–Yuvila Rd.,

2.0 mi W, 9500 ft, oak–pine, dung, 8–19 Aug 1973, A. Newton (FMNH). Paratypes: MEXICO:

Oaxaca: 1 male, Ixtlan de Juarez, 6.6mi. N, 8300 ft, oak woodland, human dung, 10–18 Aug 1973,

A. Newton; 2 males, Jct. Mex. 175–Yuvila Rd., 4.1mi. W, 9300 ft, oak–fir–pine forest, 8–19 Aug

1973, A. Newton (DEBU, FMNH).

- 87 -

Etymology: The species epithet is from the Latin for hairy, referring to the hirsute sternites 4 and 5 of the male.

Rudolfina remiforma Paiero & Marshall n. sp.

Description: Length: 1.4–1.8 mm. Eye height 2.0× genal height. Head with 4 interfrontal setae and row of 7 small setulae along inner margin of orbital plate. Gena with 1 strong setae and 8 smaller setulae. Acrostichal setulae in 4–6 rows. Costagial seta extending to humeral break; second costal sector shorter than third (20:65); length of M1 between crossveins dm-cu and r-m 1.4× dm-cu; CuA1 stub vein ~2.5–3.0× M1 stub vein. Male mid femur with 6 weak setae in ventrobasal cluster. Male mid tibia with ventral comb of 4–5 robust setae on apical 1/3.

Male abdomen (Fig. 5.19): Posterior margin of sternite 5 with small tooth on each side of medial emargination; emargination ~1/3× width of sternite 5. Transverse part of sternite 6 weakly arcuate.

Epandrium regular, not swollen, with pair of long setae ventrolaterally and another 2–3 smaller pair dorsolateral to longer pair; subanal plate narrow, incomplete with narrow medial space. Surstylus with small anterior lobe and elongate posterior lobe; anterior lobe with 4–5 elongate setae apically; posterior lobe paddle–like and drawn out into rounded tip, with posterior margin weakly expanded near mid–length, anterior margin greatly expanded just before midlength, and 1 large setae near base. Cercus small and conical, projecting posteriorly; apical seta present. Postgonite simple, apex narrowly rounded. Dorsal sclerite without distinct swellings and not extending distally beyond acrophallus (Fig. 5.19 E–G).

Female abdomen (Fig. 5.20): Tergite 7 with posterior margin entire. Tergite 8 with middle part poorly sclerotized or fused with epiproct. Epiproct tulip–shaped, with length of anterior narrowed portion equal or subequal to posterior portion. Cercus small, as long or only slightly longer than flattened apical seta; fused anterolaterally to epiproct. Sternite 7 with posterior margin broadly rounded. Spermatheca lenticular, inner surface with conical projection that connects with duct; paired spermathecae each with duct ~2× length of spermatheca before common duct; single spermatheca duct similar in length..

- 88 -

Type Material: Holotype (male, debu01086286, FMNH) and 3 paratypes (3 females, DEBU,

FMNH): MEXICO: Chiapas: Bochil, 21mi. N, 5500 ft, pine, oak, Liquidambar, human dung, 18–

24 Aug 1971, A. Newton.

Etymology: The species name is from the Latin for “oar–shaped” and refers to the posterior lobe of the surstylus.

Rudolfina rozkosnyi (Roháček, 1975)

Synonyms: Rudolfia rozkosnyi (Roháček), Limosina rozkosnyi Roháček.

Distribution: Palaearctic: Austria, Czech Republic, Germany, Norway, Russia (North European

Territory, West Siberia), Slovakia, Sweden.

Description: See Roháček (1975).

Specimens examined: CZECH REPUBLIC: 2 males, 2 females, Jezinek Mountains, Velka–

Kotlina Valley, 900–1100m, 16 Aug 1986, S.A. Marshall; RUSSIA: Siberia: 1 female, Altai Reg.,

Teletskoya Lake, ~50km SE, 1500 m, wet area, 13–15 Jul 1991, S.A. Marshall.

Commments: This species is here newly recorded from Siberia on the basis of a single female taken in pan traps near the margin of Teletskoya Lake. Previous Russian records are from the

Northern European Territory.

Rudolfina tumida Paiero & Marshall n. sp.

Description: Length: 1.8–1.9 mm. Eye height 1.0× genal height. Head with 3 interfrontal setae, and

5 small setae along inner margin of orbital plate. Gena with 2 strong setae and 4–5 smaller setulae.

Acrostichal setulae in 4–6 rows. Costagial seta length unknown (broken on both sides of holotype); second costal sector shorter than third (35:40); Length of M1 between crossveins dm-cu and r-m

4.0× dm-cu; CuA1 stub vein~5.0× M1 stub vein. Male mid femur basally with 9 strong setae present in ovoid cluster. Male mid tibia with ventral comb of 12–13 robust setae on apical 2/3.

Male abdomen (Fig. 5.2, 5.3 and 5.21): Posterior margin of sternite 5 with pair of small teeth on each side of shallow emargination; emargination extending ¼ length and ¼ width of sternite 5.

- 89 -

Transverse part of sternite 6 weakly arcuate. Epandrium swollen, wider than preceding portion of abdomen (denuded on holotype); subanal plate incomplete. Surstylus with posterior lobe elongated, with apex and posterior margin heavily setose; broad rounded laminate lobe with distal triangular projection present anterobasally. Cercus ovoid. Postgonite elongate with apex acute. Dorsal sclerite of distiphallus with several small swellings along length.

Female: Unknown.

Type Material: Holotype (male, debu01086083, DEBU): U.S.A.: WY: Unita Co., Evanston, 8mi.

SE, 7100ft., sage–grass, riparian, carrion, 30 Jul–11 Aug 1979, S. & J. Peck.

Comments: Male R. tumida are highly distinctive for the uniquely swollen epandrium and large surstylus.

Etymology: The species epithet is the Latin for “swollen”, referring to the enlarged epandrium.

5.9 Described species in other genera previously treated as Rudolfina

Trachyopella opuntiae (Richards 1967)

Comments: Richards (1967) originally described this species as a Trachyopella (then a subgenus of

Leptocera), a placement that was followed by Roháček and Marshall (1986) in their review of

Holarctic Trachyopella. Pitkin (1989) transferred opuntiae to Rudolfina, in view of its modified female cerci. Examination of type specimens, including male and female terminalia, supports the treatment of this species as a Trachyopella.

Leptocera (Acuminiseta) prominens Duda 1925

Comments: Leptocera (Acuminiseta) prominens was provisionally treated as Rudolfina by Roháček et al. (2001), who noted that Acuminiseta does not occur in the New World. This species does not fit into Rudolfina as defined here, but belongs elsewhere in the Archiceroptera complex and will be treated in a later paper (Chapter 4).

- 90 -

5.10 Discussion

With the exception of the widespread lowland species R. exuberata, Rudolfina is a

Holarctic, and largely Nearctic genus with highest diversity in the highlands of Mexico. The few records from mountains in Guatemala are from an area recognized as a transitional zone between

Nearctic and Neotropical regions. This is consistent with patterns observed in some bats (Ortega and

Arita 1998), copepods (Aglaodiaptomus and Skistodiaptomus; see Suarez-Morales et al. 2005), and passalids (Gutierrez-Velazquez et al. 2013). There are no records of Rudolfina species (other than R. exuberata) from lowland Guatemala, southern Guatemala or any other Neotropical locality. We have studied tens of thousands of specimens collected from throughout Costa Rica over the past thirty years without finding any specimens other than R. exuberata that fit Rudolfina as currently defined; this reinforces our conclusion that the distribution of Rudolfina is effectively bounded southward by the limits of the Nearctic region. The origins of Rudolfina are likely to be in western

North America, where R. cavernicola, the basal most species, occurs, with the Nearctic and

Palaearctic faunas diverging relatively early in Rudolfina’s evolutionary history.

The number of new species in the University of Guelph insect collection and the apparent novelty of the limited material from other collections, suggest that many further new species await discovery. Southern Mexico is especially poorly represented in the current material. We have only a single specimen (R. cavernicola) from the Sierra Madre Occidental range, and we have seen no

Rudolfina from the Sierra Madre Oriental. Further collecting efforts in Mexico would greatly improve our understanding of Rudolfina.

Acknowledgements:

This work was made possible largely through the generosity of Dr. Alfred Newton

(FMNH) who kindly allowed Marshall to take the Sphaeroceridae from his Mexican beetle trap residues during the 1980s. Of the 284 non-R. exuberata Rudolfina specimens studied for this revision, 219 were from these residues. Despite ongoing efforts to obtain further material from

- 91 - throughout the range of Rudolfina and related genera, the material collected by Dr. Newton during the early 1970s remains by far the most important collection of Mexican Rudolfina.

Supplemental drawings used to support the photographs were prepared by the second author, and by illustrator Rebecca Langstaff, for an unpublished manuscript.

Funding for this study was provided by NSERC Discovery grants to SAM, and NSERC postgraduate scholarships to SMP.

5.11 References

Cumming, J.M., & Wood, D.M. (2010). Adult morphology and terminology. In B.V. Brown (ed.)

Manual of Central American Diptera Volume 1. NRC Research Press, pp.9–63.

Goloboff, P.A., Farris, J. & Nixon, K. (2008) TNT: a free program for phylogenetic analysis.

Cladistics, 24, 774–786.

Gutierrez-Velazquez, A., Rojas-Soto, O., Reyes-Castillo, P. & Halffter, G. (2013) The classic theory

of Mexican Transition Zone revisited: the distributional congruence patterns of Passalidae

(Coleoptera). Invertebrate Systematics, 27: 282–293.

Maddison, W.P. & Maddison, D.R. (2017) Mesquite: a modular system for evolutionary analysis.

Version 3.2. Available from: http:mesquiteproject.org (Accessed January 2017).

Marshall, S.A. (1991) Rudolfina digitata sp. nov., a new Nearctic sphaerocerid with a disjunct

alpine-arctic distribution. Canadian Entomologist, 123, 621–626.

Marshall, S.A. & Buck, M. (2010) Sphaeroceridae (Small dung flies). In: Brown, B.V., Borkent, A.,

Cumming, J.M., Wood, D.M., Woodley, N.E., & Zumbado, M.A. (Eds), Manual of Central

American Diptera. NRC Research Press, Ottawa, Ontario, pp. 1165–1187.

Marshall, S.A. & Fitzgerald, S. (1996) Rudolfina cavernicola, a new species of cave-associated

Sphaeroceridae (Diptera) from Colorado and Arizona. Proceedings of the Entomological

Society of Washington, 99, 641–644.

Nixon, K.C. (2002) WinClada ver. 1.00.08 Published by the author, Ithaca, NY, USA.

Ortega, J. & Arita, H.T. (1998) Neotropical-Nearctic limits in Middle America as determined by

distribution of Bats. Journal of Mammology, 79(3), 72–783. - 92 -

Papp, L. (1977) A contribution to the knowledge of species of the subfamily Ceropterinae (Diptera:

Sphaeroceridae). Acta Zoologica Academiae Scientiarum Hungaricae, 23(3–4), 371–385.

Pitkin, B.R. (1989) A review of the Sphaeroceridae (Diptera) described by O. W. Richards.

Occasional Papers on Systematic Entomology, London, 6: 1–44.

Richards, O.W. (1967) On a collection of Sphaeroceridae (Diptera) from the Galapagos Islands.

Annals and Magazine of Natural History (Series 13), 9, 531–535.

Roháček, J. (1982) A monograph and reclassification of the previous genus Limosina Macquart

(Diptera, Sphaeroceridae) of Europe, Part I, Beitrage zur Entomologie, 32, 195-282.

Roháček, J. (1983) A monograph and reclassification of the previous genus Limosina Macquart

(Diptera, Sphaeroceridae) of Europe, Part II, Beitrage zur Entomologie, 33, 3–195.

Roháček, J. (1985) A monograph and reclassification of the previous genus Limosina Macquart

(Diptera, Sphaeroceridae) of Europe, Part IV. Beitrage zur Entomologie, 35, 101-179.

Roháček, J. (1987) Replacement name for Rudolfia Roháček, 1982 (Diptera, Sphaeroceridae), with

first record of R. rozkosnyi from northern Europe. Acta Entomologica Bohemoslovaca, 84,

474–476.

Roháček, J. & Marshall, S.A. (1986) The genus Trachyopella Duda (Diptera, Sphaeroceridae) of the

Holarctic Region. Monografie III (1985), Torino: Museo Regionale di Scienze Naturali,

109 pp.

Roháček, J. & Marshall, S.A. (2017) Volumosina, a new Nearctic genus for the rare old-growth

forest fly Herniosina voluminosa Marshall (Diptera: Sphaeroceridae). Canadian

Entomologist, 149(4), 444–460.

Roháček, J., Marshall, S.A., Norrbom, A.L., Buck, M., Quiros, D.I. & Smith, I. (2001) World

catalog of Sphaeroceridae (Diptera). Slezské zemské muzeum, Opava, 414 pp. (also online

at www.uoguelph.ca/debu/catalog.htm.

Shorthouse, D.P. 2010. SimpleMappr, an online tool to produce publication-quality point maps.

[Retrieved from http://www.simplemappr.net. Accessed January 29, 2015].

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Smith, I.P., & Marshall, S.A. (2004) A review of the New World genus Pterogramma Spuler and a

revision of the Pterogramma sublugubrinum group (Diptera: Sphaeroceridae:

Limosininae). Contributions in Science, 499, 1–163.

Su, L., Xu, J. & Cong, G. (2017) A new species of the Genus Rudolfina Roháček, 1987

(Diptera, Sphaeroceridae) from North-east China, with a key to the known

Holarctic species of Rudolfina. Oriental Insects, 51(4), 391–396.

Suarez-Morales, E., Reind, J.W. & Elias-Guitierrez, M. (2005) Diversity and distributional patterns

of neotropical freshwater copepods (Calanoida: Diaptomidae). International Review of

Hydrobiology, 90(1), 71–83.

5.12 Table List

Table 1. Character states used in the phylogenetic analysis of Rudolfina.

5.13 Figure List

Figure 1. Rudolfina head and wing morphology: A) R. howdeni head (debu01086104); and B) R. exuberata wing (debu00276674).

Figure 2. Rudolfina tumida. A) male capsule, posterior view; B) male capsule, lateral view.

Illustration and photograph from debu01086083.

Figure 3. Male morphology. A) sternite 5, ventral view (Rudolfina tumida, debu01086083); B) hypandrium (R. exuberata, debu00242299); C) postgonite, lateral view (R. cavernicola, debu01086077); D) phallus (including the basiphallus and distiphallus), postgonite and phallapodeme, lateral view (R. tumida, debu01086083).

Figure 4. Rudolfina megepandria , female terminalia: A) terminal abdominal segments, dorsal view;

B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086086). - 94 -

Figure 5. Strict Consensus Tree for the six recovered trees obtained from Traditional Search (TNT).

Length = 82; Ci = 50, Ri = 52.

Figure 6. Majority Rules Consensus Tree from the six optimized trees retained from Traditional

Search (TNT). Length=72, Ci=54, Ri=60.

Figure 7. Phylogeny of Rudolfina species. Character and character states refer to those given in table

1. Tree selected from six equal length trees. Length=72, Ci=56. Ri=63.

Figure 8. Rudolfina bucki, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus, dorsal view; E) phallus, postgonite and phallapodeme, dorsolateral view; F) same, lateral view. (A–F from debu01086239).

Figure 9. Rudolfina exuberata, male terminalia: A) epandrium, surstylus and cercus, caudal view;

B) same, lateral view ventral; C) sternite 5, ventral view; D) postgonite, close up lateral; E) phallus, dorsal view; F) same, dorsolateral view; G) same, lateral view. (A–G from debu00242299).

Figure 10. Rudolfina exuberata, female terminalia: A) terminal abdominal segments, dorsal view;

B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. A–C) from debu00242299; D) from debu00242286).

Figure 11. Rudolfina howdeni, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus and postgonites; dorsal view; E) phallus and postgonite, lateral view; F) same, dorsolateral view. (Photos A–F from debu01086163).

Figure 12. Rudolfina howdeni, female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu1086100).

Figure 13. Rudolfina megepandria, male terminalia: A) epandrium, surstylus and cercus, lateral view; B) same, caudal view; C) sternites 4-7, ventral view; D) phallus and postgonite, dorsal view;

E) phallus, postgonite and phallapodeme, dorsolateral view; F) same, dorsolateral view. (A–G from debu01086085).

Figure 14. Rudolfina newtoni, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternites 4 and 5, ventral view; D) phallus, dorsal view; E) phallus,

- 95 - dorsolateral view; F) phallus and postgonite, lateral view. (A–E from debu01086234, F from debu01086234).

Figure 15. Rudolfina newtoni , female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086226).

Figure 16. Rudolfina pauca, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus, postgonites and phallapodeme, dorsal view; E) phallus, postgonites, hypandrium and phallapodeme, dorsolateral view; F) same, lateral view. (A–F from debu1086247).

Figure 17. Rudolfina pauca, female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086258).

Figure 18. Rudolfina pilosa, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus and ejaculatory apodeme, dorsal view; E) same, dorsolateral view; F) same, lateral view; G) postgonite, lateral view. (A–G from debu01086241).

Figure 19. Rudolfina remiforma, male terminalia: A) epandrium, surstylus and cercus, caudal view;

B) same, lateral view; C) sternite 5, ventral view; D) phallus and postgonites, dorsal view; E) phallus, postgonites and phallapodeme, dorsolateral view; F) same, lateral view. (A–F from debu01086286).

Figure 20. Rudolfina remiforma , female terminalia: A) terminal abdominal segments, dorsal view;

B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086288).

Figure 21. Rudolfina tumida: A) sternite 5, close up ventral; B) phallus and postgonites, dorsal view;

C) same, dorsolateral view. (A–C from debu01086083).

Figure 22. Distribution of New World Rudolfina species: A) R. cavernicola, R. digitata, and R. tumida; B) R. bucki, R. megepandria, and R. howdeni; C) R. pauca, R. pilosa, R. newtoni, and R. remiforma; and D) R. exuberata. - 96 -

- 97 -

Table 5.1. Character states used in the phylogenetic analysis of Rudolfina.

Character 0 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 outgroup 2 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 0 0 0 0 0 0 0 0

R. bucki 3 0 1 1 0 0 1 0 1 0 1 1 1 1 0 1 1 1 1 1 0 0 0 0 0 1 ? ? ? ? ? ? ? ? ? ? ?

R. cavernicola 2 0 1 1 0 0 1 1 1 0 0 1 0 0 0 0 0 1 0 0 1 1 0 0 1 1 0 1 1 1 0 0 1 0 1 0 1

R. digitata 2 0 1 1 0 0 1 0 1 0 0 1 0 0 0 1 0 1 0 1 1 2 0 0 1 1 1 1 1 1 0 0 1 0 1 1 1

R. exuberata 2 1 1 0 1 1 1 0 2 0 0 0 0 1 1 1 1 0 1 1 0 0 0 1 0 1 1 1 1 1 0 1 1 1 1 1 1

R. howdeni 2 0 1 2 0 0 1 0 1 1 0 1 0 0 1 1 1 1 0 1 0 0 0 0 0 1 1 1 1 1 1 0 1 0 1 0 1

R. megepandria 3 0 1 2 0 0 1 0 1 0 1 1 1 0 1 1 0 1 1 1 0 0 0 0 0 1 1 1 1 1 0 0 1 0 1 1 1

R. newtoni 1 1 1 1 0 0 1 0 2 1 0 0 0 1 0 0 0 1 0 1 0 0 0 1 0 1 0 1 1 1 1 1 1 0 1 1 1

R. pauca 1 1 1 0 0 0 1 0 2 0 0 1 0 1 1 1 1 0 1 1 0 0 1 1 0 1 1 1 1 1 0 1 1 1 1 1 1

R. pilosa 3 0 1 1 0 0 1 0 2 1 0 1 0 1 0 0 1 1 0 1 0 0 1 0 0 1 ? ? ? ? ? ? ? ? ? ? ?

R. remiforma 2 0 1 0 1 1 1 0 1 0 0 0 0 1 1 1 1 1 0 1 0 0 0 1 0 1 1 1 1 1 0 1 1 1 1 1 1

R. rozkosnyi 2 0 1 1 0 0 1 0 1 0 0 1 0 0 0 1 0 1 0 1 1 2 0 0 1 1 0 1 1 1 0 0 1 0 1 1 1

R. tumida 0 0 1 1 0 0 1 0 1 0 1 0 0 0 1 1 1 1 0 1 1 2 1 0 1 1 ? ? ? ? ? ? ? ? ? ? ?

5.14 Figures

Figure 5.1. Rudolfina head and wing morphology: A) R. howdeni head (debu01086104); and B) R.

exuberata wing (debu00276674).

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Figure 5.2. Rudolfina tumida. A) male epandrium, cercus and surstylus, posterior view; B) same, lateral view. Illustration and photograph from debu01086083.

- 99 -

Figure 5.3. Male morphology. A) sternite 5 (Rudolfina tumida, debu01086083); B) hypandrium (R. exuberata, debu00242299); C) postgonite, lateral view (R. cavernicola, debu01086077); D) phallus

(including the basiphallus and distiphallus), postgonite and phallapodeme, lateral view (R. tumida, debu01086083).

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Figure 5.4. Rudolfina megepandria , female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086086).

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Figure 5.5. Strict consensus tree for the six trees obtained from traditional search (TNT).

Figure 5.6. Majority Rules consensus tree from the six optimized trees retained from Traditional

Search (TNT).

- 102 -

Figure 5.7. Phylogeny of Rudolfina species. Character and character states refer to table 1. One of six equal length trees. Length=72, Ci=56. Ri=63.

- 103 -

Figure 5.8. Rudolfina bucki, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus, dorsal view; E) phallus, postgonite and phallapodeme, dorsolateral view; F) same, lateral view. (A–F from debu01086239).

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Figure 5.9. Rudolfina exuberata, male terminalia: A) epandrium, surstylus and cercus, caudal view;

B) same, lateral; C) sternite 5; D) postgonite, lateral; E) phallus, dorsal view; F) same, dorsolateral view; G) same, lateral view. (A–G from debu00242299).

- 105 -

Figure 5.10. Rudolfina exuberata, female terminalia: A) terminal abdominal segments, dorsal view;

B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. A–C) from debu00242299; D) from debu00242286).

- 106 -

Figure 5.11. Rudolfina howdeni, male terminalia: A) epandrium, surstylus and cercus, caudal view;

B) same, lateral view; C) sternite 5, ventral view; D) phallus and postgonites; dorsal view; E) phallus and postgonite, lateral view; F) same, dorsolateral view. (Photos A–F from debu01086163).

- 107 -

Figure 5.12. Rudolfina howdeni, female terminalia: A) terminal abdominal segments, dorsal view;

B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu1086100).

- 108 -

Figure 5.13. Rudolfina megepandria, male terminalia: A) epandrium, surstylus and cercus, lateral view; B) same, caudal view; C) sternites 4-7, ventral view; D) phallus and postgonite, dorsal view;

E) phallus, postgonite and phallapodeme, dorsolateral view; F) same, dorsolateral view. (A–G from debu01086085).

- 109 -

Figure 5.14. Rudolfina newtoni, male terminalia: A) epandrium, surstylus and cercus, caudal view;

B) same, lateral view; C) sternites 4 and 5, ventral view; D) phallus, dorsal view; E) phallus, dorsolateral view; F) phallus and postgonite, lateral view. (A–E from debu01086234, F from debu01086234).

- 110 -

Figure 5.15. Rudolfina newtoni , female terminalia: A) terminal abdominal segments, dorsal view;

B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086226).

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Figure 5.16. Rudolfina pauca, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus, postgonites and phallapodeme, dorsal view; E) phallus, postgonites, hypandrium and phallapodeme, dorsolateral view; F) same, lateral view. (A–F from debu1086247).

- 112 -

Figure 5.17. Rudolfina pauca, female terminalia: A) terminal abdominal segments, dorsal view; B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086258).

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Figure 5.18. Rudolfina pilosa, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus and ejaculatory apodeme, dorsal view; E) same, dorsolateral view; F) same, lateral view; G) postgonite, lateral view. (A–G from debu01086241).

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Figure 5.19. Rudolfina remiforma, male terminalia: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5, ventral view; D) phallus and postgonites, dorsal view; E) phallus, postgonites and phallapodeme, dorsolateral view; F) same, lateral view. (A–F from debu01086286).

- 115 -

Figure 5.20. Rudolfina remiforma, female terminalia: A) terminal abdominal segments, dorsal view;

B) same as previous, lateral view; C) same as previous, ventral view; D) spermathecae. (A–D from debu01086288).

- 116 -

Figure 5.21. Rudolfina tumida: A) sternite 5, close up ventral; B) phallus and postgonites, dorsal view; C) same, dorsolateral view. (A–C from debu01086083).

- 117 -

Figure 5.22. Distribution of New World Rudolfina species: A) R. cavernicola, R. digitata, and R. tumida; B) R. bucki, R. megepandria, and R. howdeni; C) R. pauca, R. pilosa, R. newtoni, and R. remiforma; and D) R. exuberata.

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CHAPTER 6 – A REVISION OF ARCHICEROPTERA PAPP 1977

6.1 Abstract

The genus Archiceroptera Papp (Diptera: Sphaeroceridae: Limosininae), an almost entirely

Neotropical genus ranging from southwestern USA to Argentina, is revised to include 29 species, including the following 27 new species: A. adamas, A. addenda, A. barberi, A. basilia, A. bilobata,

A. bisetosus, A. braziliensis, A. brevivilla, A. browni, A. caliga, A. calligraphia, A. cobolorum, A. crenulata, A. curvivilla, A. dolabra, A. llama, A. maniba, A. masoni, A. megacercus, A. megavilla,

A. mexicorona, A. mitarakai, A. paracercus, A. pussula, A. ternum, A. triclavus, and A. unicinata. A species phylogeny based on morphological characters is provided and all species are keyed.

6.2 Introduction

The genus Archiceroptera Papp 1977 was originally described to include two South

American species: A. mahukani Papp and A. venezolana (Richards). Based on a study of over 3600 specimens, we here redefine Archiceroptera to include 29 species, 27 of which are newly described.

Archiceroptera belongs to a group of genera characterized by the presence of an epandrial process that extends from the right side along the posterior margin of sternite 8 towards the hypandrium

(Marshall and Cui 2005). The strongest synapomorphies for Archiceroptera within this clade are: vein M1 reaching the wing margin as a tracable fold, section of M1 between r-m and dm-cu > 3.0× length of dm-cu, and female cercus elongate with a strong apical seta. The female sternite 8 is medially divided (except in the putatively plesiomorphic A. addenda species group) into a pair of elongate glabrous lateral sclerites. Within Archiceroptera, four species groups are here recognized

(Archiceroptera mahukani group, the A. addenda group, the A. ternum group and the A. brevivilla subgroup).

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6.2.1 Biology

Most Archiceroptera species occur at low elevations (< 500m) but collections range from

0–2000m. Most of the specimens used in this revision were collected using dung and carrion traps.

Larval habitats remain unknown.

6.2.2 Related genera

Archiceroptera, along with Rudolfina and several unplaced species groups, were originally treated in the Archiceroptera genus complex (Sphaeroceridae: Limosininae) sensu Marshall and Buck

(2010), and these clades all belong to a larger clade including Aptilotella Duda, Bitheca Marshall,

Bromeloecia Spuler, Pterogramma Spuler, and Robustagramma Marshall & Cui (Marshall and Cui

2005). This larger clade is loosely referred to as the epandrial process group (EPG) because it is characterized by the presence of a finger-like process extending from the lower right corner of the epandrium along the posterior margin of sternite 8. A morphological phylogeny of the EPG

(Chapter 3) found the Archiceroptera genus complex was rendered paraphyletic by other genera.

In Marshall and Buck's (2010) key to the Neotropical genera of Sphaeroceridae,

Archiceroptera will key out to the Archiceroptera Papp genus complex and run to couplets 32

(Archiceroptera mahukani group), 72 (A. brevivilla group), 76 or 78 (A. ternum and A. addenda groups). Archiceroptera can be separated from other members of the Archiceroptera genus complex by the characters given in the diagnosis.

6.3 Materials and Methods

All specimens examined were dried and point-mounted. Abdomens of some specimens were removed and cleared by immersion into hot 10% potassium hydroxide for 6-10 minutes before being neutralized with 10% acetic acid for 10 minutes, rinsed in deionized water, and then placed into glycerin for examination. Dissected genitalia were stored in glycerine in microvials pinned below the specimen.

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Species Descriptions.

All label data are presented in a consistent manner, not verbatim from the labels; in a few cases, obvious spelling errors were corrected. Short-forms or abbreviations used on the labels are, where possible, given in full. Geographical coordinates are given only if present on the original label.

Specimens were given unique identifiers and their collection data was captured within the BIOTA database (BIOTA 3.0, Colwell 2012); this data will ultimately be hosted on Canadensys

(www.canadensys.net) but is not repeated in the text except for holotypes or as image reference.

Collection data for paratypes and other specimens examined were organized alphabetically by country, state/province, and locality name. Species distribution maps (Figs. 6.62–6.64) were generated using SimpleMappr (Shorthouse 2010).

Terminology

Terminology for external morphology follows Cumming and Wood (2010) with some modification.

Mid tibial dorsal chaetotaxy follows Buck and Marshall (2009). Male and female genitalia follow

Smith and Marshall (2004), with modifications from Cumming and Wood (2010), as labelled in

Figs. 6.5–6.7. The CuA1 and M1 stub veins are short portions of these veins that project distally beyond cell dm. The subanal plate refers to the fusion of the epandrium below the anal opening.

Female tergite 8 is treated here as tripartite, with the medial part articulating posteriorly with the epiproct (this medial sclerite is present but poorly developed in the A. addenda species group). The female sternite 8 is entire (plesiomorphic species) or medially desclerotized with two lateral triangular sclerites; a transverse posterior sclerite is also present in the A. addenda and A. mahukani species groups. Body length was measured from the anterior portion of the frons to the tip of the abdomen.

Depositories of Material Examined

Acronyms of Depositories: DEBU (School of Environmental Sciences, University of Guelph,

Guelph, Ontario, Canada); CNCI (Canadian National Collection, Ottawa, Ontario, Canada), FMNH - 121 -

(Field Museum of Natural History, Chicago, Illinois); INBC (Instituto Nacional de Biodiversidad,

Santo Domingo de Heredia, Costa Rica); MHNM (Museum National d’Histoire Naturelle, Paris,

France); MIZA (Museo del Instituto de Zoología Agrícola Francisco Fernández Yépez; Universidad

Central de Venezuela, Maracay, Venezuela); MUSM (Museo de Historia Natural, Universidad

Nacional Mayor de San Marcos, Lima, Peru); MZSP (Museu de Zoologia, Universidade de São

Paulo, São Paulo, São Paulo, Brazil); NHMW (Vienna Museum of Natural History, Vienna,

Austria), QCAZ (Departamento de Biología, Pontífica Universidad Católica del Ecuador, Quito,

Ecuador); ROME (Royal Ontario Museum, Toronto, Ontario); UASC (Museo de Historia Natural

Noel Kempff Mercado, Santa Cruz de la Sierra, Bolivia), UNAM (Universidad Nacional Autónoma de México, Mexico City, Mexico); USNM (United States National Museum of Natural History,

Smithsonian Institute, Washington, D.C., U.S.A.); UVGC (Colleción de Artrópodos, Universidad de

Valle de Guatemala, Guatemala City, Guatemala). Material is deposited in DEBU unless otherwise noted.

Illustrations

A Canon PowerShot S5IS with an ocular mount mounted on a Leitz Laborlux 11 Compound

Microscope was used to capture images of the genitalia in glycerin. Image layers where aligned and combined using Zerene Stacker version 1.04 (Zerene Systems LLC, Richland, WA, U.S.A.) with the

DMax algorithm. Species plates are organized alphabetically within species group.

6.4 Archiceroptera Papp 1977

Type species: Archiceroptera mahukani Papp 1977

6.4.1 Diagnosis

Archiceroptera is diagnosed by the following characters: interfrontal setae in 3–6 pairs, two or more inclinate orbital setulae in a single row; R4+5 straight or weakly curved just before meeting costa; M1

- 122 - traceable to wing margin; male cercus free and articulating, either triangular or with distinct distal medial process; female cercus glabrous and widened basally (usually appearing droplet-shaped) with a strong flattened apical seta; and female sternite 8 reduced to a pair of lateral triangular sclerites (except in the A. addenda species group). The mid tibia in all Archiceroptera (as with most genera in the epandrial process group) has three strong anterodorsal setae (one near basal ¼, one near mid length and one near distal ¼) and one distal posterodorsal seta, although many species have additional seta both anterodorsally and posterodorsally; ventrally all species have a distinct row of robust dark setae apically and most males have a long midventral seta. The modified female cercus and sternite 8 are considered defining characters.

6.4.2 Redescription

General: Colour variable from yellow brown to dark brown. 1.2–3.6 mm.

Head: Ocelli usually well developed (reduced in A. venezolana and absent in A. browni) with pair of

3–15 setulae on ocellar triangle between ocellar setae. Interfrontal setae in 3–6 equally long pairs

(anterior pair usually slightly shorter). Inclinate orbital setulae in usually in 3–6 pairs (except A. braziliensis which has only 1–2 pairs). Two strong orbital setae with 5–17 additional exclinate setulae along orbital plate. Outer vertical seta strong and exclinate, inner vertical seta strong and inclinate. Occipital and paravertical setae weak and inclinate. Postvertical setae inclinate or cruciate.

Gena with vibrissa, 1–2 strong subvibrissal setae, and 7–17 additional setulae.

Thoracic Chaetotaxy: Postpronotum with 2 strong setae; additional setulae sometimes present.

Notopleuron with 2 setae. Supra-alar seta in 1 presutural seta and two 2 postsutural setae.

Intrapostalar seta weak, ½ length of prescutellar dorsocentral. Dorsocentral setae in 1–6 pairs

(usually 1 prescutellar pair, but additional setae present in A. venezolana, A. browni, and A. mahukani). Acrostichal setulae in 4–10 rows; prescutellar acrostichal seta present. Katepisternum with anterior seta weakly developed or indistinct from nearby setulae; posterior seta strong, well developed. Scutellum with basal and apical pairs; setulae present near basal scutellar seta

(Archiceroptera s.str. and A. bilobata) or absent.

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Wing: Costa terminating at apex of R4+5 or shortly after, with 1–2 strong costagial setae (1 usually slightly stronger than the other in Archiceroptera s. str. and ternum species group; 1 distinctively stronger in addenda group); C2 > 1.2× C3. R4+5 straight or weakly curved. M1 traceable to wing margin. Distance between r-m and dm-cu > 3.0× dm-cu. Cell cup absent. Alula narrow with straight hind margin.

Legs: Fore femur with series of 4–7 posterodorsal setae and series of 4–8 posteroventral setae

(posteroventral series usually continue basally but setae are weaker and less distinct). Mid femur anteriorly with basal series of 8–12 robust setae extending to apical ¼ and distal series of 3–5 setae on apical ¼; in male, basal cluster of ventral setae usually present (variable between species). Mid tibia chaetotaxy variable: basal-most seta anterodorsal or dorsal (derived), with 1–3 anterodorsal and 1–3 posterodorsal setae present at midlength, and 2–5 distal setae (distal anterodorsal and distal dorsal setae always present; predistal anterodorsal, predistal dorsal, and distal posterodorsal present in some species); male usually with ventral comb of 6–17 setae on apical 2/3 or less (absent in some species); midventral seta present in known females and some males. Hind femur with apical series of 3–6 strong setae in a few species (A. venezolana, A. mahukani). Hind tibia often without dorsal setation but preapical seta and other dorsal setae present in some species (e.g., A. venezolana, A. megacerca, A. bilobata); ventrally with short apical spur.

Male Abdomen: Sternite 4 with posterior margin generally entire, but modified in some species.

Sternite 5 variable between species. Transverse portion of sternite 6 straight or arched. Synsternite

6+7 with elongate medially projecting process present near midlength (modified in some species and often with additional sclerite adjacent medially to apex of process). Ring sclerite present, associated with right portion of synsternite 6+7. Epandrium with distinct, acute ventral process

(except A. browni which has a relatively quadrate process) on right side, extending along posterior margin of sternite 8. Surstylus variable between species, separate beneath anal opening (except A. crenulata). Cerci not fused with epandrium, variable between species. Pregonite small, triangular.

Postgonite basally rectangular, with apical 1/2 relatively acute. Hypandrium forming a broad, largely hyaline triangular plate, with dark, well-sclerotized inverse “Y” medially, and with hypandrial arms extending from tips of Y to posterolateral corners (e.g., Fig. 6.40). Ejaculatory - 124 - apodeme short and finger-like (often damaged or lost in dissection). Basiphallus short and tubular, without epiphallus. Distiphallus membranous in most species (heavily sclerotized in A. addenda group) with a basal sclerite, a reduced “U-shaped” sclerite, and 3 distal pair of sclerites [one lateral flanking sclerite, starting dorsobasally before branching into pair of lateral arms, a ventral sclerite sometimes present (likely homologous to the ventral whip-like sclerites in Smith and Marshall

(2004), and a dorsal sclerite (likely homologous to Smith and Marshall’s (2004) second dorsal sclerite, with the other dorsal sclerites either absent or a part of the lateral sclerite)]; acrophallus often modified dorsolaterally, near midlength of distiphallus, and/or ventrally into flaps or processes.

Female Abdomen: Tergite 7 simple (plesiomorphic) or projecting posteromedially to fuse with anterolateral corner of tergite 8. Tergite 8 tripartite and reduced to lateral triangular sclerites and medial sclerite (absent to weakly developed in A. addenda group). Epiproct longitudinally weakened (plesiomorphic) or divided (apomorphic); mostly glabrous, with a few scattered setulae

(except A. addenda group). Cercus largely glabrous, basally wider (longitudinally divided in A. addenda group), with posterolateral extension curving inward and ending with large flattened apical seta. Sternite 7 posteriorly usually simple but modified in some species. Sternite 8 present as pair of lateral triangular sclerites (except in A.mahukani) that medially articulate in derived species.

Hypoproct medially divided; lateral sclerites triangular to rectangular. Spermatheca (2+1) variable but generally barrel-shaped or ovoid (sometimes flattened) with a small rounded protuberance at duct junction; paired spermathecae with short or obsolete stems before fusing into common duct.

6.5 Phylogeny

6.5.1 Morphological

A character matrix (Table 1) of 58 morphological characters organized by body region and sex was generated using Mesquite (version 3.10; Maddison and Maddison 2011), and exported for analysis in TNT (Goloboff et al. 2008) using traditional Search, with 10 random seeds and 5000 replications using the tree bisection reconnection (TBR) swapping algorithm.. Trees were optimized in

WINCLADA (Nixon 1999–2002). Character states were polarized using Thoracochaeta as an - 125 - outgroup to the EPG and Pectinosina as an outgroup within the EPG. The multistate characters 1, and 8–12 are treated as ordered (*).

6.5.2 Morphological characters and character states used in the phylogenetic analysis of

Archiceroptera

Head

1. *Ocellar development: (0) well developed; (1) reduced (size approximate to diameter of

ocellar seta base); (2) absent.

Thorax

2. Dorsocentral setae: (0) 3 or more; (1) 2; (2) 1.

3. Katepisternum – anterior seta development: (0) well developed; (1) present, but weak; (2)

absent.

4. Scutellum – extra scutellar setulae: (0) none present; (1) 1 or more present.

Wing

5. Costagial seta development: (0) 2 equal or subequal costagial setae present; (1) one

costagial seta greatly enlarged.

6. Costagial seta – maximum length: (0) longest seta not reaching humeral crossvein; (1)

longest seta extending between humeral crossvein and subcosta; (2) extending to or beyond

subcosta

Legs

7. Mid tibia - Basal seta placement: (0) anterodorsal, (1) dorsal

8. *Mid tibia – Anterodorsal cluster at midlength: (0) 2 or more seta present, (1) 1 seta

present; (2) none present.

9. *Mid tibia – Posterodorsal cluster at midlength: (0) 2 or more seta present; (1) 1 seta

present; (2) no setae present.

10. *Mid tibia – Predistal anterodorsal seta development: (0) strongly developed; (1) reduced;

(2) absent.

- 126 -

11. *Mid tibia – Predistal dorsal seta development: (0) strongly developed; (1) reduced; (2)

absent.

12. *Mid tibia – Distal posterodorsal seta development: (0) strongly developed; (1) reduced;

(2) absent.

13. Mid tibia – Ventral seta comb (male) development: (0) present; (1) absent.

14. Mid tibia – Midventral seta present in male: (0) yes; (1) no.

15. Mid basitarsus – basal posteroventral setulae development: (0) distinctly longer than apical

setulae; (1) indistinguishable from apical setulae or absent.

16. Hind tibia – dorsal seta: (0) absent; (1) present.

Male Abdomen

17. Sternite 5 – medial portion of posterior margin: (0) entire; (1) tab present; (2) emarginate.

18. Sternite 5 – lateral corners of emargination (when present): (0) simple; (1) tab/dentate.

19. Sternite 5 – secondary sclerite development: (0) absent; (1) present.

20. Sternite 5 – width of posteromedial emargination (if present): (0) < 1/4 width; (1) 1/4–1/2

width; (2) > 1/2 width.

21. Sternite 5 – lateral portion of posterior margin: (0) simple; (1) emarginate.

22. Sternite 6 – transverse portion: (0) straight; (1) broadly arcuate; (2) acutely arcuate.

23. Synsternite 6+7 – left medial process: (0) tip simple, acute; (1) tip modified. In some

species, the apical portion has been modified and is sometimes separated from the tip.

24. Epandrium – subanal plate complete: (0) no; (1) yes

25. Male Cercus – shape/development: (0) triangular; (1) triangular, but with distinct basal

constriction; (2) basally ovate, apical 2/3 acuminate

26. Surstylus – anterior lobe: (0) well-developed; (1) reduced, obsolete.

27. Surstylus – process on anterior lobe: (0) absent; (1) short (length< 2.0× basal width); (2)

elongate (length > 2.0× basal width)

28. Surstylus – posterior lobe: (0) simple, unmodified; (1) modified, with distinct projection.

29. Postgonite – shape: (0) basally quadrate with apical ½ acuminate; (1) entirely elongate.

- 127 -

30. Postgonite – dorsal margin morphology: (0) smooth; (1) swollen, protuberant near

midlength; (2) emarginate near midlength

31. Postgonite – development: (0) well developed; (1) reduced

32. Distiphallus – dorsal sclerite projecting beyond apex of acrophallus: (0) no; (1) yes

33. Distiphallus – acrophallus dorsolaterally near midlength: (0) rounded; (1) acutely angled.

34. Distiphallus – acrophallus with lateral projections: (0) no; (1) yes.

35. Distiphallus – acrophallus dorsolaterally reflexed near midlength of distiphallus: (0) no; (1)

yes.

36. Distiphallus – ventral acrophallus development: (0) poorly developed or absent; (1) well

developed and projecting ventrally.

37. Distiphallus – acrophallus sclerotization: (0) membranous; (1) sclerotized

Female Abdomen

38. Tergite 7 – posterior margin medially: (0) entire; (1) weakly emarginate (< 1/4 length of

sternite); (2) deeply emarginate (> 1/4 length of sclerite); (3) medially desclerotized

39. Tergite 7 – longitudinally divided: (0) no; (1) yes.

40. Tergite 7 – length across tergite: (0) relatively equal length throughout; (1) distinctly longer

medially.

41. Tergite 7 – posterolateral corner: (0) square, not projecting; (1) acutely angled.

42. Tergite 7 – posterolateral corner articulation with tergite 8: (0) broadly meeting tergite 8;

(1) acutely projecting to tergite 8.

43. Tergite 7 – length reduced laterally: (0) no; (1) yes

44. Tergite 8 – ventral projection: (0) rounded; (1) acutely angled.

45. Medial sclerite of tergite 8 – development: (0) well developed; (1) reduced in size; (2)

absent.

46. Medial sclerite of tergite 8 – shape: (0) elongate rectangular; (1) elongate triangular.

47. Epiproct – longitudinally divided: (0) no; (1) yes.

48. Epiproct – lateral sclerite morphology: (0) round/oval; (1) quadrate; (2) transverse; (3)

triangular. - 128 -

49. Epiproct – anteriorly extended: (0) no; (1) yes.

50. Epiproct – anterior margin of each lateral sclerite with lobes on inner surface: (0) absent;

(1) present.

51. Cercus – apical seta: (0) regular; (1) flattened.

52. Cercus – preapical seta flattened and appressed to base of apical seta: (0) no; (1) yes.

53. Cercus – Shape: (0) ovoid; (1) droplet-shaped.

54. Cercus – base longitudinally with distinct medial desclerotization: (0) no, (1) yes.

55. Sternite 8 – general shape: (0) transverse; (1) divided into pair of lateral triangular sclerites.

56. Sternite 8 – lateral sclerites articulating medially: (0) no; (1) yes.

57. Spermatheca – general shape: (0) barrel-shaped; (1) kidney-shaped; (2) ovoid; (3)

triangular.

58. Spermatheca – flattened: (0) no; (1) yes.

6.5.3 Molecular sequences

Cytochrome c oxidase subunit I (COI) 5P sequence data for Archiceroptera was obtained through two methods. Identified material for the four Archiceroptera species groups were submitted to the

Barcode Institute of Ontario for extraction, amplification and sequencing, where adequately preserved material was available. As no identified material was previously available on BOLD systems (http://www.boldsystems.org), public specimens/sequences with associated images were examined for unidentified Archiceroptera to increase the number of sequences available for study.

The prominens group was included as the outgroup and sequences were obtained from both submitted material and examination of unidentified public material. A total of 20 Archiceroptera and 4 prominens group sequences were imported from BOLD into Mesquite (Maddison and

Maddison 2017) and aligned by hand. Two datasets of this sequence data were exported into

PhyML (Guidon et al. 2010) for maximum likelihood analysis; one including all codons and one excluding the 3rd codon. The GTR+G substitution model with SPR tree improvement was used for the all inclusive dataset, while the dataset excluding the 3rd codon used the TN93+G substitution model and SPR tree improvement, based on the AIC estimates provided by PhyML. - 129 -

6.5.2 Discussion

Seventy equally parsimonious trees were recovered for the morphological data (length 220, consistency index = 34, retention index = 62) from 1,207,586 rearrangements, summarized here as a strict consensus tree (Fig. 6.1) and a majority rules consensus tree (Fig. 6.2). Bootstrap and

Jackknife values > 50 are given on the strict consensus tree. Characters were optimized on one of the 70 trees (Fig. 6.3). This tree was selected because it included the following species pairs, which are considered here to be sister species: A. venezolana and A. browni as sister species (based on shared reduction of the ocelli), A. megacerca and A. paracerca as sister species (based on the shared mid tibial chaetotaxy and male surstylus characteristics), A. adamas and A. maniba as sister species

(based on shared male sternite 5 characteristics), and A. calligraphia and A. dolabra as sister species

(based on shared male sternite 5 characteristics).

The two molecular phylogenies included substantially less species and provided slightly differing topologies from each other and from the morphological analysis. Both molecular and morphological analyses recovered the A. addenda group + [A. mahukani group + A. ternum group], but the topology of the larger A. ternum group varied between all datasets. The inclusive dataset

(Fig. 6.4A) and the dataset that excluded the 3rd codon (Fig. 6.4B) both recovered the A. brevivilla subgroup as sister to the rest of the A. ternum group, but differed in the topology of the rest of the A. ternum group. This may be a result of poor molecular representation from within the speciose A. ternum group, and inclusion of representatives of other clades would be required in future analyses to determine if this is a sampling artifact or if the A. brevivilla subgroup is sister to the A. ternum group.

Based on both the morphological and molecular analyses, the following clades are recognized: the A. addenda species group, A. mahukani species group, the A. brevivilla species subgroup, and the A. ternum species group. The first three clades were recovered in all 78 trees, while the A. ternum species group was recovered in 61 of the trees. Based on the tree depicted in

Figure 6.3, the A. addenda group was recovered as the basal-most clade supported by the following

- 130 - synapomorphies: strong anterior katepisternal seta, male sternite 5 with posterior secondary sclerite, distiphallus heavily sclerotized, and female cercus longitudinally desclerotized near the base. The remainder of Archiceroptera forms a clade supported by the following synapomorphies: the male cercus is ovoid with astrong ventral process, the distiphallus has a long dorsal sclerite, female tergite

7 is posterolaterally approximated or fused to the anterolateral corner of tergite 8, female tergite 8 with medial sclerite, epiproct longitudinally divided into a pair of sclerites, and female sternite 8 reduced to pair of elongate triangular sclerites. The Archiceroptera mahukani species group is supported by the following characters: dorsocentral setae in 3 or more pairs, anterior katepisternal seta extremely reduced or absent, and scutellum with 2 or more setulae near basal seta. The incomplete linkage between female tergite 7 and 8 supports its treatment as intermediate between the A. addenda group and the A. ternum group. The A. ternum group is well supported by the following synapomorphies: female tergite 7 projecting acutely to and fused with tergite 8 (e.g., Fig.

6.32B; secondarily broadened in a few species), lateral halves of epiproct with small lobes on anteromedial corner, and female sternite 8 with lateral sclerites medially articulating. The A. brevivilla species group is a well supported subgroup of the A. ternum species group based on the synapomorphic reduction of the mid tibial chaetotaxy, reduction of the male cerci and several female genitalic characters.

Archiceroptera megacercus and A. paracercus were recovered together and share a uniquely swollen dorsal margin of the postgonite along with extremely acuminate male cercus. Both species occur in Costa Rica and Ecuador. These two species were initially considered to be related to the Andean clade composed of A. bisetosus, A. basilia and A. bilobata, as they all have the female tergite 7 medially desclerotized and the same mid tibial chaetotaxy (a single strong anterodorsal seta and single strong posterodorsal seta near the midlength), but these two clades were recovered separately. The elongation of the epiproct of A. megacercus and A. paracercus could be considered an intermediate stage between the elongated anterior portion of the epiproct of the A. bisetosus group and the anteriorly rounded epiproct of the rest of Archiceroptera.

- 131 -

6.6 Key to the species of Archiceroptera Papp

Archiceroptera species identification is based largely on thoracic chaetotaxy, tibial chaetotaxy and genitalia. Identification of females to species can be difficult and some species cannot be currently identified using female morphological characters.

1. Scutellum with 2 or more basal setulae in addition to basal pair of marginal setae. Dorsocentral setae in 3 or more pairs (e.g. Fig. 6.18B). Hind tibia with 3+ large setae on dorsal surface. (A. mahukani group)…2

- Scutellum with 0–1 basal setulae. Dorsocentral seta in 1 prescutellar pair. Hind tibia rarely with more than 2 large setae on dorsal surface…4

2. Ocelli normal (> socket diameter of ocellar setae; Fig. 6.16C). Scutellum with long (> 0.5× length of basal seta) and short setulae (Fig. 6.16A). Spermathecae with small protuberance at junction with spermathecal duct (Fig. 6.17D)…A. mahukani Papp, 1977

- Ocelli reduced (< socket diameter of ocellar setae) or absent. Scutellum with all setulae short (<

0.4× length of basal seta). Females of A. venezolana with spermatheca without protuberance at junction with spermathecal duct (Fig. 6.7D; A. browni females unknown)…3

3. Ocelli present (Fig. 6.18A). Thorax with 3 or more strong dorsocentral setae. Scutellum with 2–6 small setulae near basal scutellar seta; scutellar setae distinctly longer than medial scutellar length

(Fig. 6.18C). Male surstylus posterodistally broadly rounded (Fig. 6.5E)…A. venezolana Richards,

1963

- Ocelli absent (Fig. 6.14C). Thorax with 2 strong dorsocentral setae and 1 weak anterior pair.

Scutellum with 18–20 small setulae near basal scutellar seta (Fig. 6.14B); scutellar setae shorter than medial scutellar length. Male surstylus posterodistally with narrow lobe (Fig. 6.15E)…A. browni Paiero & Marshall, n. sp.

4. Mid tibia with distinct posterodorsal setae near midlength and 4 or more apical setae; basal-most seta situated usually situated dorsally…5

- 132 -

- Mid tibia without posterodorsal setae on basal 2/3, with 2–3 apical setulae on apical 1/3; basal- most seta apparently anterodorsal (A. brevivilla species group)…31

5. Scutellum with single setula near base of basal seta. Male surstylus distinctly bilobed with small finger-like process near base of anterior lobe (Fig. 6.25D). Female tergite 7 with posterior margin sinuate, and weak medial desclerotized line extending from posterior margin to midlength; epiproct with elongate anterior projection ~1/2 length of posterior section (Fig. 6.26A)...A. bilobata Paiero &

Marshall n. sp.

- Scutellum without setula. Male surstylus variously shaped but not bilobate. Female tergite 7 variable, usually without medial desclerotization; epiproct variable (if with elongate anterior projection, tergite 7 is desclerotized medially, e.g., Fig. 6.28A)...6

6. Mid tibia with basal-most seta dorsal AND/OR wing with 2 equal or subequal costagial setae.

Male sternite 5 simple posteriorly, without additional sclerite. Female: tergite 8 medially divided, with distinct lateral triangular sclerites and small medial sclerite (e.g., Fig. 6.22A); cercus with large flattened apical seta with smaller flattened preapical seta appressed to base (difficult to see without dissection)...7

- Mid tibia with 2–3 anterodorsal and posterodorsal setae near mid length, but no basal dorsal seta present. Wing with 1 large costagial seta. Male with distinct sclerite posterior to sternite 5 (e.g., Fig.

6.8B). Female: tergite 8 entire or narrowly divided (e.g., Fig. 6.13A); cercus apically with 2 distinct flattened setae (e.g. Fig. 13C) and smaller preapical seta not appressed to larger apical seta. (A. addenda species group)...38

7. Hind tibia with 2 or more large dorsal setae on basal 2/3. Mid tibia EITHER with dorsal basal, and 1 anterodorsal and 1 posterodorsal setae near midlength (smaller seta basal to large anterodorsal and posterodorsal seta sometimes weakly developed), OR with dorsal basal seta and 2 anterodorsal setae near midlength (no posterodorsal setae present at midlength)...8

- Hind tibia with only uniformly small setulae dorsally. Mid tibia with 1–3 anterodorsal and posterodorsal setae present near midlength...10

- 133 -

8. Mid tibia with 2 anterodorsal setae and no posterodorsal setae at midlength. Hind tibia with 5–6 anterodorsal setae, 5–6 posterodorsal setae, and predistal dorsal seta. Male sternite 5 with 14-18 strong setae medially on posterior margin; surstylus anteriorly with elongate finger-like process, with small setae along length (Fig. 6.33D). Female tergite 7 entire and distinctly longer medially

(Fig. 6.34A); sternite 7 with posterior margin broadly emarginate (Fig. 6.34C)...A. cobolorum Paiero

& Marshall, n. sp.

- Mid tibia usually with 1 anterodorsal seta and 1 posterodorsal seta at midlength (rare cases have the basal anterodorsal and posterodorsal present but reduced). Hind tibia with proximal anterodorsal and posterodorsal setae on basal third (anterodorsal seta slightly distal to posterodorsal). Male sternite 5 without series of long posteromedial setae (Fig. 6.40B and 6.45B); surstylus boot-shaped, with anterior triangular lobe bare (Figs. 6.40BD and 6.45E). Female (unknown for A. paracerca) with tergite 7 deeply emarginate and not distinctively longer medially (Fig. 6.41A); sternite 7 with posterior margin broadly rounded (Fig. 6.41C) ...9

9. Male sternite 5 with shallow posteromedial emargination delimited on each side by small rounded tabs (Fig. 6.40B); anterior lobe of surstylus triangular and evenly narrowed towards tip (Fig. 6.40D); postgonite with large quadrate base longer than apical part (Fig. 6.40E). Female tergite 7 with deep oval emargination extending from posterior margin to basal ¼ (Fig. 6.41A)...A. megacercus Paiero

& Marshall, n. sp.

- Male sternite 5 with posterior margin entire or only weakly emarginate (Fig. 6.54B); anterior lobe of surstylus triangular with distinctly preapical constriction (Fig. 6.54E); postgonite with rounded circular base and elongate apical part that is longer than base (Fig. 6.54F). Female unknown...A. paracerca Paiero & Marshall, n. sp.

10. Mid tibia with 1 anterodorsal and 1 posterodorsal bristle present at midlength (smaller seta rarely present); male with ventral comb weak or indistinct, and midventral seta present. Female tergite 7 medially longitudinally divided and epiproct with narrow anterior projection (Figs. 6.24A and 6.28A)...11

- Mid tibia with 2+ anterodorsal and 2+ posterodorsal setae present at midlength; male with distinct

- 134 - ventral comb, and midventral seta absent (most species) or present (A. masoni). Female tergite 7 not longitudinally divided, and epiproct without distinct anterior projection (e.g., Fig. 6.32A)...12

11. Male sternite 5 with pair of elongate setae on disc anterior to distinct medial emargination of posterior margin (Fig. 6.27B); surstylus (in lateral view) with dark, narrow posteroapical process, and anterior lobe preapically narrowed to make a short anterior process (Fig. 6.27D); cercus (in posterior view) triangular with 3–4 large seta (Fig. 6.27C); postgonite wedge-shaped (Fig. 6.27E).

Female epiproct with narrow anterior projection ~1/2 as long as broad posterior section (Fig.

6.28A)...A. bisetosus Paiero & Marshall n. sp.

- Male sternite 5 posteriorly with pair of apically rounded elongate processes and without elongate setae on disc (Fig. 6.23B); surstylus (in lateral view) anteriorly triangular, posteriorly rounded, and posterior half setose (Fig. 6.23D); cercus, in posterior view, abruptly constricted on apical 1/3, not swollen basally (Fig. 6.23C); postgonite narrowed on apical third, with apical ¼ elongate and narrow (Fig. 6.23G). Female epiproct with narrow anterior projection as long as broad posterior section (Fig. 6.24A)...A. basilia Paiero & Marshall, n. sp.

12. Males...13

- Females...24

13. Posterior margin of sternite 5 with distinctly projecting lobes adjacent to medial emargination

(e.g., Figs. 6.45B and 6.57B)...14

- Sternite 5 posteriorly truncate, weakly emarginate or large emargination, but never with distinct lobes present (e.g., Figs. 6.40B and 6.59B)...18

14. Posterior margin of sternite 5 with long posteriorly projecting lobes on each side of emargination; medial emargination broadly open and not distinctly constricted posteriorly (e.g., Fig.

6.35B and 6.41C)...15

- Posterior margin of sternite 5 with medially projecting lobes on each side of emargination; medial emargination posteriorly constricted, with posterior width < ½ greatest width (e.g., Figs. 6.21B and

6.48B)...17

- 135 -

15. Mid tibia with predistal dorsal and apical posterodorsal setae reduced, distinctly smaller than other distal setae. Sternite 5 posterior margin with rounded lobes on each side of narrow, but shallow, emargination (Fig. 6.41C). Surstylus boot-shaped, with short anterior process (Fig. 6.41D).

Transverse part of sternite 6 weakly arcuate (Fig. 6.41C). Postgonite with apical 1/3 wedge-shaped

(Fig. 6.41E)...A. mexicorona Paiero & Marshall, n. sp.

- Mid tibia with predistal dorsal and apical posterodorsal setae well developed, as large or slightly shorter than other distal setae. Sternite 5 posterior margin with acute lobes on each side of a deep and broad emargination (e.g., Fig. 6.45B). Surstylus strap-like (Fig. 6.45D) or weakly clavate (Fig.

6.18D), with small anterior process on apical 1/3. Transverse part of sternite 6 strongly arcuate medially (Fig. 6.45B). Postgonite with apical 1/3 elongate and narrow...16

16. Anterior katepisternal seta absent. Mid basitarsomere with posteroventral setae equal in length.

Sternite 5 emargination triangular, posterolateral corners of emargination with tab with 4–6 setae

(claw-like in appearance, Fig. 6.45B)...A. maniba Paiero & Marshall, n. sp.

- Anterior katepisternal seta weak, but present. Mid basitarsomere with basal posteroventral seta longer than distal setae. Sternite 5 with diamond-shaped emargination (Fig. 6.18B); posterolateral corners of emargination acute...A. adamas Paiero & Marshall, n. sp.

17. Sternite 5 well sclerotized and emargination well defined, with rounded clavate lobe on each side of emargination; medially projecting lobes consisting of anterior section that is setose, and posterior section with series of 15–20 dark flattened setae along posterior margin (anterior and posterior sections separated by lighter sclerotization). Surstylus with elongate setae on posterior surface shorter than surstylus (Fig. 6.48D). Cercus (in posterior view) gradually narrowed (Fig.

6.48C)...A. ternum Paiero & Marshall, n. sp.

- Sternite 5 with emargination poorly defined (sometimes more elongate or rounded, or indistinct); posterior margin with medially projecting lobes on each side of emargination acute, nearly meeting medially, and with pale setae along posterior margin (Fig. 6.21B). Surstylus with elongate setae on posterior surface longer than surstylus (Fig. 6.21D). Cercus (in posterior view) distinctly constricted near basal 1/3 (Fig. 6.21C)...A. barberi Paiero & Marshall, n. sp.

- 136 -

18. Sternite 5 posterior margin deeply emarignate; medial length of emargination > 1.0× greatest width (e.g., Fig. 6.31B)...19

- Sternite 5 posterior margin entire or weakly emarginate; if emarginate, emargination shallow with greatest length < 2/3 greatest width (e.g., Fig. 6.38B)...21

19. Sternite 4 posteromedially with broad triangular emargination (Fig. 6.35A). Surstylus anteriorly with triangular, acutely angled projection (Fig. 6.35D). Postgonite ventrally flat, with small dorsal swelling near apical 1/5 (Fig. 6.35E)... A. dolabra Paiero & Marshall, n. sp.

- Sternite 4 posteromedially entire (e.g., Fig. 6.31A). Surstylus anteriorly with broad triangular projection (Figs. 6.31D and 6.44D). Postgonite ventrally sinuate (e.g., Fig. 6.31E); apical 1/5 smooth dorsally...20

20. Sternite 5 deeply emarginate, with emargination ended just short of anterior margin of sternite

(Fig. 6.31B). Transverse portion of sternite 6 weakly arcuate (Fig. 6.31B). Surstylus boot-like, with distinct posteroapical process extending beyond anterior triangular process (Fig. 6.31D)...A. calligraphia Paiero & Marshall, n. sp.

- Sternite 5 with broad triangular emargination extending ~1/2 length of sternite (Fig. 6.43B).

Transverse portion of sternite 6 abruptly and distinctly arcuate (Fig. 6.44B). Surstylus weakly clavate, with distal 1/3 wider than basal 2/3 (Fig. 6.44D)...A. mitarakai Paiero & Marshall, n. sp

21. Surstylus anteriorly with short, basally projecting process (Fig. 6.50D). Sternite 6 with transverse portion distinctly arcuate; left corner of arcuate portion swollen (Fig. 6.50B) ...A. uncinata Paiero & Marshall, n. sp.

- Surstylus either without distinct anterior process, or process broad and/or projecting anteriorly.

Sternite 6 with transverse portion sinuate or straight, without any distinctive swellings...22

22. Mid tibia with midventral seta present. Sternite 5 posteromedially weakly sinuate with fringe of weak setulae on middle 1/3 (Fig. 6.38B). Surstylus with small, narrow, anteriorly projecting finger- like process (Fig. 6.38D)...A. masoni Paiero & Marshall, n. sp.

- 137 -

- Mid tibia with mid ventral seta absent. Sternite 5 posteromedially with broad triangular emargination (e.g., Fig. 6.46B). Surstylus with broad anterior projection (e.g., Fig. 6.46D)...23

23. Sternite 5 emargination usually terminating anteriorly at dark sclerotized fold (Fig. 6.29B).

Surstylus boot-like, with anterior projection evenly curved (Fig. 6.29D)...A. caliga Paiero &

Marshall n. sp.

- Sternite 5 emargination without anterior fold (Fig. 6.46B). Surstylus with anterior projection distally sinuate, ending in small nipple-like swelling (Fig. 6.46D)...A. pussula Paiero & Marshall n. sp.

24. (Couplets 24-30,females, not known for A. dolabra and A. mitarakai). Tergite 7 narrowing laterally (lateral length < 0.75 medial length); posteromedially bilobate (Fig. 6.22C and 6.43C) ...25

Tergite 7 not distinctly narrowed laterally (lateral length > 0.9× medial length); posteromedially weakly sinuate or truncate (e.g., Fig. 6.20B)...26

25. Mid tibia with predistal dorsal and apical posterodorsal setae strong, equal or only slightly smaller than other distal setae. Sternite 7 posteromedially narrowly rounded (Fig. 6.22C)...A. barberi Paiero & Marshall, n. sp.

- Mid tibia with predistal dorsal and apical posterodorsal setae reduced, smaller than other distal setae. Sternite 7 posteromedially acutely angled (Fig. 6.43C)...A. mexicorona Paiero & Marshall, n. sp.

26. Epiproct rectangular, anterior margin transverse (Fig. 6.32A). Sternite 7 posteromedially acute, narrowly rounded (Fig. 6.32C)...A. calligraphia Paiero & Marshall, n. sp.

- Epiproct triangular or rounded, anterior margin rounded or acute (e.g. Fig. 6.49A and 6.39A).

Sternite 7 posteromedially broadly rounded (e.g., Fig. 6.49C and 6.30C)...27

27. Medial part of tergite 8 parallel sided (Fig. 6.49A) ...A. ternum Paiero & Marshall, n. sp.

- Medial part of tergite 8 triangular with posterior margin distinctly wider than anterior margin (e.g.,

Figs. 6.20C and 6.30C)...28

- 138 -

28. Spermatheca triangular (Fig. 6.30D)...A. caliga Paiero & Marshall, n. sp.

- Spermatheca ovoid (e.g., Fig. 6.20)...29

29. Tergite 7, in lateral view, with anterolateral corner abruptly rounded (anterior width of rounding

< 0.5× length of tergite; Fig. 6.20B and 6.37B)...30

- Tergite 7, in lateral view, with anterolateral corner broadly rounded (anterior width of rounded portion > 0.5× length of tergite; Fig. 6.47B and 6.51B)...A. pussula and A. uncinata (currently not separable)

30. Tergite 8 with anterior margin weakly sinuate laterally, before anterolateral corner (Fig.

6.20B)...... A. adamas Paiero & Marshall n. sp.

- Tergite 8 with anterior margin evenly rounded before anterolateral corner (Fig. 6.37B)...A. maniba

Paiero & Marshall n. sp.

31. Male sternite 5 posteriorly with row of 6–9 dark robust setae (Fig. 6.52B); surstylus base tapering apically, with long anterior finger-like projection (as long or longer than surstylus length;

Fig. 6.52D). Female sternite 7 with tip deeply emarginate (Fig. 6.53C)...A. braziliensis Paiero &

Marshall, n. sp.

- Male sternite 5 posteriorly with > 10 pale setae along posterior margin (e.g., Figs. 6.54B and

6.56B). Surstylus with anterior projection variable. Female sternite 7 entire, rounded...32

32. Male sternite 5 posteromedially with broad emargination and dense setal cluster of 10+ pale setae confined to poorly sclerotized “pad” present on each side of posteromedial emargination (Fig.

6.58B); surstylus anteriorly with 3 small finger-like projections (Fig. 6.58D). Female tergite 7 posteromedially broadly rounded (Fig. 6.59A); spermatheca triangular in profile (Fig. 6.59D)...A. llama Paiero & Marshall, n. sp.

- Male sternite 5 posteromedially sinuate or with projecting tabs but never with broad emargination and setose pads (e.g., Fig. 6.56B); surstylus anteriorly with single anterior projection. Female tergite

7 truncate; spermatheca ovoid...33

- 139 -

33. Males...34

- Females...36

34. Surstylus anteriorly with elongate narrow process as long as basal width of surstylus (Fig.

6.60D); posterior margin of sternite 5 weakly sinuate, with medial emargination filled with transverse row of setae (Fig. 6.60B); distiphallus with lateral acrophallic flaps recurved dorsally

(Fig. 6.60F-H) ...A. megavilla Paiero & Marshall, n. sp.

- Surstylus anteriorly with short (straight or curved) finger-like projection no longer than ½ basal width of surstylus (e.g., Fig. 6.56D); posterior margin of sternite 5 as above (A. brevivilla, Fig.

6.54B) or with flattened setae present on 2 broadly rounded posterior lobes (A. curvivilla, Fig.

6.56B); lateral acrophallic flaps projecting laterally, not recurved dorsally (Figs. 6.54F–H and

6.56F–H) ...35

35. Surstylus anteriorly with straight, short finger-like process (Fig. 6.54D). Sternite 5 posteromedially with unbroken series of 20–25 setae present (Fig. 6.54B). Cercus basally swollen

(Fig. 6.54C)...A. brevivilla Paiero & Marshall, n. sp.

- Surstylus anteriorly with curved finger-like process (Fig. 6.56D). Sternite 5 posteromedially projecting and medially sinuate (appearing bilobed); each “lobe” with series of 9–11 setulae (Fig.

6.56B). Cercus base flat, not swollen (Fig. 6.56C)...A. curvavilla Paiero & Marshall, n. sp.

36. Sternite 8 with posterior margin angulate, narrowly rounded (Fig. 6.55C). Tergite 8 with posterolateral corner distinctly projecting (Fig. 6.55B)...A. brevivilla Paiero & Marshall n. sp.

- Sternite 8 with posterior margin broadly rounded (Fig. 6.23C and 6.27C). Tergite 8 with posterolateral corner rounded, not projecting (Fig. 6.57B and 6.61B)...A. curvivilla and A. megavilla

(females of these species are not separable)

37. Male sternite 5 width ~2.5× length; secondary sclerite on posterior margin with flat, crenulate lobe to the left of middle, and a broad, weakly bilobed process medially (Fig. 6.10B). Postgonite extremely reduced (Fig. 6.10H), ~1/2 length of cercus. Female epiproct not narrowed medially (Fig.

6.11A). Paired spermathecae extremely small (< 1/10th of single spermatheca; Fig.

- 140 -

6.11D)...Archiceroptera crenulata Paiero & Marshall n. sp.

- Male sternite 5 width ~3.0× length; secondary sclerite on posterior margin various, not as described above. Postgonite well developed, > ½ length of cercus (e.g., Fig. 6.12D). Female epiproct narrowed medially (e.g., Fig. 6.13A). Paired spermathecae well developed, equal in size to single spermatheca (e.g., Fig. 6.13D)...2

38. Male sternite 5 with pair of irregular processes (Fig. 6.8B). Male cercus only ½ length of surstylus. Surstylus broadly rounded apically (Fig. 6.8D). Female with epiproct divided medially

(Fig. 6.9A). Female cercus with medial sclerite cerci elongate triangular (Fig.

6.9A)...Archiceroptera addenda Paiero & Marshall n. sp.

-Male sternite 5 with 3 acute processes (one large acute medial process and two lateral rounded processes; Fig. 6.12B). Male cercus ~2/3 length of surstylus (Fig. 6.12D); surstylus triangular; with constriction on apical 2/5 to make tip appear abruptly widened (Fig. 6.12E). Female epiproct entire, transverse, with broad posteromedial emargination (Fig. 6.13A); medial sclerite of cerci elongate oval (Fig. 6.13A)...Archiceroptera triclavus Paiero & Marshall n. sp.

6.7 Species Descriptions

6.7.1 Archiceroptera addenda species group

The A. addenda species group is apparently the sister group to the rest of Archiceroptera, and is characterized by derived female terminalia (epiproct medially divided, tergite 8 is tripartite). It includes three newly described species: A. addenda, A. crenulata and A. triclavus. Members of this species group can be recognized in having 3–4 median anterodorsal and 3–4 median posterodorsal setae, a strong costagial seta, a secondary sclerite posterior to the male sternite 5, and a strongly sclerotized distiphallus. The females tergite 8 and the epiproct are both plesiomorphic for

Archiceroptera; the medial part of tergite 8 is absent or poorly developed, and the epiproct is either entire (A. crenulata and A. triclavus) or only narrowly divided (A. addenda). - 141 -

A. addenda species group description:

Head: Frons with 4–5 interfrontal setae. Gena with 2 strong anterior setae and additional smaller posterior setae.

Thorax: Scutum with 1 prescutellar dorsocentral seta. Acrostichal setulae in 6 rows between prescutellar dorsocentral seta. Anterior katepisternal seta strongly developed (> 1/2 posterior katepisternal seta). Scutellum without setulae.

Mid tibia with dorsal basal seta, 2–3 median anterodorsal and 2–3 median posterodorsal setae; strong predistal anterodorsal, strong predistal dorsal, and strong distal posterodorsal setae present; male without ventral comb; midventral seta present in female (sometimes present in male). Mid basitarsus with basal posteroventral setae slightly longer than distal setae. Hind tibia with small preapical seta; additional seta sometimes present.

Wing: C2:C3 ratio = 4:3. Costa with 2 strong costagial setae; 1 longer and extending beyond humeral crossvein. Cell dm without CuA1 stub vein.

Male Abdomen: Sternite 4 posteromedially entire. Sternite 5 with additional posterior sclerite; posterior sclerite shape species specific. Surstylus variable. Cercus elongate triangular. Distiphallus with acrophallus heavily sclerotized.

Female Abdomen: Tergite 7 posteromedially entire; posterolaterally separate from tergite 8. Tergite

8 medially divided, with medial sclerite poorly developed or absent. Epiproct transverse, hirsute; medially entire or divided. posteromedially with arcuate emargination almost reaching anterior margin. Cercus strap-like, with basal 1/3 longitudinally weakened or divided; strong apical and weaker preapical setae weakly separated at their base. Sternite 7 broadly rounded. Sternite 8 weakly sclerotized; medially divided into pair of lateral sclerites (triangular but shape variable between species) and transverse posterior sclerite. Spermatheca with stems short (< 1/2 spermathecal width).

Archiceroptera addenda Paiero & Marshall, n. sp.

Figs. 6.8–6.9

Size: 2.0–2.8 mm. - 142 -

Head: Ocellar triangle with 6–7 setae. Frons with 3–5 inclinate orbital setulae. Eye:gena ratio =

2.5:1. Gena with 7–14 small setae.

Legs: Fore femur with 6–7 posterodorsal and 5–7 posteroventral setae. Male mid femur ventrally with 15–25 unmodified setae on basal 1/2. Mid tibia with 3 median anterodorsal and 2–3 median posterodorsal setae; midventral seta present in both sexes. Hind tibia with 4–5 posterodorsal setae.

Wing: Distance between r-m and dm-cu ~5.0× dm-cu. Cell dm without CuA1 stub vein.

Male Abdomen (Fig. 6.8): Sternite 5 with broad shallow emargination; secondary sclerite with two irregular lateral lobes (right lobe elongate with weak apical emargination; left lobe with basolateral tooth and narrowed on apical 1/5). Transverse portion of sternite 6 straight; right lateral portion recurved and fused with small quadrate sclerite. Synsternite 6+7 with narrowly-rounded, medially projecting process ending in irregular sclerite. Cercus triangular, with long seta (sometime 2) near apical 1/3 and smaller seta near basal 1/3. Surstylus (in lateral view) broadly oval, not entending anteriorly; surface hirsute with 15–17 small setae. Postgonite elongate, acuminate (sometimes received within excavation of acrophallus. Distiphallus: dorsal sclerite not extending beyond apex of acrophallus (usually divergent); acrophallus heavily sclerotized (inner sclerites difficult to distinguish), expanded basally with broad, rounded lobes to encompass postgonites.

Female Abdomen (Fig. 6.9): Tergite 7 posterolaterally quadrate. Tergite 8 with lateral sclerites posteriorly acute and small medial sclerite near posterior margin. Epiproct medially divided. Cercus with basal 1/3 longitudinally desclerotized; preapical seta ~2/3 length of apical seta. Sternite 8 with lateral sclerites triangular. Spermatheca barrel-shaped, surface striate; pair with stems short.

Type Material: Holotype (male, debu00260837, QCAZ) + 1 paratype (1 male): ECUADOR:

Esmeraldas: La Chiquita, 11 km SE San Lorenzo, 5 m, carrion trap, day 2, 7–8 Jun 1975, S. Peck.

Additional Paratypes: BOLIVIA: La Paz: 1 male, Arroyo Tuhiri W Mapiri, 15°17'27"S,

68°15'29"W, 10 Apr 2001, S.A. Marshall (UASC). ECUADOR: Esmeraldas: 4 males, 7 females,

La Chiquita, 11 km SE San Lorenzo, 5 m, dung, 9–10 Jun 1975, S. Peck (QCAZ); 6 males, 6 females, La Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, 10–11 Jun 1975, S. Peck; 1 male, La

Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, day 3, 8–9 Jun 1975, S. Peck; 1 male, 2 female,

La Chiquita, 17km SE San Lorenzo, 5m, dung, 7–8 Jun 1975, S. Peck; Napo: 1 female, Tiputini - 143 -

Biodiversity Stn., 0°36'50"S, 76°9'1"W, sweep, May 2011, S.A. Marshall. FRENCH GUIANA: St.

Laurent du Maroni: 1 male, 2 females, Maripasoula, Mitaraka, MIT-DZ, 2°14'2"N, 54°27'1"W,

306m, tropical moist forest (plateau-slope-cleared), FIT, 1 Mar 2015, Touroult & Poirier (MHNM);

2 males, Maripasoula, Mitaraka, MIT-DZ, 2°14'2"N, 54°27'1"W, 306m, tropical moist forest near

DZ, FIT, 6–10 Mar 2015, Touroult & Poirier. PANAMA: 4 males, 6 females, Canal Zone, Madden

Forest, carrion trap, 10–13 Jun 1977, S. Peck. PERU: Cusco: 1 male, Cock-of-the-Rock Lodge, NE

Paucartambo, 13°3'18"S, 71°32'42"W, 1120 m, FIT, 4–9 Nov 2007, D. Brzoska; Loreto: 2 males, 1 female, Campamento San Jacinto, 175–215 m, FIT, 10 Jul 1993, R. Leschen; 2 males, 5 females,

Campamento San Jacinto, 175–215 m, FIT, 11 Jul 1993, R. Leschen; 2 males, Campamento San

Jacinto, 175–215 m, FIT, 3 Jul 1993, R. Leschen (MUSM); 1 male, Campamento San Jacinto, 175–

215 m, FIT, 3 Jun 1993, R. Leschen (MUSM); 4 males, 2 females, Campamento San Jacinto, 175–

215 m, FIT, 5 Jul 1993, R. Leschen; 5 males, 2 females, Campamento San Jacinto, 175–215 m, FIT,

7 Jul 1993, R. Leschen; 2 females, Campamento San Jacinto, 175–215 m, in primary forest, FIT, 8

Jul 1993, R. Leschen (MUSM); 1 female, Campamento San Jacinto, 175–215 m, FIT, 9 Jul 1993, R.

Leschen (MUSM); 3 males, 6 females, Teniente López, FIT, 23 Jul 1993, R. Leschen; 1 male, 1 female, Teniente López, FIT, 26 Jul 1993, R. Leschen; 3 males, 1 female, Teniente López, 1.5 km

N, 2°31'S, 76°10'W, 230–305 m, FIT, 18 Jul 1993, R. Leschen (MUSM); 1 male, 1 female, Teniente

López, 1.5 km N, 2°31'S, 76°10'W, 230–305 m, FIT, 20 Jul 1993, R. Leschen; 2 females, Teniente

López, 1.5 km N, 2°31'S, 76°10'W, 230–305 m, FIT, 22 Jul 1993, R. Leschen; 1 male, 2 females,

San Jacinto, FIT, 12 Jul 1993, R. Leschen; Madre de Dios: 1 female, CICRA, trail 2, 12°33'40"S,

70°6'23"W, 267m, Malaise, 22–23 Nov 2013, J. Caballero; 1 male, Pantiacolla Lodge, Alto Madre de Dios River, 12°39'18"S, 71°13'54"W, 420 m, FIT, 14–19 Nov 2007, D. Brzoska.

Etymology: The species’ name refers to the additional sclerite posterior to male sternite 5.

Archiceroptera crenulata Paiero & Marshall, n. sp.

Figs. 6.10–6.11

Size: 1.5–1.9 mm.

- 144 -

Head: Ocellar triangle with 8–10 setae. Frons with 3–5 inclinate orbital setulae. Eye:gena ratio =

2.5:1. Gena with 6–12 small setae.

Legs: Fore femur with 5–6 posterodorsal and 6–8 posteroventral setae. Male mid femur ventrally with 20–25 regular setae in cluster on basal 1/3, with short series of setae apically on posteroventral margin. Mid tibia with 3 median anterodorsal and 2–3 median posterodorsal setae; midventral seta present only in female. Hind tibia simple (except for preapical seta).

Wings: Distance between r-m and dm-cu ~4.0× dm-cu.

Male Abdomen (Fig. 6.10): Sternite 5 posteriorly with broad, shallow emargination; secondary sclerite with medioventral, furcate lobe and a posterior crenulate lobe to the left of middle.

Transverse portion of sternite 6 straight; right lateral portion recurved, simple. Synsternite with with broadly rounded, medially projecting process (often difficult to discern from posteior margin of transverse portion and secondary sclerite). Epandrium broadly meeting below anal opening. Cercus triangular with 1–2 setae near basal third and 1 near apical third. Surstylus (in lateral view) strap- like, elongate, with small finger-like process anterobasally and distal posterior corner with short triangular lobe. Postgonite small, triangular. Distiphallus: dorsal sclerite not reaching apex of acrophallus; acrophallus weakly sclerotized, ventrally with elongate bifid process, and dorsolaterally rounded projections near midlength of distiphallus.

Female Abdomen (Fig. 6.11): Tergite 7 posterolaterally quadrate. Tergite 8 with lateral sclerites posterolaterally rounded; medial sclerite absent or extremely weak and small. Epiproct medially entire. Cercus longitudinally divided on basal 1/2, with inner sclerite elongate; preapical seta ~1/2 length of apical seta. Sternite 8 with lateral sclerites triangular with ventral surface emarginate

(boomerang-shaped). Spermathecae ovoid with small swelling at duct junction; sclerotized portion of duct < 1/4 length of spermatheca); pair extremely reduced, with stems very short.

Type Material: Holotype (male, debu01084487): COLOMBIA: Leticia, 28 Feb 1974, V. Nealis.

Paratypes: COLOMBIA:1 male, Leticia, dung traps, 28 Feb 1974, V. Nealis. FRENCH

GUIANA: St. Laurent du Maroni: 1 male, Maripasoula, Mitaraka, MIT-A-SL, 2°14'18"N,

54°27'8"W, 352m, tropical moist forest (slope), yellow pan traps, 3–8 Mar 2015, M. Pollet

(MHNM); 1 female, Maripasoula, Mitaraka, MIT-C-TOP, 2°13'59"N, 54°26'38"W, 433m, tropical - 145 - moist forest (plateau), white pan traps, 2–8 Mar 2015, M. Pollet. VENEZUELA: Bolivar: 3 males

3 females, km 40 Sta. Elena Icabaru Rd., 100m, 4–6 Aug 1986, B. Gill (DEBU, MIZA); 1 male, km

40 Sta. Elena Icabaru Rd., 220m, 4–6 Aug 1986, B. Gill (MIZA).

Etymology: The species epithet refers to the crenulate margin of the secondary sclerite posterior to sternite 5.

Archiceroptera triclavus Paiero & Marshall, n. sp.

Figs. 6.12–6.13

Size: 1.7–2.1 mm.

Head: Ocellar triangle with 5–8 setae. Frons with 2–4 inclinate orbital setulae. Eye:gena ratio = 3:1.

Gena with 8–10 small setae.

Legs: Fore femur with 6–7 posterodorsal and 5–7 posteroventral setae. Male mid femuir ventrally with 15–25 regular setae on basal 1/3, and extending along posteroventral margin. Mid tibia with 2 median anterodorsal and 3 median posterodorsal setae; midventral seta present in both sexes. Hind tibia with 4–5 posterodorsal setae.

Wings: Distance between r-m and dm-cu ~5.0× dm-cu.

Male Abdomen (Fig. 6.12): Sternite 5 posteromedially weakly emarginate; secondary sclerite trilobed with pair of lateral rounded lobes and central truncate lobe; central lobe with acute process projecting off ventral surface. Transverse portion of sternite 6 straight; right lateral portion recurved, simple. Synsternite with with narrowly rounded, medially projecitng process; process with apical

1/3 curved anteriorly. Cercus triangular, with long seta near apical 1/3. Surstylus (in lateral view) strap-like, elongate, with distinct constriction near distal 1/3. Postgonite elongate, acuminate.

Distiphallus: dorsal sclerite not projecting beyond apex of acrophallus, but tips abruptly divergent; acrophallus heavily sclerotized, with inner sclerites nearly indistinguishable from acrophallus.

Female Abdomen (Fig. 6.13): Tergite 7 posterolaterally rounded. Tergite 8 with lateral sclerites broadly rounded; medial sclerite narrow and elongate, extremely weak and posterior to tergite 7.

Epiproct entire; posteromedially with arcuate emargination that nearly extends to anterior margin.

Cercus nearly longitudinally divided with narrowly bridge near midlength; inner sclerite ovoid; - 146 - preapical setae ~1/2× length of apical seta). Sternite 8 lateral sclerites triangular. Spermatheca oblong, with distinct ribbing and elongate conical projection towards duct junction; sclerotized portion of duct < 1/4 length of spermatheca), difficult to separate from conical projection; paired spermathecae with short stems.

Type Material: Holotype (male, debu01084483, MZSP) + 1 Paratype (1 female): BRAZIL:

Bahia: Porto Segure, 15 km NE, Ecological Reserva “Pau-Brasil”, 1° Atlantic Forest, Shannon-trap,

19–27 Feb 1986, Daltoas, Cristina & Amaorim. Additional Paratypes: BRAZIL: Bahia: 1 female,

Encruzilhada, malaise trap, Nov 1973, Seara & Roppa (MZSP); Pará: 1 female, Tucuruí, Jan 1979,

M. Alvarensa. GUYANA: Potaro-Siparuni: 1 female, Mount Wokomung, 5°6'35”N, 59°49'15”W,

1234m, 1° rainforest, human dung, pitfall trap, 27 Oct–1 Nov 2004, B. Hubley (ROME);

Mazaruni-Potaro: 1 male, 1 female, Potaro River, E side, downstream Tukeit Falls, 300', 1° rainforest, dry stream bed, malaise-coarse, 26–30 Sep 1990, Hubley & Coote; Rupununi: 1 male,

Kurupukari, Essequibo River, 200', 1° rainforest clearing, malaise, 7–11 Oct 1990, Coote & Hubley

(ROME).

Etymology: The species epithet refers to the three nail-like processes on the extension of sternite 5;

“clavus” is Latin for nail.

6.7.2 Archiceroptera mahukani species group

The A. mahukani species group includes three species, including one newly described species (A. browni) and the only two species previously described in the genus Archiceroptera. The group is characterized by three or more dorsocentral setae and two or more setulae on the scutellum near the basal pair of setae. Species in this group also have more dorsal setae on the mid and hind tibia than other Archiceroptera.

A. mahukani species group description

Head: Frons with 4–6 interfrontal setae. Gena with 2 enlarged long setae and additional smaller posterior setae.

- 147 -

Thorax: Scutum with 2–7 dorsocentral setae (including prescutellar pair). Acrostichal setulae in 6-8 rows between prescutellar dorsocentral setae. Anterior katepisternal seta absent. Scutellum with 2 or more setulae near base of basal scutellar seta.

Legs: Mid tibia with dorsal basal seta, 2–3 median anterodorsal and 2–3 median posterodosal setae; predistal anterodorsal and predistal dorsal setae present; male with ventral comb; midventral seta present only in female. Mid basitarsus with basal posteroventral setavariable in length to distal setae. Hind tibia with 2 or more dorsal setae.

Wings: Costa with 2 strong costagial setae. Cell dm without CuA1 stub vein present.

Male abdomen: Sternite 4 posterior margin entire. Sternite 5 posteromedially with tab-like projection. Surstylus variable. Cercus with basal ¼-1/3 ovoid, with apical portion narrow and acuminate. Distiphallus with well-developed membranous acrophallus.

Female abdomen: Tergite 7 posteromedially entire; posterolaterally either closely approximated with, or broadly fused to tergite 8. Tergite 8 with medial sclerite present. Epiproct medially desclerotized, with lateral portions transverse, broadly rounded anteriorly; with scattered small setulae. Cercus ovoid/tear-drop shaped with strong flattened apical seta and smaller preapical seta

(not flattened). Sternite 7 with posterior margin rounded. Sternite 8 entire or weakly divided medially into pair of triangular sclerites. Spermathecae barrel-shaped with swelling at duct junction and small finger-like invagination at opposite ends; sclerotized portion of duct < 1/2 length of spermatheca; paired spermathecae with stems short.

Archiceroptera browni Paiero & Marshall, n. sp.

Figs. 6.14–6.15

Description:

Size: 2.4mm

Head: Ocelli absent with 13 small setulae on ocellar triangle. Frons with 6 interfrontal seta pairs and 6 inclinate orbital setulae. Eye:gena ratio = 5:2. Gena with 11 smaller setae.

Thorax: Dorsocentral setae in 2 pair (including prescutellar); acrostichal setae in 8 rows between prescutellar dorsocentral setae. Scutellum with 18-20 setae near basal seta. - 148 -

Legs: Fore femur with 5 posterodorsal and 5 posteroventral setae. Mid femur anteriorly with series of 5 stout setae extending from base to apical 3/4; apicoventral series of 3-5 setae on apical 1/4.; male, ventrally, with 12 setae in small, circular basal cluster. Mid tibia with 3 median anterodorsal and 2 median posterodorsal setae; predistal anterodorsal seta weaker than predistal dorsal seta; distal posterior seta absent; male, ventrally, with 6-7 dark setae on apical 1/3. Mid basitarsus with basal posteroventral setae not enlarged. Hind tibia with 2-3 predistal anterodorsal setae, 4-5 posterodorsal setae (apical 2 weak, absent on one side), and with predistal dorsal seta present.

Wings: Costagial setae, extending to humeral crossvein. C2:C3 ratio = 4:3. Distance between r-m and dm-cu ~4.0× dm-cu.

Male Abdomen (Fig. 6.15):. Sternite 5 posteromedial medially narrowed and with 4-5 flattened setae on lateral thirds. Transverse portion of sternite 6 straight, right side simple. Cercus with basal

1/3 ovoid and with large distinct seta; apical 2/3 narrow and acute. Surstylus (in lateral view) quadrate with acutely projecting anterodistal corner and posterodistal corner apparently with finger- like projection (projection actually lateral view of posterior surface). Postgonite basal half robust, rounded ventrally and apically acute. Distiphallus: dorsal sclerite with apical ¼ extending beyond apex; acrophallus with round, ventrolateral projections present, and ventral acrophallus well developed.

Female Abdomen: unknown.

Type Material: Holotype (male, debu01077561): ECUADOR: Pichincha: Río Palenque Stn., 47 km S Santo Domingo, light, 1 May 1987, B. Brown (QCAZ).

Comments: Archiceroptera browni has a number of characters that are unique within

Archiceroptera, the most obvious being the absence of ocelli. Ocellar reduction occurs in several different limosinine clades (e.g., Aptilotella, Howickia), and is usually associated with loss of flight

(see Luk & Marshal 2014) and an associated shift to a terricolous habit; the well-developed wings of

A. browni are not consistent with that pattern. The collection of the holotype at a light trap might be reflective of nocturnal habits. Also unique in this species is the reduced size of the thoracic chaetotaxy compared with other Archiceroptera and the reduced epandrial process, which is

- 149 - relatively distinct in all other members of the larger Archiceroptera-group (with the exception of A. venezolana). Both of these are apparently apomorphic.

Etymology: This species is a patronym for Dr. Brian Brown, the collector of the type specimen and previous graduate student at Guelph.

Archiceroptera mahukani Papp 1977

Fig. 6.16–6.17

Description: See Papp 1977. In an effort to have comparable information for this species available here, with included morphological data from the female incorporated, a review of the morphology is given here based on the available female and the original description.

Length: 2.3–2.4 mm.

Head: Ocellar triangle with 15 small setae on ocellar triangle. Frons with 5 interfrontal seta and 5 inclinate orbital setulae. Eye:gena ratio = 4:3. Gena with 13 smaller setae.

Thorax: Dorsocentral setae in one strong prescutellar pair with 5–6 weaker dorsocentral setae anterior to prescutellar seta. Acrostichal setae in 6 rows between prescutellar dorsocentrals.

Scutellum with 6–7 large setulae near basal seta.

Legs: Fore femur with 5–6 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 10 stout setae extending from base to apical 3/4; apicoventral series of 5 setae on apical

1/4.; male, ventrally, with 20 robust setae in elongate basal cluster extending to near midlength. Mid tibia with 2–3 median anterodorsal and 2–3 median posterodorsal setae; predistal anterodorsal, predistal dorsal and distal posterodorsal setae strong, equal in length; male, ventrally with distal seta comb (not fully visible in image). Mid basitarsus with basal posteroventral seta stronger than apical setae. Hind tibia with 2–4 anterodorsal and 2–4 posterodorsal setae present (variable between sides on female); predistal dorsal seta sometimes present.

Wings: Costagial setae almost reach humeral crossvein. C2:C3 ratio = 5:3. Distance between r-m and dm-cu ~4.0–4.5× dm-cu.

Male Abdomen: Unknown (neither included in original description nor visible in images). “Male sternite 6 very small” (Papp 1977). - 150 -

Female Abdomen (Fig. 6.17): Tergite 7 narrowly rounded posterolaterally and closely appressed to base of tergite 8. Tergite 8 with lateral sclerites posterolaterally acutely projecting, narrowly rounded; medial sclerite transversely oval, poorly sclerotized. Sternite 7 broadly rounded. Sternite 8 medially fused into transverse band. Hypoproct not visible. Paired spermathecae with stems < 1/10th spermathecal length.

Type Material: Holotype (male): PARAGUAY: Puerto Presidente Stroessner, collected by lamplight on cut-down (burnt) clearing, at night, 5–6 Jan 1966, S. Mahunka (HNHM).

Specimens examined: BOLIVIA: La Paz: 1 female, Heath River Wildlife Research Centre,

12°40'S, 68°42'W, 3 km, 29 Apr–12 May 2007, S.M. Paiero

Comments: Papp (1977) discussed the differences between A. mahukani and A. venezolana, giving support of their differences using number of interfrontal setae, dorsocentral setae, extrascutellar setae, and tibial chaetotaxy. As he only had the holotype of A. mahukani and Richards’ (1963) limited description, he was unable to examine the extent of the variation within either species; the tibial chaetotaxy and interfrontal setae are not useful to separate the two species. The distributions of the two species appear to be allopatric, with A. venezolana occuring in the northern half of South

America and A. mahukani occuring farther to the south (Bolivia and Paraguay). The male internal genitalia remain unknown.

Archiceroptera venezolana (Richards) 1963

Ceroptera venozolana Richards, 1963: 232 [(printer’s error); both sexes]. Type locality:

Venezuela, Guanace, Estado Portuguesa. HT male (CASC).

Ceroptera venezolana. Richards, 1967: 7 [justified emendation, Neotropical catalog].

Archiceroptera venezolana. Papp, 1977: 382 [generic combination].

Figs. 6.5–6.7, and 6.18

Size: 1.4–3.6 mm.

Head: Ocelli reduced with 3–5 small setae on ocellar triangle. Frons with 4–6 interfrontal seta and

3–4 inclinate orbital setulae. Eye:gena ratio = 21:10. Gena with and 9–10 smaller setae.

- 151 -

Thorax: Dorsocentral setae in 3–5 pairs (including strong prescutellar); acrostichal setae in 8 rows between prescutellar dorsocentrals. Scutellum with 2–6 setulae near basal seta.

Legs: Fore femur with 4–6 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 9–11 stout setae extending from base to apical 3/4; apicoventral series of 6–7 setae on apical 1/4.; male, ventrally, with 15–20 robust setae in circular basal cluster, with another 7–10 weaker setae along posterior margin. Mid tibia with 2 median anterodorsal and 2–3 median posterodorsal setae; predistal anterodorsal, predistal dorsal and distal posterodorsal setae strong and equal in length; male ventral comb with 11–13 robust setae on apical 1/2. Mid basitarsus with basal posteroventral setae slightly larger than apical setae. Hind tibia with 4–6 anterodorsal, 5–7 posterodorsal, and 1–2 predistal dorsal setae present.

Wings: Costagial setae almost reaching humeral crossvein. C2:C3 ratio = 7:4. Distance between r- m and dm-cu ~4.0× dm-cu.

Male Abdomen (Figs. 6.5–6.6): Sternite 5 on disc adjacent to tab with numerous long setae; posteromedial tab short (1/6 sternite width), covered in fine setae; disc. Transverse portion of sternite 6 weakly arcuate, right lateral portion recurved, simple distally, left side with broadly acute medially projecting process. Cercus with basal ¼ triangular, with strong medial seta; apically narrow, acuminate. Surstylus (in lateral view) with broad, irregular anterior lobe; posteriorly with 8–

10 long setae. Postgonite basal 1/2 quadrate, with apical half acuminate. Distiphallus: dorsal sclerite with apical ¼ projecting beyond the acrophallus; acrophallus with dorsolateral and ventrolateral projections (both narrowly rounded).

Female Abdomen (Fig. 6.7): Tergite 7 posterolaterally broadly fused with tergite 8. Tergite 8 with lateral sclerites rounded posterolaterally; medial sclerite elongate, broadly rectangular. Sternite 7 with posterior margin narrowly rounded. Sternite 8 medially divided into pair of triangular sclerites.

Paired spermathecae with stems short (< 1/6 length of spermatheca) before fusing with common duct.

Comments: The male and female genitalia are described here for the first time.

Type Material Examined: Holotype (male) (images only): Venezuela: Estado Portuguesa, Guanare

(misspelled in Richards 1963 as “Guanace”), 10–13.IX.1957, B. Malkin (CASC). Non-type - 152 - material examined: COLOMBIA: Amazonas: 1 male, Leticia, 28 Feb 1974, V. Nealis, 2 females;

Rio Raposo, light, Jul 1964, V.H. Lee. COSTA RICA: Guanacaste: 1 female, Estacion Pitilla, 9 km S Sta. Cecilia, 700m, "II curso, L-N-330200.380200", May 1990; Heredia: 1 female, La Selva

Biological Station, 1* forest, blacklight, 28 Apr 1989, B.V. Brown; 1 male, La Selva Biological

Station, 3 km S Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 2–5 Jun 2001, S. Chatzimanolis; 2 males, Puerto Vieja, La Selva Biological Station, black light, 23 Apr 1989, B.V. Brown.

ECUADOR: Emeraldas: 1 male, La Chiquita, 11 km SE San Lorenzo, 5 m, dung, 9–10 Jun 1975,

S. Peck; 1 male, La Chiquita, dung, 7–8 Jun 1975, S. Peck; Napo: 1 male, Coca, Río Napo, 250 m,

May 1965, L.E. Peña; 3 males, 2 females, Tiputini Biodiversity Stn., 0°36'50"S", 76°9'1"W, May

2011, S.A. Marshall; 4 male, 6 females, Tiputini Biodiversity Stn., vicinity Yasuni National Park,

0°38'S, 76°10'W, pitfall trap (human dung), 14–19 Feb 1998, D.C. Darling; 2 males, 3 females,

Yasuní National Park, Yasuní Research Station, 0°38'S, 76°36'W, rainforest, malaise trap, 3–20 Nov

1998, Pape & Viklund; Pichincha: 1 female, Maquipucuna Biological Reserve, river trail,

0°7'34”N, 78°37'57”W, 1200 m, near stream, 26–28 Apr 2002; 3 males, Palenque, day 3 trap, 24–25

Mar 1976, S. Peck; 12 males 13 females, Rio Palenque, dung, 25 Feb 1979, S.A. Marshall; 1 male,

5 females, Rio Palenque, dung, 27 Feb 1979, S.A. Marshall; 3 males 9 females, Rio Palenque, dung trap, 22–23 Feb 1976, S. Peck; 7 males 1 female, Rio Palenque, dung trap, 25–26 Feb 1976, S.

Peck; 1 male, Rio Palenque, 22 Feb 1976, S. Peck; 19 males 17 females, Rio Palenque, 25–26 Jan

1976, S. Peck; 1 male, Rio Palenque, 26 Feb 1976, J. Glaser; 1 male, Rio Palenque Reserve Station,

Malaise trap, Feb 1983, M. Sharkey & L. Masner; 1 female, Rio Palenque Science Center, 47km S

Santo Domingo, 180m, 1* lowland rainforest, Malaise head, 29 Apr–5 May 1987, B. Brown & L.

Coote; 1 male, Rio Palenque Science Center, 47km S Santo Domingo, 180m, primary rainforest,

FIT, 2–4 May 1987, B. Brown & L. Coote; 1 male, Río Palenque Stn., 47 km S Santo Domingo, dung, 27 Feb 1979, S.A. Marshall; 2 males 1 female, Río Palenque Stn., 47 km S Santo Domingo, traps 3–5, day 1, 22–23 Feb 1976, S. Peck; 1 male, Rio Palenque Stn., 47km S Santo Domingo,

250m, rainforest, malaise-FIT, 5 May–25 Jul 1985, S. & J. Peck; 1 female, Rio Palenque Stn., 47km

S Santo Domingo, 250m, carrion trap, day 4, 27–28 May 1975, S. Peck; 6 males 1 female, Rio

Palenque Stn., 47km S Santo Domingo, 250m, dung, 17–25 Feb 1979, S.A. Marshall; 12 males 4 - 153 - females, Rio Palenque Stn., 47km S Santo Domingo, 250m, 17–25 Feb 1979, S.A. Marshall; 1 male

2 females, Santo Domingo, 4km SE, 500m, 3 forest dung pans, 8–11 Jun 1976, S. Peck; 6 males 8 females, Tinalandia, 1120m, wet lower montane rainforest, Malaise head ROM870006, 9–13 May

1987, L.D. Coote & B.V. Brown; 2 males, Tinalandia, 16 km SE Santo Domingo, 680 m, rainforest, malaise-FIT, 4 May–25 Jul 1985, S. & J. Peck. FRENCH GUIANA: St. Laurent du Maroni: 1 male, Mitaraka, MIT-A-RBF1, 2°14'11”N, 54°27’7”W, 306m, tropical wet forest (bas fond), yellow pan traps, 4–8 Mar 2015, M. Pollet; 1 male 1 female, Mitaraka, MIT-A-SL, 2°14'18”N, 54°27’8”W,

352m, tropical moist forest (slope), blue pan traps, 3–8 Mar 2015, M. Pollet; 3 males, Mitaraka,

MIT-A-TOP, 2°14'20”N, 54°27'11”W, 361m, tropical moist forest (plateau), blue pan traps, 3–8

Mar 2015, M. Pollet; 1 male, Mitaraka, MIT-C-RBF2, 2°14’3”N, 54°26'53”W, 299m, tropical wet forest (bas fond), yellow pan traps, 6–10 Mar 2015, M. Pollet; 5 males, Mitaraka, MIT-C-SL,

2°14’8”N, 54°26'42”W, 373m, tropical moist forest (slope), blue pan traps, 2–8 Mar 2015, M.

Pollet; 4 males 6 females, Mitaraka, MIT-C-TOP, 2°13'59”N, 54°26'38”W (MHNM), 433m, tropical moist forest (plateau), blue pan traps, 2–8 Mar 2015, M. Pollet; 1 male 4 females, Mitaraka,

MIT-C-TOP, 2°13'59”N, 54°26'38”W, 433m, tropical moist forest (plateau), white pan traps, 2–8

Mar 2015, M. Pollet (MHNM); 3 females, Mitaraka, MIT-C-TOP, 2°13'59”N, 54°26'38”W, 433m, tropical moist forest (plateau), yellow pan traps, 2–8 Mar 2015, M. Pollet; 1 female, Mitaraka, MIT-

DZ2, 2°14’3”N, 54°27’2”W, 296m, tropical moist forest (bas fond), blue pan traps, 2–10 Mar 2015,

M. Pollet; 2 males, Mitaraka, MIT-DZ-RBF1, 2°14’4”N, 54°27’2”W, 270m, tropical wet forest (bas fond), blue pan traps, 2–10 Mar 2015, M. Pollet; 1 female, Mitaraka, MIT-DZ-RBF2, 2°13'59”N,

54°27’0”W, 283m, tropical wet forest (bas fond), blue pan traps, 5–10 Mar 2015, M. Pollet; 1 female, Mitaraka, near base camp & along trails, tropical moist forest, SLAM trap, 1 Mar 2015,

Touroult & Poirier; 1 male, Mitaraka, near base camp & along trails, partially opened areas near base camp & DZ, & rock savannah, SLAM trap, 12–20 Aug 2015, P.-H. Dalens; 1 male, Mitaraka, near base camp & along trails, tropical moist forest, SLAM trap, 14 Mar 2015, Touroult & Poirier; 1 male, Mitaraka, near base camp & along trails, tropical moist forest, SLAM trap, 1–6 Mar 2015,

Touroult & Poirier. GUYANA: 1 male 1 female, Kabocalli, Iwokrama Forest Reserve, 60 m, FIT,

3–5 Jun 2001, Brooks & Falin; 1 female, Kartabo, Bartica Dist., trap lante, 9 Nov 1920. - 154 -

JAMAICA: 1 male, St. Thomas, Portland Gap, 3500ft, 20 Jul–1 Aug 1974, S. Peck. PANAMA: 1 female, Canal Zone, Barro Colorado, 20 Jul 1924, N. Banks. PERU: Loreto: 2 males, Campamento

San Jacinto, 175–215 m, in primary forest, FIT, #54, 8 Jul 1993, R. Leschen; 2 males 2 females,

Campamento San Jacinto, 175–215 m, #66, FIT, Qd.24, 9 Jul 1993, R. Leschen; 2 females,

Campamento San Jacinto, 175–215 m, FIT, 7 Jul 1993, R. Leschen; 2 females, Campamento San

Jacinto, 175–215 m, FIT #10, 3 Jul 1993, R. Leschen; 3 females, Campamento San Jacinto, 175–

215 m, FIT #31, 5 Jul 1993, R. Leschen; 1 maleCampamento San Jacinto, 175–215 m, FIT, #67, 9

Jul 1993, R. Leschen; 1 male 1 female, Campamento San Jacinto, FIT, #87, 12 Jul 1993, R.

Leschen; 1 male, Teniente López, riverine forest, FIT, #199, 24 Jul 1993, R. Leschen; Madre de

Dios: 1 male 1 female, Zona Reserva Manu, Pakitza, 11°57'S, 71°17'W, 400 m, malaise trap 3, 18–

23 Feb 1992, B. Brown & D. Feener; 1 male, Rio Tambopata, Explorers Inn, primary rainforest, 17

Jan 1981, Gärdenfors, Hall & Samuelsson. VENEZUELA: Bolivar: 1 male, 2 females, 10km S El

Dorado, 200m, 17 Jul–7 Aug 1986, B.D. Gill; 1 female, 125km S El Dorado, 1100m, 18 Jul–7 Aug

1986, B.D. Gill; 1 male, 22km S El Dorado, lowland rainforest, FIT, 25 Jun–12 Jul 1987, S. & J.

Peck; 9 males 3 females, 33km S El Dorado, 220m, 2–7 Aug 1986, B.D. Gill; 7 males, 5 females, km40 Sta. Elena Icabaru Rd., 220m, 4–6 Aug 1986, B.D. Gill; 31 males 14 females, km40 Sta.

Elena Icabaru Road, 1000m, 4–6 Aug 1986, B.D. Gill; 6 males 14 females, km40 Sta. Elena Icabaru

Road, 100m, 4 Aug 1986, B.D. Gill; 4 females, Quebrada de Jaspe, 19–20 Jul 1986, B. Gill.

6.7.3 Archiceroptera ternum-species group

Species from this group are known from the USA south to Argentina. This group includes all Archiceroptera species outside the two basal clades (A. addenda and A. mahukani species groups). All 23 species in this group are here newly described. Members of this group can be recognized by the combination of a distinct dorsal basal seta, female tergite 7 fused laterally with tergite 8, and female sternite 8 reduced to a pair of lateral triangular sclerites that narrowly meet medially. The A. brevivilla species subgroup is a derived species group within the A. ternum group that has a reduced mid tibial chaetotaxy.

- 155 -

Archiceroptera ternum species group description:

Head: Frons with 4–7 interfrontal seta. Gena with 2 strong anterior setae and additional smaller posterior setae.

Thorax: Prescutellar dorsocentral seta strong. Acrostichal setulae in 4-10 rows between prescutellar dorsocentrals. Anterior katepisternal seta absent or present. Scutellum with 1 setulae near basal seta

(A. bilobata) or without setulae (all other species).

Legs: Mid tibia with dorsal basal seta, 1–3 median anterodorsal, and 1–4 median posterodorsal setae; strong predistal anterodorsal present; predistal dorsal present or absent; distal posterodorsal setae present or absent; male with or without ventral comb; midventral seta present in females and some males. Mid basitarsus with basal posteroventral setae variable in length compared with distal setae. Hind tibia dorsally variable (usually simple, but 1–7 dorsal setae and/or preapical seta present in some species).

Wings: Costa with 2 strong costagial setae; dorsal seta usually longer. Cell dm with small CuA1 stub vein present or absent.

Male abdomen: Sternite 4 posteromedially usually entire (emarginate in some species). Sternite 5 variable (usually posterior margin medially emarginate). Transverse portion of sternite 6 nearly straight to strongly arcuate. Surstylus variable. Cercus variable (triangular to basally ovoid with apical portion narrowed). Distiphallus with acrophallus largely membranous.

Female Abdomen: Tergite 7 posteromedially variable; posterolaterally fused to anterolateral corner of tergite 8. Tergite 8 tripartite, with medial sclerite well sclerotized. Epiproct medially desclerotized into paired sclerites; with scattered setulae. Cercus variable (often tear-drop shaped) with flattened apical seta with small, flattened preapical seta appressed to base of apical seta.

Sternite 7 posteriorly usually rounded (emarginate in some species). Sternite 8 medially divided into pair of elongate triangular sclerites that narrowly meet medially. Spermatheca shape variable, with swelling at duct junction; paired spermathecae with stems extremely short or obsolete.

Archiceroptera adamas Paiero & Marshall, n. sp.

Figs. 6.19–6.20 - 156 -

Size: 1.5–2.1 mm.

Head: Ocellar triangle with 7–9 setae. Frons with 4 interfrontal seta and 2–3 inclinate orbital setulae. Eye:gena ratio = 1:1. Gena with 11–12 smaller setae.

Thorax: Acrostichal setulae in 6 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with posterodorsal and posteroventral setae indistinct. Mid femur anteriorly with series of 12 stout setae extending from base to apical 3/4; apicoventral series of 4–5 setae on apical

1/4.; male, ventrally, with 10–11 robust setae in elongate basal cluster that extends to basal 1/3. Mid tibia with dorsal basal seta, 2 median anterodorsal and 2 median posterodorsal; with strong predistal dorsal seta and strong distal posterodorsal seta; male, ventrally, with 10–11 robust setae on apical

3/4; midventral seta present only in female. Mid basitarsus with basal posteroventral seta slightly larger than apical setae. Hind tibia with no distinct setae.

Wings: Costagial setae not reaching humeral crossvein. C2:C3 ratio = 5:4. Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm with small CuA1 stub vein present.

Male Abdomen (Fig. 6.19): Sternite 4 posteriorly entire. Sternite 5 posteromedially with diamond- shaped emargination that extends to basal ¼ of sternite; emargination flanked by acute posteromedially-directed processes (which may have small apical crenulations present). Transverse portion of sternite 6 narrowly, but strongly, arcuate; right lateral portion recurved and fused with quadrate sclerite. Synsternite 6+7 with narrowly rounded medially projecting process, with triangular sclerite immediately distal to tip. Cercus basal 1/3 triangular with strong distinct seta; apical 2/3 narrowed slightly and medially curved at apical 1/3. Surstylus (in lateral view) with small anterior lobe with short, rounded process; posterior portion with 14–16 setae along posterior surface. Postgonite basal ½ quadrate with rounded ‘heel’; apical 1/2 acuminate with upper surface flat. Distiphallus: dorsal sclerite with apical ¼ extending beyond apex; acrophallus with broadly rounded dorsolateral flaps and with large, triangular ventral portion.

Female Abdomen (Fig. 6.20): Tergite 7 posteromedially broadly sinuate; posterolaterally narrowly fused to tergite 8. Tergite 8 with lateral sclerites broadly rounded; medial sclerite broadly triangular.

Epiproct with lateral sclerites anterolaterally rounded. Cercus basally quadrate; flattened apical setae - 157 - with smaller, flattened preapical seta appressed to base of apical seta. Sternite 7 broadly rounded.

Spermatheca ovoid, slightly flattened; sclerotized portion of duct ~1/3 length of spermatheca.

Type Material: Holotype (male, debu00385853): Maripasoula, Mitaraka, MIT-C-TOP,

2°13'59"N, 54°26'38"W, 433m, tropical moist forest (plateau), blue pan traps, 2–8 Mar 2015, M.

Pollet (MHNM). Paratypes: FRENCH GUIANA: St. Laurent du Maroni: 1 male, Maripasoula,

Mitaraka, MIT-A-TOP, 2°14'20"N, 54°27'11"W, 361m, tropical moist forest (plateau), blue pan traps, 3–8 Mar 2015, M. Pollet (MHNM); 1 male, Maripasoula, Mitaraka, MIT-C-SL, 2°14'08"N,

54°26'42"W, 373m, tropical moist forest (slope), yellow pan traps, 2–8 Mar 2015, M. Pollet.

GUYANA: Potaro-Siparuni: 1 male, 1 female, Mount Wokomung, 5°7'53"N, 59°48'31"W, 698m,

1° forest, pitfall trap (human dung), 21–26 Oct 2004, B. Hubley; Rupununi: 1 male, Kurupukari,

Essequibo River, 200', 1° forest, dung traps, 9 Oct 1990, B. Hubley (ROME); (distr. unknown): 1 male, Kabocalli, Iwokrama Forest Reserve, 60 m, FIT, 3–5 Jun 2001, Brooks & Falin

Comments: The enlarged sclerite at the apex of the medially projecting process on the left side of synsternite 6+7 has some resemblance to the sclerites found in the A. addenda group. One paratype has a fungal growth (Laboulbeniales) in the middle of sternite 5 (Fig. 6.19 A–B).

Etymology: The species epithet is the Latin for “diamond” in reference to the diamond shape emargination of the male sternite 5

Archiceroptera barberi Paiero & Marshall, n. sp.

Figs. 6.21–6.22

Size: 1.3–2.8 mm.

Head: Ocelli well developed with 3–5 setae. Frons with 3–5 interfrontal setae; 3–5 inclinate orbital setulae. Eye:gena ratio = 1:1. Gena with 12–17 smaller setae.

Thorax: Acrostichal setulae in 6 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 4–5 posterodorsal and 5–7 posteroventral setae. Mid femur anteriorly with series of 8–11 stout setae extending from base to apical 3/4; apicoventral series of 4–6 setae on apical 1/4.; male, ventrally, with 15–17 robust setae, with 7–13 in cluster on basal 1/3 and remaining - 158 -

4–6 extending distally along posterior margin. Mid tibia with 2 median anterodorsal and 2 median posterodorsal (basal seta sometimes weak or absent) setae; weak predistal dorsal and weak distal posterodorsal setae present; male ventral comb with 12–15 robust setae on apical 1/3; midventral seta present only in females. Mid basitarsus with basal posteroventral setae slightly larger than apical setae. Hind tibia with 1–2 weak predistal dorsal setae present on some individuals.

Wings: Costagial setae with longer seta extending to humeral crossvein. C2:C3 ratio = 2:1. Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm usually without CuA1 stub vein.

Male Abdomen (Fig. 6.21): Sternite 4 posteriorly entire. Sternite 5 posteromedially with several small fine setae on each side of medial desclerotized area; desclerotized area elongate oval, extending to basal ½ of sternite (extent of desclerotization somewhat variable and not always apparent in undissected specimens). Transverse portion of sternite 6 straight to weakly arcuate, right lateral portion recurved and fused with quadrate sclerite. Synsternite 6+7 with acute, medially projecting process on left. Cercus with basal 1/3 ovoid, with long, strong seta; apical 2/3 elongate, flattened (length = ~3× width). Surstylus (in lateral view) with anterior lobe weakly developed with a triangular distal portion and short anteriorly-projecting finger-like process. Postgonite with basal

½ quadrate, apical ½ acuminate, robust. Distiphallus: dorsal sclerite not projecting beyond apex; acrophallus dorsolaterally with broadly rounded process recurved dorsally.

Female Abdomen (Fig. 6.22): Tergite 7 posteromedially produced with weak medial emargination; posterolaterally narrowly fused with tergite 8. Tergite 8 with lateral sclerites posterolaterally narrowly rounded; medial sclerite elongate, triangular. Epiproct with lateral sclerites oval. Cercus ovoid, with large, apical flattened setae and smaller flattened preapical seta appressed to base of apical seta. Sternite 7 posteriorly broadly rounded. Spermatheca kidney-shaped; sclerotized portion of duct ~1/5 length of spermatheca.

Type Material: Holotype (male, debu01081889) and 36 Paratypes (23 males, 14 females):

PANAMA: Chiriquí: Hartmann's Finca, 15 km NW Hato de Volcán, 1200 m, dung trap, 20–31

May 1977, S. Peck. Additional Paratypes: BELIZE: 4 males, 3 females, Mtn. Pine Ridge, Hidden

Valley Inst., 2500ft, grass-pine dung traps, 10–15 Jan 1991, S.A. Marshall. COLOMBIA:

Cundinamarca: 1 male, Finca Bella Vista, near Sasaima, 13 Apr 1965, P.R. Craig. COSTA - 159 -

RICA: Alajuela: 1 male 1 female, Florencia Forest, dung tp., 28 Feb 1980, H. Howden; 2 males 5 females, Volcán Tenorio, N slope near Bijagua Biological Station, 700 m, rainforest, RET over Atta mound, 16–20 Jun 2000, S.A. Marshall; 1 female, Volcán Tenorio, N slope near Bijagua Biological

Station, 700 m, rainforest, sweeping trail, 17 Jun 2000, S.A. Marshall; 1 male, Volcán Tenorio, N slope near Bijagua Biological Station, 700 m, pan traps in tree fall, 18 Jun 2000, Buck & Marshall;

1 male 1 female, Volcán Tenorio, N slope near Bijagua Biological Station, 700 m, sweeping trail,

18 Jun 2000, S.A. Marshall; Cartago: 1 male, Tapantí National Park, Ranger Stn., 1200 m, human dung, 11 Oct 1999, M. Buck; 1 male, Tapantí National Park, Ranger Stn., 1200 m, human dung, 12

Oct 1999, M. Buck; 8 males 7 females, Tapantí National Park, Ranger Stn., 1200 m, human dung, hand & traps, 9–12 Oct 1999, Buck & Marshall; 1 male 2 females, Turrialba Catie, 600 m, 26 Feb

1980, H. & A. Howden; 1 male, Turrialba Catie, 600 m, 28 Feb 1980, H. & A. Howden; 2 females,

Parque Nacional Tapanti, 9°43'21”N, 83°46'30”W, 1600m, 20–27 Jan 2013, ZADBI-505 #106710,

Malaise trap, ZADBI; Guanacaste: 1 male, Guanacaste Cons. Area, Pitilla Field Station, Malaise,

29 Jan 1996, J. Noyes; 4 males, 5 females, Guanacaste Cons. Area, Ricon de la Vieja, Las Pailas,

1400 m, Clusea rosea forest litter, 18–20 Feb 1996, R. Anderson; Heredia: 1 female, 10km W

Puerto Viejo, La Selva Verde, 2–4 Mar 1991, H. & A. Howden; 1 male, La Selva, 10°25'48"N,

84°1'12"W, Malaise, 8–15 May 1989, B. Brown & D. Feener; 4 males 19 females, La Selva

Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 11–14 Jun 2001, S.

Chatzimanolis (INBC, DEBU); 2 males, 7 females, La Selva Biological Station, 3 km S Puerto

Viejo, 10°26'N, 84°1'W, 80 m, FIT, 14–17 Jun 2001, S. Chatzimanolis; 4 females, La Selva

Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 18–20 Jun 2001, S.

Chatzimanolis; 2 males, 3 females, La Selva Biological Station, 3 km S Puerto Viejo, 10°26'N,

84°1'W, 80 m, FIT, 20–23 Jun 2001, S. Chatzimanolis; 2 males, La Selva Biological Station, 3 km S

Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 2–5 Jun 2001, S. Chatzimanolis (INBC); 1 male, La

Selva Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 5–8 Jun 2001, S.

Chatzimanolis; 1 male, 1 female, La Selva Biological Station, 3 km S Puerto Viejo, 10°26'N,

84°1'W, 80 m, FIT, 8–11 Jun 2001, S. Chatzimanolis; Puntarenas: 1 female, Coto Brus, Z.P. Las

Tablas, Estacion Biologica Las Alturas, 8°57'7”N, 82°50'4”W, 1500–1600 m, 10–17 Dec 2012, - 160 -

ZADBI-326 #105646, Malaise trap, ZADBI; 1 male, 4 females, Las Alturas, 8°57'N, 82°58'W, 1600 m, malaise trap, 11–14 Aug 1995, S.A. Marshall; 1 male, 1 female, Los Alturas, 1600 m, ground

Eciton raid, 15 Aug 1995, S.A. Marshall; ; 1 male, Osa Peninsula, Rincón, 2.5 km S, 8°42'1"N,

83°30'50"W, ~50 m, secondary forest, dung pans, 11 Aug 2001, M. Buck; San José: 1 male, San

Carlos, Riosparaíso Reserve, trail to Río Blanco, 9°34'15"N, 84°7'29"W, 420 m, 22 Feb 2006, S.M.

Paiero; 1 male, Zurqui de Moravia, 10°2'58”N, 84°0'57”W, 1600m, 24–30 May 2013, Tower path,

ZADBI-785, #106841, Malaise trap #1, ZADBI; 2 males, Zurqui de Moravia, 10°2'58”N,

84°0'57”W, 1600m, 11–18 Oct 2013, Tower path, ZADBI-1280 #107959, Malaise trap #1, ZADBI

(INBC); 1 male, Zurqui de Moravia, 10°2'58”N, 84°0'57”W, 1600m, 12–19 Apr 2013, Creek 2 north, Malaise trap #2, ZADBI-711 #106716, ZADBI (INBC). ECUADOR: Emeraldas: 1 female,

La Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, 10–11 Jun 1975, S. Peck (QCAZ); 1 male, La

Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, day 3, 8–9 Jun 1975 (QCAZ), S. Peck; 3 males,

La Chiquita, dung, 7–8 Jun 1975, S. Peck; Pichincha: 1 male, Maquipucuna Biological Reserve, river trail, 0°7'34"N, 78°37'57"W, 1200 m, near stream, pans/ dung, 26–28 Apr 2002, S.A.

Marshall; 1 male, Rio Palenque, 25–26 Jan 1976, S. Peck; 1 male, Tinalandia, 1120m, wet lower montane rainforest, Malaise head, 9–13 May 1987, L.D. Coote & B.V. Brown (QCAZ).

GUATEMALA: Baja Verapaz: 2 males, Biotopo Quetzal, 15°12'49"N, 90°13'6"W, 8 m, 1690 m, cloud forest, malaise trap, 7–10 May 2009; 3 males, 4 females, Purulhá, 7 km NE, 1500 m, FIT, 20

May–8 Jun 1991, B.D. Gill; 7 males, 17 females, Purulhá, 7.4 km S, 1650 m, FIT, 2 Jul 1993, Ashe

& Brooks; 9 males, 13 females, Purulhá, 7.4 km S, 1650 m, FIT, #189, 2–3 Jul 1993, Ashe &

Brooks (UVGC, DEBU); 1 male, Purulhá, 8 km S, dung trap, 25 May 1991, H. & A. Howden; 1 male, Purulhá, 8 km S, FIT, 23–25 May 1991, H. Howden; 2 females, Purulhá, 8 km S, FIT, 27 May

1991, H. & A. Howden; 2 males, Purulhá, 8 km S, FIT, 29 May 1991, H. & A. Howden;

Guatemala: 3 females, Santa Catarina Pinula, dung traps, 11–13 Jun 1991, B.D. Gill; Zacapa: 7 males, 2 females, La Unión, 3.5 km SE, 1500 m, FIT, 23–25 Jun 1993, Ashe & Brooks; 3 males, 8 females, La Unión, 3.5 km SE, 1500 m, FIT, 25–27 Jun 1993, Ashe & Brooks. HONDURAS:

Olancho: 1 male, La Muralla National Park, 15°5'49"N, 86°44'17"W, 1450 m, FIT, 4–7 Jul 2002,

Smith & Ocampo. MEXICO: Chiapas: 1 female, Bonampok Rd, 100km S Palenque, 230m, - 161 - rainforest, FIT, 8–24 Jul 1983, B.D. Gill; 1 male, 1 female, Laguna Belgica, 16 km NW

Ocozocoaulta, 970m, FIT, 13 Jun 1990, H. & A. Howden & Gill (UNAM); 1 male, Lagunas de

Montebello Parque Nacional, La Eucantada, 4900ft, oak/pine/liquidambar, human dung, 21–24 Aug

1971, A. Newton; 1 male, 1 female, Ocozocoautla, 11mi NW, 3400ft, oak-evergreen forest, human dung, 19–25 Aug 1971, A. Newton (FMNH); 1 female, San Cristóbal de las Casas, dung traps, 26–

28 May 1990, B. Gill; 1 male, San Cristóbal de las Casas, 7087 ft, 12 Jun 1969, B.V. Peterson;

Guerrero: 3 males, 1 female, Mazatlan, 4mi W, 4800ft, oak, tropical deciduous forest, human dung, 30 Aug–5 Sep 1971, A. Newton; Hidalgo: 3 males, 1 female, Tenango de Doria, 7mi SW,

7000ft, cloud oak forest, human dung, 2–6 Jul 1971, A. Newton; 9 males, 7 females, Tlanchinol,

2.5mi N, 5100ft, cloud forest, dung, 6–11 Jul 1973, A. Newton (FMNH, UNAM); Mexico: 6 males,

4 females, Temascalteper, 3mi NE, 6300ft, oak-madrono-pine, human dung, 2–7 Sep 1971, A.

Newton; 4 males, 6 females, Tenancingo, 1mi NE, 7100ft, oak-pine-madrono, human dung, 31

Aug–6 Sep 1971, A. Newton; Morelos: 1 female, Tres cumbres, 4mi W, km 6, 8900ft, oak, human dung, 29 Aug–4 Sep 1971, A. Newton; Oaxaca: 1 female, El Camaron, 9mi E, 4300ft, human dung,

23 Aug–6 Sep 1973, A. Newton; 1 male, Ixtlan de Juarez, 14.2 mi S, 7600ft, oak woodland, dung,

10–18 Aug 1973, A. Newton; 1 female, Valle Nacional, 12mi S, 3200 ft, tropical montane forest, shrimp carrion, 22–31 Jul 1971, A. Newton; Puebla: 2 males, Huanchinango, 5mi W, 6000ft, hardwood-pine, human dung, 3–7 Jul 1971, A. Newton; 6 males, 8 females, Xicotepec de Juarez,

3mi S, 4000ft, tropical evergreen forest, human dung, 3–8 Jul 1971, A. Newton (FMNH, UNAM);

Queretaro: 2 females, Landa de Matamoros, 19mi E, 5400ft, pine madrono, human dung, 21–27

Jun 1971, A. Newton; Sinaloa: 2 males, 2 females, Concordia, 31mi NE, 4700ft, tropical deciduous forest, dung trap, Sep 1973, A. Newton; Veracruz: 8 males, 6 females, Huatusco, 4mi N, 4100ft, cloud forest, dung, 11–16 Jul 1971, A. Newton; 1 female, Sumidero, FIT, 7–9 Jun 1990, B.D. Gill;

12 males, 4 females, Teocelo, 10mi SW, 4400ft, oak, wet, human dung, 11 Jul 1971, A. Newton.

PANAMA: Chiriquí: 4 males, 5 females, Hartmann Finca, 15 km NW Hato de Volcán, dung trap,

20–25 May 1977, S. Peck; 2 males, 2 females, Hartmann's Finca, 1550 m, dung trap, 31 May 1977,

S. Peck; 5 males, 2 females, Hartmann's Finca, 1700 m, 28 Jun–3 Jul 1981, B. Gill; 1 male,

Hartman's Finca, 28 Jun–3 Jul 1981, B.D. Gill; 21 males, 14 females, Lagunas, 5km SW Hato del - 162 -

Volcan, 1360m, dung, 22–26 May 1977, S. Peck; 5 males, 7 females, Lagunas, 5km SW Hato del

Volcan, 1360m, dung, 22–27 May 1977, S. Peck; 1 male, Lagunas, W Hato del Volcan, carrion trap#2, 22–29 May 1977, S. Peck. TRINIDAD & TOBAGO: Tobago: 2 male, 2 females,

Charlotteville, Man-O-War Bay cottages, littoral rainforest, UV light, 26–30 Jun 1993, S. & J. Peck.

VENEZUELA: Aragua: 1 male, Rancho Grande Biological Station, 10°21'N, 67°41'W, 1200 m, flight intercept trap, 14 May 1998, Ashe, Brooks & Hanley; 1 female, Tiara, Env, 1250m, 12 Apr

1994, L. Masner; Lara: 1 male, 1 female, Yacambu, 1200m, cloud forest, 7 May 1981, H.K.

Townes (MIZA); 1 male, 2 females, Yacambu, 1200m, 10 May 1981, H.K. Townes.

Comments: There appears to be some subtle allometric variation between larger and smaller A. barberi males, but this appears to be variation between individuals and not suggestive of cryptic species.

Etymology: The species name honours dipterist and University of Guelph alumni Kevin Barber, and has been a manuscript name (originally Rudolfina barberi) since 1982.

Archiceroptera basilia Paiero & Marshall, n. sp.

Figs. 6.23-6.24

Size: 2.8–3.1 mm.

Head: Ocellar triangle with 3–5 small setae. Frons with 3–5 interfrontal seta and 3–5 inclinate orbital setulae. Eye:gena ratio = 1.8:1. Gena with 13–14 smaller setae.

Thorax: Acrostichal setulae in 6 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 4–5 posterodorsal and 5–6 posteroventral setae. Mid femur anteriorly with series of 11–13 stout setae extending from base to apical 3/4; apicoventral series of 4–7 setae on apical 1/4; male, ventrally, with linear series of 5–6 weakly strengthened setae on basal 1/3. Mid tibia with 1 median anterodorsal and 1 median posterodorsal setae; strong predistal dorsal and weak distal posterodorsal setae present; male ventral comb with 9–10 weak setae on apical 1/2; midventral seta present in both sexes. Mid basitarsus without enlarged distinct posterobasal setae.

Hind tibia simple. - 163 -

Wings: Costagial setae with longer seta reaching humeral vein. C2:C3 ratio = 2:1. Distance between r-m and dm-cu ~5.0× dm-cu. Cell dm without CuA1 stub vein.

Male Abdomen (Fig. 6.23): Sternite 4 posterior margin entire. Sternite 5 posteromedially with pair of broadly rounded, weakly clavate processes (length of process = ~1/4 length of sternite); margin lateral to processes broadly emarginate. Transverse portion of sternite 6 broadly arcuate; right lateral portion recurved and fused with quadrate sclerite. Synsternite 6+7 with trilobed medially projecting process. Cercus with basal 2/3 quadrate, narrowed on apical 1/3 to 1/3 basal width; apically rounded. Surstylus (in lateral view) with anterior lobe triangular, broadly rounded anteriorly; posteriorly with 12–14 long setae. Postgonite with basal ½ quadrate, concave below; apically acuminate, anteriorly curved. Distiphallus: dorsal sclerite with apical 1/3 extending beyond apex; acrophallus dorsolaterally with narrow, basally-angled processes with narrowly rounded tips.

Female Abdomen (Fig. 6.24): Tergite 7 medially desclerotized, posteriorly with broad sinuate emargination (~1/3 tergite width); posterolaterally narrowly fused with tergite 8. Tergite 8 with lateral sclerites posterolaterally acutely angled; medial sclerite narrower than anterior portion of epiproct. Epiproct with lateral sclerites triangular; anteriorly extended into elongate stem ( = length of broader, posterior portion). Cercus basally rectangular, weakly narrowed near midlength..

Sternite 7 broadly rounded. Spermathecae ovoid; sclerotized portion of ducts ~1/3 length of spermathecae.

Type Material: Holotype (male, debu00140620) and paratype (1 female): ECUADOR: Napo:

Cosanga, 2.5 km W, 0°35'24"S, 77°53'19"W, 2150 m, dung/pans, 5–7 Nov 1999, S.A. Marshall

(QCAZ). Additional Paratypes: ECUADOR: Napo: 1 female, Maquipucuna Biological Reserve, river trail, 0°7'34"S, 78°37'57"W, 1200 m, pans/dung, 26–28 Apr 2002, S.A. Marshall; 8 females,

Baeza 17km NE, 1400m, 3–6 Mar 1976, dung trap, S. Peck; 3 females, El Chaco, 2000m, 15–23

Feb 1983, Malaise trap, Masner & Sharkey (QCAZ, DEBU). PERU: Cusco: 1 female, Wayquecha

Biological Station, ~9km NE Challabamba, 13°10'20"S, 71°35'0"W, 2600–2700m, Trecha Ferdiz, dung pans, 3 Dec 2011, S.A. Marshall (MUSM).

Comments: The trilobed process on synsternite 6+7 is unique within Archiceroptera, although several other species do have variously modified processes. - 164 -

Etymology: The species epithet is Latin for royalty or crown, in reference to the crown-like appearance of the male sternite 5.

Archiceroptera bilobata Paiero & Marshall, n. sp.

Figs. 6.25–6.26

Size: 2.0–2.8 mm.

Head: Ocellar triangle with 5–6 small setae. Frons with 4 interfrontal seta and 3–4 inclinate orbital setulae. Eye:gena ratio = 1:1. Gena with 11–12 smaller setae.

Thorax: Dorsocentral seta in 1 prescutellar pair. Acrostichal setulae in 6 rows between prescutellar dorsocentrals. Scutellum with 1 setula near basal seta. Anterior katepisternal seta present, weaker than posterior seta.

Legs: Fore femur with 5–7 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 11–12 stout setae extending from base to apical 3/4; apicoventral series of 5 setae on apical

1/4.; male, ventrally, with 10–13 robust setae in cluster on basal 1/3, with 3–4 additional setae along posterior margin. Mid tibia with 1 median anterodorsal and 1 median posterodorsal setae; predistal dorsal and distal posterior setae present; male ventral comb with 14–15 robust setae on apical 2/3; midventral seta present in female. Mid basitarsus with basal posteroventral setae slightly longer than distal setae. Hind tibia with 1–2 distal dorsal setae usually present.

Wings: Costa with strionger costagial seta reaching humeral cross vein. C2:C3 ratio = 2:1. Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm without CuA1 stub vein (corner of cell rounded).

Male Abdomen (Fig. 6.25): Sternite 4 posteriorly entire. Sternite 5 posteromedially with broad triangular emargination extending ½ length and ½ width of sternite. Transverse portion of sternite 6 broadly arcuate, with right lateral portion expanded as a quadrate sclerite with posteromedially recurved process. Synsternite 6+7 with broadly rounded process with irregular sclerite distal to tip.

Cercus with basal 2/3 triangular-ovoid with strong seta, distally projecting as acute process; short, no more than ½ length of surstylus. Surstylus (in lateral view) bilobed with small finger-like process anterobasally; anterior lobe setose, ~2/3 length of posterior lobe; posterior lobe with 15–20 setae on posterior surface. Postgonite with basal ½ quadrate, narrowing into acute distal half. Distiphallus: - 165 - distiphallus with apical ¼ extending beyond apex; acrophallus with elongate, acute lateral projection near midlength.

Female Abdomen (Fig. 6.26): Tergite 7 medialy desclerotized, with small emargination at posterior end of desclerotized area; posterolaterally broadly fused to tergite 8. Tergite 8 with lateral sclerites with posterolateral corner narrowly rounded; medial sclerite elongate, narrow rectangular.

Epiproct with lateral sclerites extended anteriorly (~1/2 length of broader posterior portion) into stem. Cercus ovoid, with large apical flattened setae with small flattened preapical seta appressed at base. Sternite 7 posteriorly broadly rounded. Spermatheca barrel-shaped with small invaginations on ends; sclerotized portion of duct ~1/2 length of spermatheca.

Type Material: Holotype (male, debu00378707, QCAZ) and 149 Paratypes (92 males, 57 females, QCAZ & DEBU): ECUADOR: Pinchincha: Alluriquin, 23km E, Chiriboyo Ret., 4600ft, dung, 19–27 Jun 1975, S. Peck. Additional Paratypes: ECUADOR: Pichincha: 9 males, 11 females, Allurquin, 28km E, Chiribaga Rd., 5200ft, moss forest, carrion trap, 19–27 Jun 1975, S.

Peck; 2 males, Tinalandia, 16 km SE Santo Domingo, 680 m, pasture, dung, 28–29 Jun 1975, S.

Peck.

Comments: This species is unique within the A. ternum group for its small acrophallus, additional scutellar seta, distinctly bilobate surstylus, and reduced male cercus.

Etymology: The name refers to the distinctly bilobate surstylus.

Archiceroptera bisetosus Paiero & Marshall, n. sp.

Figs. 6.27-6.28

Size: 2.0–2.3 mm.

Head: Ocellar triangle with 3 setae. Frons with 4 interfrontal seta and 3–4 inclinate orbital setulae.

Eye:gena ratio = 1:1. Gena with 8–11 smaller setae.

Thorax: Acrostichal setulae in 8 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta not apparent, indistinguishable from surrounding setulae.

Legs: Fore femur with 4–6 posterodorsal and 4–6 posteroventral setae. Mid femur anteriorly with series of 11 stout setae extending from base to apical 3/4; apicoventral series of 4 setae on apical - 166 -

1/4.; male, ventrally, with 1–2 robust setae present near base. Mid tibia with dorsal seta more anterodorsal than dorsal; 1–3 median anterodorsal (1 strong, basally with 1–2 additional, weak setae sometimes present) and 1 median posterodorsal seta; weak predistal dorsal and weak distal posterodorsal setae present; male without ventral comb; midventral seta present in both sexes. Hind tibia usually with uniformly small setulae only (1 weak preapical seta sometimes present).

Wings: Costa with longer costagial setae not reaching humeral crossvein. C2:C3 ratio = 7.5:4.

Distance between r-m and dm-cu ~5.0× dm-cu. Cell dm without CuA1 stub vein (corner evenly rounded).

Male abdomen (Fig. 6.27): Sternite 4 posterior margin sntire. Sternite 5 posteromedially with shallow arcuate emargination and pair of long setae basal to emargination; emargination ~1/5 width of sternite. Transverse portion of sternite 6 broadly arcuate; right lateral portion. Synsternite 6+7 with narrow, medially projecting process with forked tip; posterior fork branch 1/3 anterior fork branch. Cercus triangular, shorter than surstylus, with series of 4 setae (setae shortening distally).

Surstylus (in lateral view) anterior lobe triangular, with small anterior process; posteriorly with 12–

15 elongate setae, distally with dark process. Postgonite with basal ½ quadrate; apical ½ triangular.

Distiphallus: dorsal sclerite with apical 1/3 projecting beyond apex of acrophallus; acrophallus dorsolaterally with acute, narrow process near midlength.

Female Abdomen (Fig. 6.28): Tergite 7 medially longitudinally desclertozied; posterior margin with broad rectangular emargination medially; posterolaterally narrowly fustedto tergite 8. Tergite 8 with lateral sclerites acutely rounded; medial sclerite narrow, triangular. Epiproct with lateral sclerites with anterior stems ~2/3 length of broad, posterior portion; posterior portions with anterolateral margins broadly rounded. Cercus elongate, strap-like; apical setae flattened with small flattened preapical seta appressed to base. Sternite 7 posteriorly broadly rounded. Spermatheca barrel-shaped; sclerotized portion of duct ~1/2 length of spermatheca.

Type Material: Holotype (male, debu00189531, UASC) and paratypes (2 males 2 females;

DEBU & UASC): BOLIVIA: La Paz: Cumbre Alto Beni, 28 km E Caranavi, ~1400m,

15°40’31”S, 67°29’21”W, dung pans, 14 Apr 2001, S.A. Marshall (UASC). Additional Paratypes:

- 167 -

BOLIVIA: La Paz: 1 male, 1 female, Caranavi, ca. 10 km NW, road to ENTEL tower, 15°46'35”S,

67°35'48”W, 1400 m, dung pans, 13 Apr 2001, S.A. Marshall

Etymology: The name refers to the two elongate setae present near the posterior margin of the male fifth sternite.

Archiceroptera caliga Paiero & Marshall, n. sp.

Figs. 6.29–6.30

Size: 1.8–2.8 mm.

Head: Ocellar triangle with 3–5 setae. Frons with 4–5 interfrontal seta and 4–5 inclinate orbital setulae. Eye:gena ratio = 2:1. Gena with 11–16 smaller setae.

Thorax: Acrostichal setulae in 6 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weakly developed.

Legs: Fore femur with 5–6 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 10 stout setae extending from base to apical 3/4; apicoventral series of 4–6 setae on apical

1/4; male ventral comb with 7–8 robust setae extending to midlength. Mid tibia with 2 median anterodorsal and 2 (rarely 3) median posterodorsal setae; strong predistal dorsal, and weak distal posterodorsal setae present; maleventral comb with 7–8 robust setae on apical 1/2; midventral seta present in female only. Hind tibia usually simple but sometimes with weak preapical seta present.

Wings: Costa with stronger costagial setae not reaching humeral crossvein. C2:C3 ratio = 10:7.

Distance between r-m and dm-cu ~5.0× dm-cu. Cell dm with CuA1 stub vein usually absent.

Male Abdomen (Fig. 6.29): Sternite 4 posteriorly entire. Sternite 5 posteromedially with broad triangular emargination extending ~2/5 length and ½ width of sternite; transverse seam at anterior extent of emargination. Transverse portion of sternite 6 sinuate; right lateral portion. Synsternite

6+7 side with broadly acute, medially projecting process, with small narrow sclerite apically. Cercus triangular, elongate, with 1 strong seta on basal 1/3. Surstylus (in lateral view) bootshaped; anterior lobe triangular (with 4–5 setae on surface); posteriorly with small triangular lobe apically, and 18–

21 long setae on distaly 2/3. Postgonite with basal ½ quadrate, ventrually sinuate; apical ½

- 168 - acuminate, dorsal surface entire. Distiphallus: dorsal sclerite with apical 1/5 projecting beyond apex of acrophallus; acrophallus dorsolaterally obtusely angled near aedeagal midlength.

Female Abdomen (Fig. 6.30): Tergite 7 posteromedially entire; posterolaterally narrowly fused to tergite 8. Tergite 8 with posterolateral corners rounded; medial sclerite rectangular, elongate (poorly sclerotized). Epiproct with lateral sclerites oval. Cercus ovoid. Sternite 7 posteromedially narrowly rounded.. Spermatheca triangular-ovoid; sclerotized portion of duct ~1/4 length of spermatheca.

Type Material: Holotype (male, debu01085225) + 18 Paratypes (18 males): ECUADOR:

Pichincha: Rio Palenque, dung, 25 Feb 1979, S.A. Marshall (QCAZ). Additional Paratypes:

BRAZIL: Pará: 1 male, Tucuruí, Jan 1979, M. Alvarensa (MZSP); Parana, 1 male, Londrina,

Mata dos Godoy, 28–31 Jan 1990, S.A. Marshall. COLOMBIA: Amazonas: 2 males, Leticia, 28

Feb 1974, V. Nealis. COSTA RICA: Alajuela: 2 males, Florencia Forest, dung tp., 28 Feb 1980,

H. Howden; 1 male, Volcán Tenorio, N slope near Bijagua Biological Station, 700 m, pan traps in tree fall, 18 Jun 2000, Buck & Marshall (INBC); Guanacaste: 1 male, Cacao Field Station, 1250m, dung trap, 12–15 Feb 1996, S.A. Marshall (INBC); 1 male, Estacion Pitilla, 9 km S Sta. Cecilia,

700m, Aug 1994, C. Moraga; 1 male, Estacion Pitilla, 9 km S Sta. Cecilia, 700m, Jul 1994, C.

Moraga; 2 males, Guanacaste Cons. Area, Pitilla Field Station, Malaise, 29 Jan 1996, J. Noyes;

Heredia: 1 male, La Selva, 50–100m, carrion trap, 18 Feb 1980, H.F. Howden; Limon: 4 males, 3 females, Estrella Valley, Pandora, carrion trap, 20 Feb 1984, H. Howden; 1 male, Bribri, 4 km NE,

50 m, Dec 1989–Mar 1990, P. Hanson; Puntarenas: 2 males, Golfo Dulce Forest Reserve, 24 km

W Piedras Blancas, 200 m, Dec 1990, P. Hanson; 1 male, Osa Peninsula, Rincón, 2.5 km S,

8°42'1"N, 83°30'50"W, ~50 m, secondary forest, dung pitfalls, 10 Aug 2001–11 Aug 2002.

ECUADOR: Emeraldas: 3 males, La Chiquita, 11 km SE San Lorenzo, 5 m, carrion, 9–10 Jun

1975, S. Peck (QCAZ); 4 males, La Chiquita, 11 km SE San Lorenzo, 5 m, carrion trap, day 2, 7–8

Jun 1975, S. Peck; 1 male, La Chiquita, 11 km SE San Lorenzo, 5 m, dung, 9–10 Jun 1975, S. Peck;

6 males, La Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, 10–11 Jun 1975, S. Peck; 9 males, La

Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, day 3, 8–9 Jun 1975, S. Peck; 9 males, 1 female,

La Chiquita, 17km SE San Lorenzo, 5m, dung, 7–8 Jun 1975, S. Peck (QCAZ); Napo: 3 males, 1 female, Jatun Sacha Reserve, 6 km E Misahuallí, 1°4'S, 77°37'W, 450 m, by stream, dung pans, 5–7 - 169 -

May 2002, S.A. Marshall; 4 males, 3 females, Jatun Sacha Reserve, 6 km E Misahuallí, 1°4'S,

77°37'W, 450 m, varzea, dung pans, 2–7 May 2002, M. Buck; 6 males, 1 female, Tena, 12 km SW,

500m, dung trap, 8–11 Aug 1976, S. Peck; 14 males, 7 females, Tena, 12 km SW, 500m, dung trap

30–33, day 2–5, 8–11 Jul 1976; 2 males, Tiputini Biodiversity Stn., vicinity Yasuni National Park,

0°38'S, 76°10'“W, pitfall trap (human dung), 14–19 Feb 1998, D.C. Darling (ROME); Pichincha: 1 male, Allurquin, 28km E, Chiribaga Rd., 5200ft, moss forest, carrion trap, 19–27 Jun 1975, S. Peck;

7 males, Palenque, day 3 trap, 24–25 Mar 1976, S. Peck; 16 males, Rio Palenque, dung, 27 Feb

1979, S.A. Marshall; 4 males, Rio Palenque, dung trap, 22–23 Feb 1976, S. Peck (QCAZ); 10 males, Rio Palenque, dung trap, 25–26 Feb 1976, S. Peck; 2 males, Rio Palenque, J. Glasser trap, 26

Feb 1976, S. Peck; 36 males, Rio Palenque, 25–26 Jan 1976, S. Peck; 4 males, 2 females, Rio

Palenque, 26 Feb 1976, J. Glaser; 1 male, Rio Palenque Stn., 47 km S Santo Domingo, 160m, 1° lowland rainforest, malaise head, 30 Apr–5 May 1987, Coote & Brown; 1 male, Río Palenque Stn.,

47 km S Santo Domingo, carrion, 27–28 May 1975, S. Peck; 3 males, Río Palenque Stn., 47 km S

Santo Domingo, traps 3–5, day 1, 22–23 Feb 1976, S. Peck; 8 males, Rio Palenque Stn., 47km S

Santo Domingo, 250m, dung, 17–25 Feb 1979, S.A. Marshall; 3 males, Rio Palenque Stn., 47km S

Santo Domingo, 250m, 17–25 Feb 1979, S.A. Marshall (QCAZ); 3 males, Rio Palenque Stn., 47km

S Santo Domingo, 250m, 17–25 Feb 1979; 6 males, Tinalandia, 1120m, wet lower montane rainforest, Malaise head, 9–13 May 1987, L.D. Coote & B.V. Brown; 1 male, Tinalandia, 680m, dung, 22–28 Jun 1975, S. Peck; 4 males, Tinalandia, 16 km SE Santo Domingo, 680 m, forest, dung traps 32, 16–28 Jun 1975, S. Peck; 3 males, Tinalandia, 16 km SE Santo Domingo, 680 m, rainforest, malaise-FIT, 4 May–25 Jul 1985, S. & J. Peck; 7 males, Tinalandia, 16 km SE Santo

Domingo, 680 m, dung trap, 21–22 Jun 1975, S. Peck. PANAMA: Chiriquí: 3 males, 1 female,

Hartmann's Finca, 15 km NW Hato de Volcán, 1200 m, dung trap, 20–25 May 1977, S. Peck; 2 males, 1 female, La Fortuna Dam, 1000 m, 5–6 Jul 1981, B. Gill; Colón: 2 males, Santa Rita Ridge,

10 mi SE Colón, 270 m, dung trap, 10–12 Jun 1977, S. Peck; (Prov. unknown): 14 males, 5 females, Canal Zone, Madden Forest, carrion trap, 10–13 Jun 1977, S. Peck. PERU: Loreto: 1 male, Campamento San Jacinto, 175–215 m, FIT, 7 Jul 1993, R. Leschen (MUSM); 1 male,

Campamento San Jacinto, 175–215 m, FIT, 5 Jul 1993, R. Leschen (MUSM); 9 males, Teniente - 170 -

López, FIT, 24 Jul 1993, R. Leschen (MUSM); 2 males, Teniente López, 1.5 km N, 2°31'S,

76°10'W, 230–305 m, FIT, 20 Jul 1993, R. Leschen; 2 males, San Jacinto, FIT, 12 Jul 1993, R.

Leschen (MUSM); Madre de Dios: 4 males, Zona Reserva Manu, Pakitza, 11°57’S, 71°17’W,

400m, Malaise trap, 18–23 Feb 1992, B. Brown & D. Feener .

Comments: The two records from Brazil appear anomalous but this may just reflect lack of available specimens from large areas of the neotropics.

Etymology: The species epithet is from the Latin for boot, referring to the boot-shaped surstylus.

Archiceroptera calligraphia Paiero & Marshall, n. sp.

Figs. 6.31–6.32

Size: 1.5–2.8 mm.

Head: Ocellar triangle with 3–5 setae. Frons with 4 interfrontal seta and.5–6 interfrontal pairs of setae. Eye:gena ratio = 2.2:1. Gena with 10–13 smaller setae.

Thorax: Acrostichal setulae in 8 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 5–7 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 9–10 stout setae extending from base to apical 3/4; apicoventral series of 4–5 setae on apical 1/4.; male, ventrally, with 10–12 robust setae in cluster on basal 1/3. Mid tibia with 2–3 median anterodorsal and 2 median posterodorsal setae; strong predistal dorsal and strong distal posterodorsal setae present; male ventral comb with 10–11 robust setae on apical 2/3; midventral seta present in female only. Mid basitarsus with basal posteroventral setae slightly longer than apical setae. Hind tibia usually with weak, preapical dorsal seta present.

Wings: Costa with stronger costagial setae not reaching humeral crossvein. C2:C3 ratio = 5:3.

Distance between r-m and dm-cu ~4.5–5.0× dm-cu. Cell dm sometimes with small CuA1 stub vein present.

Male Abdomen (Fig. 6.31): Sternite 4 entire. Sternite 5 posteromedially with acute, conical emargination extending anteriorly almost to anterior margin; emargination ~1/5 width of sternite; posterior margin weakly convex lateral to emarignation. Transverse portion of sternite 6 weakly - 171 - arcuate; right lateral portion recurved, fused with poorly defined quadrate sclerite. Synsternite 6+7 with narrow medially projecting process with tip posteriorly angled. Cercus basally wide with small seta, abruptly narrowing at basal 1/5 to ½ basal width; apical 4/5 gradually narrowed to narrowly rounded tip. Surstylus (in lateral view) boot-shaped; anterior lobe triangular and distal posteror corner with elongate, slightly curved process. Postgonite with basal ½ quadrate, apical ½ acuminate and with dorsal edge of distal process straight. Distiphallus: dorsal sclerite with distal 1/5 extending beyond apex of acrophallus; acrophallus with dorsolateral lobes broadly rounded.

Female Abdomen (Fig. 6.32): Tergite 7 posteromedially entire or weakly sinuate; posterolaterally narrowly fused to tergite 8. Tergite 8 with lateral sclerites posterolaterally rounded; medial sclerite broadly triangular. Epiproct with lateral sclerites elongate rectangular. Cercus elongate oval.

Sternite 7 narrowly rounded. Spermatheca ovoid; sclerotized portion of duct ~1/4 length of spermatheca.

Type Material: Holotype (male debu001085417, FMNH) + 14 paratypes (2 males, 12 females,

FMNH): MEXICO: Chiapas: Lagunas de Montebello Parque Nacional, Aqua Tinta, 4900ft, oak- pine, human dung, 21–24 Aug 1971, A. Newton (FMNH). Additional Paratypes: BELIZE: Cayo:

1 male, 3 females, Caves Branch, forest, dung, 23–29 Aug 1972, S. & J. Peck. COSTA RICA:

Alajuela: 2 males, Florencia Forest, dung tp., 28 Feb 1980, H. Howden; Cartago: 1 male,

Turrialba, Catie, Florence Forest, 600m, cup traps, 28 Feb 1980, H. & A. Howden; Heredia: 1 male, 10km N Puerto Viejo, La Selva Verde, FIT, 3 Mar 1991, H. & A. Howden; 1 male, La Selva

Biological Station, 3 km S Puerto Viejo, 80 m, FIT, 14–17 Jun 2001, S. Chatzimanolis (INBC); 1 male, Puerto Vieja, La Selva Biological Station, SHo+SOR 2nd growth, Malaise trap, 22–25 Apr

1989, Brown & Feener (INBC); Puntarenas: 2 males, Osa Peninsula, Rincón, 2.5 km S, 8°42'1"N,

83°30'50"W, ~50 m, prim. forest, dung pitfalls, 11 Aug 2001, M. Buck (INBC); 7 males, Osa

Peninsula, Rincón, 2.5 km S, 8°42'1"N, 83°30'50"W, ~50 m, secondary forest, dung pans, 11 Aug

2001, M. Buck; 1 male, Osa Peninsula, Rincón, 2.5 km S, 8°42'1"N, 83°30'50"W, ~50 m, secondary forest, fish pitfalls, 10–11 Aug 2001. ECUADOR: Guayas: 13 males, 9 females, 78 km N Santa

Elena, 27 km S Puerto López, 500ft, dung trap, 25–27 Jul 1976, S. Peck (QCAZ, DEBU).

GUATEMALA: Baja Verapaz: 1 male, Chilasco, 6.6km W, 1700m, dung, 30 May 1991, H. - 172 -

Howden; 1 female, Purulhá, 8 km S, dung trap, 25 May 1991, H. & A. Howden; Guatemala: 16 males, 7 females, Guatemala City, Universidad del Valle, dung traps, 11–13 Jun 1991, B.D. Gill

(UVGC, DEBU); 1 male, 2 females, Santa Catarina Pinula, dung traps, 11–13 Jun 1991, B.D. Gill.

HONDURAS: Francisco: 8 males, 9 females, Uyuca, 5200ft, pines, dung trap, 30 May 1994, H.

Howden; 5 males, 2 females, Uyuca, dung traps, 10 Jun 1994, H. & A. Howden; 24 males, 29 females, Cerro Uyuca, 30km E Tegucigalpa, 1800m, 4–10 Jun 1994, B. Gill; 5 males, 4 females,

Cerro Uyuca, 1800 m, dung, 30 May 1994, H. Howden; 1 male, Cerro Uyuca, 1800 m, malaise, 27

May 1994, H. Howden; 1 female, Uyuca, 1800 m, malaise trap, 6 Jun 1994, H. & A. Howden;

(Dpto. unknown): 14 males, 13 females, Cerro Monserrat, 7km SW Yusearan, dung trap, 24 May

1994, H. & A. Howden; 2 males, 6 females, Cerro Montserrat, 1800m, Malaise, 24 May 1994, H.

Howden; 8 males, 8 females, Lago de Yajoa, 2600ft, dung trap, 1–2 Jun 1994. MEXICO:

Campeche: 1 female, Escarcega, 6km W El Tormenta, 110m, evergreen tropical forest, 12–23 Jul

1983, S. & J. Peck; Chiapas: 6 males, 4 females, Bochil, 21mi. N, 5500ft, pine, oak, liquidamber, human dung, 18–24 Aug 1971, A. Newton; 2 males, 5 females, Bochil, 5mi. S, 5000ft, oak, human dung, 18–24 Aug 1971, A. Newton (UNAM); 2 males, Laguna Belgica, 16 km NW Ocozocoaulta,

970m, FIT, 13 Jun 1990, H. & A. Howden & Gill; 1 male, 1 female, Laguna Belgica, 16 km NW

Ocozocoaulta, 970m, 31 May 1990, H. & A. Howden; 16 males, 22 females, Lagunas de

Montebello Parque Nacional, La Eucantada, 4900ft, oak/pine/liquidambar, human dung, 21–24 Aug

1971, A. Newton (DEBU, FMNH); 1 male, Nahá, 16°56'57"N, 091°35'41"W, 960 m, mesophil forest, malaise trap, 29 May 2008; 1 female, Ocosingo Rd., 76km S Palenque, Rt. 195, 760m, rainforest, window trap, 5–29 Jul 1983, S. & J. Peck, & R. Anderson; 1 female, Parque Nacional

Sumidero, 1000m, 1 Jun 1990, H. & A. Howden; 1 male, Salto de Agua, 8 km SE, 17°30'45"N,

92°17'40"W, 60 m, 2° wet forest, malaise trap, 14–17 Jun 2008; 1 female, Teopisca, 5 mi NW,

16°34'48"N, 92°31'12"W, 6600ft, oak-pine-juniper woodland, human dung, 21–24 Aug 1971, A.

Newton; 5 males, 6 females, Trinitaria, 11mi. E, km 18.5, 5200ft, tropical deciduous forest, human dung, Aug 1971, A. Newton (FMNH, UNAM); Quintana Roo: 2 males, Kohunlich, 68 km W

Chetumal, 160 m, tropical seasonal forest, FIT, 14–17 Jul 1982, S. & J. Peck; 1 female, Kohunlich,

- 173 -

96km W Chetumal, seasonal tropical forest, carrion trap, 15–17 Jul 1983, S. & J. Peck; Veracruz: 1 male, Huatusco, 4mi N, 4100ft, cloud forest, dung, 11–16 Jul 1971, A. Newton.

Etymology: The species epithet refers to the shape of the emargination of the male sternite 5, which resembles the tip of a calligraphy pen.

Archiceroptera cobolorum Paiero & Marshall, n. sp.

Figs. 6.33–6.34

Size: 1.8–2.6 mm.

Head: Ocellar triangle with 5–6 small setae. Frons with 4 interfrontal seta and 4–5 inclinate orbital setulae. Eye:gena ratio = 2:1. Gena with 5–8 smaller setae.

Thorax: Acrostichal setulae in 6 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta extremely weak or absent (usually indistinct from nearby setulae).

Legs: Fore femur with 6–7 posterodorsal and 5–6 posteroventral setae. Mid femur anteriorly with series of 12–14 stout setae extending from base to apical 3/4; apicoventral series of 4–6 setae on apical 1/4.; male, ventrally, with 7–8 robust setae in dense cluster in basal 1/3. Mid tibia with 2 median anterodorsal and 2 median posterodorsal setae; predistal dorsal, and distal dorsal setae present; male ventral comb with 12–14 robust setae on apical 2/3; midventral seta present only in female. Mid basitarsus with basal posteroventral setae slightly larger than apical setae. Hind tibia with 5–7 anterodorsal and 5–7 posterodorsal setae (females usually with reduced number of setae, sometimes only the predistal posterodorsal seta present).

Wings: Costa with longer costagial setae extending to humeral crossvein. C2:C3 ratio = 5:4.

Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm with distinct, long CuA1 stub vein present.

Male Abdomen (Fig. 6.33): Sternite 4 posteriorly entire. Sternite 5 posteromedially with weak pair of tabs on each side of middle (sometimes nearly fused medially and indistinguishable); each tab with 5–8 pale flattened setae. Transverse portion of sternite 6 weakly sinuate to straight; right lateral portion recurved, simple. Synsternite 6+7 with small, broadly acute process. Cercus with basal ¼ transverse with 1 strong, long seta; apical ¾ acuminate. Surstylus (in lateral view) with long finger- - 174 - like process on anterior surface, and triangular process on posteroventral corner; posterior surface setose with 8–10 long setae. Postgonite narrow and elongate; basal ½ elongate rectangular with venter rounded; apical ½ acuminate and dorsal surface curved. Distiphallus: dorsal sclerite with apical 1/3 extending beyond apex; acrophallus broadly expanded with dorsolateral quadrate process recurved dorsally and with ventrolateral process rounded.

Female Abdomen (Fig. 6.34): Tergite 7 posteromedially broadly rounded narrowing laterally; posterolaterally broadly fused tergite 8.Tergite 8 with lateral sclerites acute, with small ventral subapical tooth; medial sclerite oblong ovoid. Epiproctwith lateral sclerites transverse. Cercus ovoid. Sternite 7 posteromedially broadly emarginate (~1/2 sternite width). Single spermatheca ovoid with large invaginations on both ends, paired spermathecae kidney-shaped with large invaginations on both ends; sclerotized portion of duct ~1/4 length of spermatheca.

Type Material: Holotype (male, debu01086490, QCAZ) + 102 Paratypes (40 males, 62 females;

QCAZ, DEBU): ECUADOR: Napo: Baeza, 15km NW, 2200m, dung trap, 2–6 Mar 1976, S. Peck.

Additional Paratypes: BRAZIL: São Paulo: 4 males, USP Biology Station, human dung, 5–6 Feb

1979, R. Woodruff & J. Runnacles (DEBU, MZSP). COLOMBIA: Cundinamarca: 1 male,

Bogota, 12 km S, 10000 ft, dung trap, 28 Feb–6 Mar 1972, S. & J. Peck; Norte de Santander: 3 males, 4 females, Pamplona, above, 9000ft, dung trap, 9–13 May 1974, S. Peck; (Dpto. unknown):

5 males, 3 females, Tequendama Falls, 30km SW Bogota, dung trap, 27 Feb–6 Mar 1972, S. & J.

Peck. ECUADOR: Napo: 1 male, 2 females, Baeza, 27km NW, 2700 m, dung trap, 2–6 Mar 1976,

S. Peck; 1 female, Baeza, 7 km S, 2000 m, dung trap, 23 Feb 1979, H. & A. Howden (QCAZ); 3 males, 4 females, Cosanga, 2.5 km W, 0°35'24"S, 77°53'19"W, 2150 m, dung/pans, 5–7 Nov 1999,

S.A. Marshall; Pichincha: 2 females, Bellavista Reserve, 0°0'54"S, 78°40'56"W, 2200 m, 30 Oct

1999, S.A. Marshall; 2 females, Rio Palenque, dung trap, 22–23 Feb 1976, S. Peck. PERU: Cusco:

1 female, Wayqecha Biological Station, ~9km NE Challabamba, 13°10'20"S, 71°35'0"W, 2600–

2700m, Trecha Ferdiz, dung pans, 3 Dec 2011, S.A. Marshall (MUSM, DEBU).

Comments: The surstylar shape of R. cobolorum males resemble those of the brevivilla species group (especially R. megavilla).

- 175 -

Etymology: The species epithet is the Latin for “goblin”, referring to the profile of the surstylus, which has a general similarity to a facial profile of a goblin.

Archiceroptera dolabra Paiero & Marshall n. sp.

Fig. 6.35

Size: 2.7–2.8 mm.

Head: Ocelli developed with 10–12 setae. Frons with 4–5 interfrontal seta and 5–6 inclinate orbital setulae. Eye:gena ratio = 2.5:1. Gena with 12–17 smaller setae.

Thorax: Acrostichal setulae in 8 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 6–7 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 8–9 stout setae extending from base to apical 3/4; apicoventral series of 5–6 setae on apical

1/4; male, ventrally, with 14–18 robust setae in elongate cluster on basal 1/2. Mid tibia with 2 median anterodorsal and 2 median posterodorsal setae; strong predistal dorsal and strong distal posterodorsal setae present; male ventral comb with 11–12 robust setae on apical 2/3; midventral seta absent in male (female unknown). Mid basitarsus with basal posteroventral setae slightly larger than apical setae. Hind tibia simple.

Wings: Costa with longer costagial seta not reaching humeral crossvein. C2:C3 ratio = 5:3.

Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm with small CuA1 stub vein present.

Male abdomen (Fig. 6.35): Sternite 4 posteromedially with broad triangular emargination; emargionation ~4/5 width and 1/3 length of sternite. Sternite 5 posteromedially broadly emarginate; emargination broadly rounded, extending 2/5 length and 3/4 width of sternite. Transverse portion of sternite 6 weakly arcuate, with right lateral portion not recurved. Synsternite 6+7 with broadly acute, medially projecting process on left and small elongate sclerite adjacent to tip of process.

Cercus elongate, with basal 1/6 weakly triangular; large seta present near basal 1/3. Surstylus (in lateral view) with anterior lobe acutely triangular (with 3–4 setae near tip); distally rounded and with 13–15 elongate setae on posterior surface. Postgonite with basal ½ quadrate, apical half acuminate with dorsal surface with distinct swelling near apical 1/3. Distiphallus: dorsal sclerite no - 176 - projecting beyond apex of acrophallus; acrophallus dorsolaterally with broadly rounded, dorsally recurved projections.

Female: Unknown.

Type Material: Holotype (male, debu00385443): FRENCH GUIANA: St. Laurent du Maroni:

Maripasoula, Mitaraka, MIT-DZ, 2°14'2"N,7 54°27'1"W, 306m, tropical moist forest (plateau- slope-cleared), FIT, 6 Mar 2015, Touroult & Poirier (MHNM). Paratype: FRENCH GUIANA: St.

Laurent du Maroni: 1 male, Maripasoula, Mitaraka, near base camp & along trails, tropical moist forest, SLAM trap, 1–6 Mar 2015, Touroult & Poirier.

Comments: The male abdomen is similar to A. caliga.

Etymology: The species epithet is from the Latin for pickaxe, referring to the shape of the male surstylus.

Archiceroptera maniba Paiero & Marshall, n. sp.

Figs. 6.36–6.37

Size: 2.3–2.8 mm.

Head: Ocellar triangle with 6–7 setae. Frons with 4 interfrontal seta and 5–7 inclinate orbital setulae. Eye:gena ratio = 2.5:1. Gena with 11–13 smaller setae.

Thorax: Acrostichal setulae in usually in 8 rows (rarely 6) between prescutellar dorsocentrals.

Scutellum without setulae. Anterior katepisternal seta indistinguishable from nearby setulae or absent

Legs: Fore femur with 7–8 posterodorsal and 4–6 posteroventral setae. Mid femur anteriorly with series of 9–10 stout setae extending from base to apical 3/4; apicoventral series of 4–5 setae on apical 1/4.; male, ventrally, with 12–14 robust setae in cluster on basal 1/3. Mid tibia with dorsal basal seta, 2 median anterodorsal, 2 median posterodorsal, strong predistal anterodorsal, strong predistal dorsal and strong apical posterodorsal setae; male ventral comb with 14–17 robust setae on apical 2/3; midventral seta present only in female. Hind tibia simple.

Wings: Costa with longer costagial setae not reaching humeral crossvein. C2:C3 ratio = 5:3.

Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm without CuA1 stub vein present. - 177 -

Male Abdomen (Fig. 6.36): Sternite 4 posteromedially entire or weakly emarginate. Sternite 5 posteromedially with large triangular emargination flanked by elongate process; emargination broadly rounded anteriorly, extending ~1/2 width and ½ length of sternite; processes with 6–7 flattened setae. Transverse portion of sternite 6 distinctly arcuate medially, right lateral portion recuved. Synsternite 6+7 with broadly acute medially projecting process ending in a quadrate sclerite with a distinct posteromedially curved process. Cercus basally transverse, ovoid with small seta; abruptly narrowed at basal ¼ with distal ¾ narrowing. Surstylus (in lateral view) with anteror lobe elongate rectangular, with small finger-like process on dorsal surface, near midlength of lobe; posteriorly with distal corner narrowly rounded and with 14–16 long setae present on surface.

Postgonite with ventral surface broadly rounded on basal ¼; dorsal surface relatively straight.

Distiphallus: dorsal sclerite with apical 1/5 extending beyond apex of acrophallus; acrophallus dorsolaterally broadly rounded near midlength of distiphallus.

Female Abdomen (Fig. 6.37): Tergite 7 posteromedially with broad, weak emargination; posterolaterally narrowly fused to tergite 8. Tergite 8 with lateral sclerites posterolaterally broadly rounded; medial sclerite triangular. Epiproct with lateral sclerites anterolaterally broadly rounded.

Cercus oval. Sternite 7 with posterior margin broadly rounded. Spermatheca circular, partially flattened; sclerotized portion of duct ~1/3 length of spermatheca.

Type Material: Holotype (male, debu00378573, ROME) + 5 paratypes (2 males, 3 females;

ROME, DEBU): GUYANA: Potaro-Siparuni: Mount Wokomung, 5°6'35"N, 59°49'15"W, 500m,

1234m, 1° rainforest, human dung, pitfall trap, 27 Oct–1 Nov 2004, B. Hubley. Additional

Paratypes: VENEZUELA: Bolivar: 1 male, El Dorado, 135km S, 1400m, dung traps, 20 Jul–7

Aug 1986, B. Gill (MIZA); 1 male, km40 Sta. Elena Icabaru Rd., 220m, 4–6 Aug 1986, B.D. Gill

(MIZA); 1 male, km40 Sta. Elena Icabaru Road, 100m, 4 Aug 1986, B.D. Gill; 2 males, km40 Sta.

Elena Icabaru Road, 1000m, 4–6 Aug 1986, B.D. Gill;

Comments: Like several other species, A. maniba has a distinct sclerite at the apex of the medially projecting process on the left side of synsternite 6+7.

Etymology: The species epithet is from the Latin for “hands”, referring to the resemblance of the posterior extensions of sternite 5 to a pair of hands. - 178 -

Archiceroptera masoni Paiero & Marshall, n. sp.

Figs. 6.38–6.39

Size: 1.5–2.1 mm.

Head: Ocellar triangle with 3–5 setae. Frons with 5–7 interfrontal seta and 4–5 inclinate orbital setulae. Eye:gena ratio = 1.8:1. Gena with 10–12 smaller setae.

Thorax: Acrostichal setulae in 8 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 4–5 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 10–12 stout setae extending from base to apical 3/4; apicoventral series of 3–5 setae on apical 1/4.; male, ventrally, with 6–7 robust setae in double linear series on basal 1/3. Mid tibia with

2 median anterodorsal and 1 median posterodorsal setae; weak predistal dorsal and weak distal posterodorsal setae present; male ventral comb with 5–7 robust setae on apical 1/3; midventral seta present in both sexes. Mid basitarsus with basal posteroventral setae not distinctly longer than distal setae. Hind tibia simple.

Wings: Costa with longer costagial setae not reaching humeral crossvein. C2:C3 ratio = 5:4.

Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm with small CuA1 stub vein present.

Male Abdomen (Fig. 6.38): Sternite 4 posteromedially entire. Sternite 5 posteromedially with broad, shallow arcuate emargination; emargination ~1/3 width and 1/6 length of sternite. Transverse portion of sternite 6 weakly arcuate to straight; right lateral portion recuved. Synsternite 6+7 withblunt, medially projecting process; process with small posterodistal spur. Cercus elongate, triangular with large seta near midlength. Surstylus (in lateral view) anteriorly with small finger-like process near apical 1/3; posterior surface with 12–15 large setae. Postgonite with ventral surface rounded near base, posterodistal surface relatively straight, and apical ½ wedge-shaped.

Distiphallus: dorsal sclerite only slightly projecting beyond apex of acrophallus; acrophallus with dorsolateral process broadly rounded.

Female Abdomen (Fig. 6.39): Tergite 7 posteromedially entire; posterolaterally narrowly fused to tergite 8. Tergite 8 with lateral sclerites broadly rounded posterolaterally; medial sclerite triangular, - 179 - elongate. Epiproct with lateral sclerites triangular, anterolaterally broadly rounded. Cercus triangular. Sternite 7 posteriorly broadly rounded. Spermatheca barrel-shaped, with invagination at end opposite from duct; sclerotized portion of duct ~1/3 length of spermatheca.

Type Material: Holotype (male, debu001085696, UNAM) and 6 paratypes (3males, 3 females;

DEBU, UNAM): MEXICO: Sinaloa: Concordia, 20mi. E, 3000 ft, 4 Aug 1964, W.R.M. Mason.

Additional Paratypes: MEXICO: Sinaloa: 1 male, Concordia, 20mi. E, 3000ft, 8 Aug 1964,

W.R.M. Mason; 2 males, El Palmito, 15mi. W, 5000ft, 30 Jul 1964, W.R.M. Mason.

Comments: Several of the type specimens have collapsed, presumably due to air-drying. The holotype is mounted upside down.

Etymology: The species’ name is a patronym is in honour of the collector of the type series, the

Canadian hymenopterist W.R.M. Mason.

Archiceroptera megacercus Paiero & Marshall, n. sp.

Figs. 6.40–6.41

Size: 1.5–2.8 mm.

Head: Ocellar triangle with 5–6 setae. Frons with 5–6 interfrontal seta and 3–4 inclinate orbital setulae. Eye:gena ratio = 1:1. Gena with 0–11 smaller setae.

Thorax: Acrostichal setulae in 4 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 7–8 posterodorsal and 5–6 posteroventral setae. Mid femur anteriorly with series of 9–11 stout setae extending from base to apical 3/4; apicoventral series of 4–6 setae on apical 1/4.; male, ventrally, with 12–15 robust setae in cluster on basal 1/3 and 4–6 extending apically along posteroventral margin. Mid tibia with 1 median anterodorsal (rarely 2; if 2nd present very weak) and 1 median posterodorsal (rarely 2; if 2nd present very weak) setae; strong predistal dorsal and strong apical posterodorsal setae present; male ventral comb with 5–8 robust setae on apical 3/5; midventral seta present only on female. Mid basitarsus with basal posteroventral setae slightly longer than distal setae.Hind tibia with 1 anterodorsal and 1 posterodorsal seta near midlength (often reduced in female), with 2–4 weaker setae sometimes present apical to medial pair. - 180 -

Wings: Costa with longer costagial seta reaching humeral crossvein. C2:C3 ratio = 5:3. Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm usually with small CuA1 stub vein present.

Male Abdomen (Fig. 6.40): Sternite 4 posteromedially entire. Sternite 5 posteromedially with shallow emargination with small rounded teeth on lateral corners; emargination ~1/5 width and 1/10 length of sternite. Transverse portion of sternite 6 straight to weakly arcuate; right lateral portion recurved, fused with small quadrate sclerite. Synsternite 6+7 with narrow, elongate, medially projecting process. Cercus acuminate, with seta near basal 1/4. Surstylus (in lateral view) with bare, broadly conical anterior lobe; posterior portion hirsute with 16–20 elongate setae. Postgonite basal

2/3 quadrate, with apical 1/3 acuminate; anterodorsal surface with deep emargination. Distiphallus: dorsal sclerite not extending beyond apex of acrophallus; acrophallus dorsolaterally with broadly rounded, short process near midlength of distiphallus.

Female Abdomen (Fig. 6.41): Tergite 7 with posterior margin medially interruptedby deep emargination; emargination extending ¾ tergite length, broadening anteriorly; posterolaterally narrowly fused to tergite 8. Tergite 8 with lateral sclerites broadly rounded; medial sclerite elongate, rectangular. Epiproct with lateral sclerites triangular. Cercus elongate oval. Sternite 7 posteriorly broadly rounded. Spermatheca ovoid, slightly flattened; sclerotized portion of duct short, ~1/4 length of spermatheca.

Type Material: Holotype (male, debu01084917, QCAZ) + 46 Paratypes (22 males, 24 females;

DEBU, QCAZ): ECUADOR: Napo: Tena, 12 km SW, 500m, dung trap, 8–11 Aug 1976, S. Peck.

Additional Paratypes: BOLIVIA: La Paz: 1 male, Arroyo Tuhiri W Mapiri, 15°17'27"S,

68°15'29"W, 10 Apr 2001, S.A. Marshall (UASC); 1 male, Cumbre Alto Beni, 28 km E Caranavi,

15°40'31"S, 67°29'21"W, ~1400 m, dung pans, 14 Apr 2001, S.A. Marshall (UASC); 1 male, 3 females, San Antonio, ca. 8 km S Mapiri, 15°20'56"S, 68°13'31"W, secondary forest, dung pans, 11

Apr 2001, S.A. Marshall. BRAZIL: Paraná: 1 male, 1 female, Curitiba, 30 km SE, BR 277, dung traps, 6–9 Feb 1990, S.A. Marshall (MZSP); São Paulo: 1 male, 1 female, USP Biology Station, human dung, 5–6 Feb 1979, R. Woodruff & J. Runnacles. COLOMBIA: Amazonas: 5 males,

Leticia, pepper farm, dung, 1 Mar 1974, V. Nealis; 9 males, Leticia, dung traps, 28 Feb 1974, V.

Nealis. ECUADOR: Esmeraldas: 2 females, La Chiquita, 11 km SE San Lorenzo, 5 m, carrion, 9– - 181 -

10 Jun 1975, S. Peck; 1 male, La Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, 10–11 Jun 1975,

S. Peck; 4 males, La Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, day 3, 8–9 Jun 1975, S.

Peck; 3 males, 3 females, La Chiquita, 17km SE San Lorenzo, 5m, dung, 7–8 Jun 1975, S. Peck; 5 males, 5 females, La Chiquita, 17km SE San Lorenzo, 5m, dung, day 2, 7–8 Jun 1975, S. Peck

(QCAZ); Napo: 29 males, 28 females, Tena, 12 km SW, 500m, dung trap 30–33, day 2–5, 8–11 Jul

1976; 1 male, Tiputini Biodiversity Station, vicinity Yasuní National Park, 0°38'S, 76°0'W, human dung pitfalls, 14–19 Feb 1998, D.C. Darling (ROME); 1 male, Tiputini Biodiversity Stn.,

0°36'50"S, 76°9'1"W, sweep, May 2011, S.A. Marshall; 1 male, Tiputini Biodiversity Stn.,

0°36'50"S, 76°9'1"W, May 2011, S.A. Marshall; 1 male, Tiputini Biodiversity Stn., vicinity Yasuni

National Park, 0°38'S, 76°10'W, pitfall trap (human dung), 14–19 Feb 1998, D.C. Darling (ROME);

1 female, Yasuni National Park, Yasuni Research Station, rainforest, Malaise trap, 3–20 Nov 1998,

Pape & Viklund. FRENCH GUIANA: St. Laurent du Maroni: 1 female, Maripasoula, Mitaraka,

MIT-DZ, 2°14'2"N, 54°27'1"W, 306m, tropical moist forest near DZ, FIT, 6–10 Mar 2015, Touroult

& Poirier (MHNM); 1 male, Maripasoula, Mitaraka, near base camp & along trails, tropical moist forest (undergrowth), FIT, 7 Mar 2015, Touroult & Poirier (MHNM); 1 male, Mitaraka, MIT-C-

RBF2, 2°14'3"N,7 54°26'53"W, 299m, tropical wet forest (bas fond), yellow pan traps, 6–10 Mar

2015, M. Pollett. GUYANA: Potaro-Siparuni: 1 male, 1 female, Mount Wokomung, 5°7'53"N,

59°48'31"W, 698m, 1° forest, pitfall trap (human dung), 21–26 Oct 2004, B. Hubley (ROME);

(distr. unknown): 6 males, 7 females, Kabocalli, Iwokrama Forest Reserve, 100 m, FIT, 22–25

May 2001, Brooks & Falin. PERU: Loreto: 1 male, Campamento San Jacinto, 175–215 m, FIT

#42, 7 Jul 1993, R. Leschen (MUSM); 2 females, Campamento San Jacinto, 175–215 m, FIT, 5 Jul

1993, R. Leschen (MUSM); 2 females, Campamento San Jacinto, 175–215 m, FIT, 9 Jul 1993, R.

Leschen; 5 males, Teniente López, Riv. forest, FIT, #211, 26 Jul 1993, R. Leschen; 1 male, 2 females, Teniente López, riverine forest, FIT, #199, 24 Jul 1994, R. Leschen; 1 male, 1 female,

Teniente López, FIT, 23 Jul 1993, R. Leschen (MUSM); 1 male, 2 females, Teniente López, FIT, 26

Jul 1993, R. Leschen; 2 males, 3 females, Teniente López, 1.5 km N, 230–305 m, FIT, 18 Jul 1993,

R. Leschen; 1 female, Teniente López, 1.5 km N, 230–305 m, FIT, 20 Jul 1993, R. Leschen

(MUSM); 1 female, Teniente López, 1.5 km N, 230–305 m, FIT, 22 Jul 1993, R. Leschen; Madre - 182 - de Dios: 1 male, Pantiacolla Lodge, Alto Madre de Dios River, 12°39'18”S, 71°13'54”W, 420 m,

14–19 Nov 2007, FIT, D. Brzoska; San Loreto: 1 male, San Jacinto, FIT, 12 Jul 1993, R. Leschen.

VENEZUELA: Aragua: 2 males, 2 females, Rancho Grande Biological Station, 10°21'N,

67°41'W, 1200 m, flight intercept trap, 14 May 1998, Ashe, Brooks & Hanley (DEBU, MIZA); 1 male, Rancho Grande, La Cumbre cloud forest, 1500m, FIT, 1–10 Aug 1987, Borden & Peck;

Bolivar: 1 male, 2 females, 22km S El Dorado, lowland rainforest, FIT, 25 Jun–12 Jul 1987, S. & J.

Peck; 1 male, 1 female, km40 Sta. Elena Icabaru Rd., 220m, 4–6 Aug 1986, B.D. Gill (MIZA); 4 males, km40 Sta. Elena Icabaru Road, 1000m, 4–6 Aug 1986, B.D. Gill; 2 males, 1 female, km40

Sta. Elena Icabaru Road, 100m, 4–6 Aug 1986, B.D. Gill; 1 male, Quebrada de Jaspe, 19–20 Jul

1986, B. Gill (MIZA); Lara: 1 male, 1 female, Yacambu, 1200m, cloud forest, 7 May 1981, H.K.

Townes; 1 male, 1 female, Yacambu, 1200m, 10 May 1981, H.K. Townes.

Comments: One male from French Guiana has irregular mid tibial chaetotaxy (smaller setae present basal to both the anterodorsal and posterodorsal setae at the midlength) but the genitalia of the irregular specimen seem identical to other specimens.

Etymology: The species epithet refers to the enlarged male cercus.

Archiceroptera mexicorona Paiero & Marshall, n. sp.

Figs. 6.42–6.43

Size: 1.8–2.5 mm.

Head: Ocellar triangle with 3–5 setae. Frons with 4–5 interfrontal seta and 4–5 inclinate orbital setulae. Eye:gena ratio = 2:1. Gena with 8–10 smaller setae.

Thorax: Acrostichal setulae in 7–8 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 5–6 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 10–11 stout setae extending from base to apical 3/4; apicoventral series of 3–4 setae on apical 1/4.; male, ventrally, with 6–7 robust setae in linear series on basal 1/3. Mid tibia with 1–2 median anterodorsal (basal seta usually weak) and 2 median posterodorsal (basal seta usually weak) setae;, weak predistal dorsal, and weak apical posterodorsal setae present; male ventral comb 7–8 - 183 - robust setae on apical 3/5; midventral seta present only on females. Mid basitarsus with basal posteroventral setae slightly longer than distal setae. Hind tibia simple.

Wings: Costa with longer costagial seta not reaching humeral crossvein. C2:C3 ratio = 10:7.

Distance between r-m and dm-cu ~3.5× dm-cu. Cell dm usually without CuA1 stub vein.

Male Abdomen (Fig. 6.42): Sternite 4 posteromedially entire. Sternite 5 posteromedially with pair of rounded processes on each side of narrow emargination; emargination extending ~1/4 length and

1/8 width of sternite and with fine setae between tips of processes. Transverse portion of sternite 6 straight or weakly arcuate; right lateral portion recurved. Synsternite 6+7 with narrowly-rounded medially projecting process extending to; elongate sclerite. Cercus basally transverse with lateral seta, abruptly narrowly at basal 1/5, with apical 4/5 gradually narrowing. Surstylus (in lateral view) slightly boot-shaped, with anterior lobe elongate, with narrow, distal dorsal process (inner surface with additional process visible in posterior view); distal posterior corner broadly rounded; surface entiresly hirsute with 19–23 elongate setae. Postgonite with ventral surface basally rounded, sinuate near midlength; apical ½ wedge-shaped. Distiphallus: dorsal sclerite with apical 1/3 extending beyond apex of acrophallus; acrophallus with dorsolateral corners narrowly rounded.

Female Abdomen (Fig. 6.43): Tergite 7 with posterior margin broadly rounded, medially interrupted by shallow medial emargination (~1/12 tergite width); posterolaterally narrowly fused to tergite 8. Tergite 8 with posterolateral corner broadly rounded; medial sclerite small, elongate.

Epiproct with anterolaterals corner broadly rounded. Cercus quadrate; flattened apical setae with smaller flattened preapical seta appressed at base. Sternite 7 posteromedially acute, narrowly rounded. Spermatheca ovoid; sclerotized portion of duct ~2/5 length of spermatheca.

Type Material: Holotype (male, debu01085706, UNAM) and 11 paratypes (4 males 7 females;

DEBU, UNAM): MEXICO: Guerro: Ixtapa, 45km NE, 10–12 Jul 1987, B. Gill.

Comments: Females of this species resemble A. barberi, a widespread, commonly collected species.

Etymology: The species epithet is a combination of two words: mexi- refers to the type country, and -corona refers to the crown-like shape of the male sternite 5.

- 184 -

Archiceroptera mitarakai Paiero & Marshall, n. sp.

Fig. 6.44

Size: 2.1–2.3 mm.

Head: Ocellar triangle with 5 setae, with smaller pair of ocellar setae present. Frons with 4–5 interfrontal seta and 4–5 inclinate orbital setulae. Eye:gena ratio = 2:1. Gena with 9 smaller setae.

Thorax: Acrostichal setulae in 8 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 5 posterodorsal and 4 posteroventral setae. Mid femur anteriorly with series of 10–11 stout setae extending from base to apical 3/4; apicoventral series of 3–4 setae on apical

1/4.; male, ventrally, with 13 robust setae in cluster on basal 1/3. Mid tibia with 2 median anterodorsal and 2 median posterodorsal setae; weak predistal dorsal and strong distal posterodorsal setae present; male ventral comb with 7–8 robust setae on apical 2/3; midventral seta absent in male

(female unknown). Mid basitarsus with basal posteroventral setae not distinctly longer than distal setae. Hind tibia simple (some setae are slightly strengthened in preapical region).

Wings: Costa with longer costagial seta not reaching humeral crossvein. C2:C3 ratio = 5:3.

Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm with small CuA1 stub vein present.

Male abdomen (Fig. 6.44): Sternite 4 posteromedially entire. Sternite 5 posteromedially with broad arcuate emargination; emargination ~3/5 length and ½ width of sternite. Transverse portion of sternite 6 medially arcuate; right lateral portion recurved. Synsternite 6+7 with medially projecting process narrowly rounded apically, ending near elongate apical sclerite. Cercus elongate, triangular, with slight constriction and long seta near basal 1/3. Surstylus (in lateral view) elongate, with short, round anterior lobe (acutely rounded on distal dorsal corner; posterior surface with 12–15 long setae. Postgonite with ventral surface rounded basally, sinuate at midlength; dorsal surface with slight constriction near midlength. Distiphallus: dorsal sclerite with apical 1/8 extending beyond apex of acrophallus; acrophallus dorsolaterally broadly rounded at midlength of distiphallus.

Female: Unknown

Type Material: Holotype (male, debu00394580, MHNM): FRENCH GUIANA: St. Laurent du

Maroni: Maripasoula, Mitaraka, MIT-DZ-RBF1, 2°14'4"N, 54°27'2"W, 270m, tropical wet forest - 185 -

(bas fond), yellow pan traps, 2–10 Mar 2015, M. Pollet. Paratype: FRENCH GUIANA: 1 male,

Mitaraka, MIT-DZ, 2°14’2”N 54°27’1”W, 306m, tropical moist forest (plateau-slope-cleared), FIT,

1 Mar 2015, Touroult & Poirier.

Etymology: The species epithet refers to the type locality.

Archiceroptera paracercus Paiero & Marshall, n. sp.

Fig. 6.45

Size: 2.0–2.4 mm.

Head: Ocellar triangle with 5 setulae. Frons with 4 interfrontal setae and 3–4 inclinate orbital setulae. Eye:gena ratio = 2:1. Gena with 5–8 smaller setae.

Thorax: Acrostichal setae in 6 rows between prescutellar dorsocentrals. Scutellum without setulae.

Anterior katepisternal seta present but weak.

Legs: Fore femur with 7–8 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 9–10 stout setae extending from base to apical 3/4; apicoventral series of 3–4 setae on apical 1/4; male, ventrally, with 13–15 robust setae in elongate basal cluster. Mid tibia with 1 median anterodorsal and 1 median posterodorsal setae; strong predistal dorsal, and weak distal posterodorsal setae present; male ventral comb with 10–12 robust setae extending to basal 1/3; midventral seta present in females only. Mid basitarsus with basal posteroventral setae weak, slightly longer than distal setae.Hind tibia dorsally usually simple (weak preapical seta present on some individuals).

Wings: Costa with longer costagial seta reaching humeral crossvein. C2:C3 ratio = 2:1. Distance between r-m and dm-cu ~5.0× dm-cu. Cell dm without CuA1 stub vein.

Male abdomen (Fig. 6.45): Posterior margin of sternite 5 with weak medial emargination; emargination ~1/5 sternite width and < 1/10th sternite length. Transverse portion of sternite 6 straight or weakly arcuate; right lateral portion simple and recuved.Synsternite 6+7 with narrow, elongate, medially projecting process. Cercus acuminate, with strong seta near basal 1/4. Surstylus

(in lateral view) with setose strap-like posterior portion and bare triangular anterior lobe, with small preapical constriction. posterior portion with 14–19 elongate setae and small apical triangular - 186 - process. Postgonite basal 2/3 quadrate, with apical 1/3 acuminate; anterodorsal surface with deep emargination. Distiphallus: dorsal sclerite not extending beyond apex of acrophallus; acrophallus dorsolaterally with broadly rounded, short process near midlength of distiphallus.

Female Abdomen: Unknown

Type Material: Holotype (male, debu00260801, QCAZ) + 3 Paratypes (3 males; DEBU, QCAZ):

ECUADOR: Esmeraldas: La Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, day 3, 8–9 Jun

1975, S. Peck. Additional Paratypes: COSTA RICA: Guanacaste: 1 male, Guanacaste

Conservation Area, Ricon de la Vieja, Las Pailas, 1400m, Clusea rosea forest litter, 18–20 Feb

1996, R. Anderson (INBC); ECUADOR: Esemeraldas: 1 male, La Chiquita, 11 km SE San

Lorenzo, 5 m, dung trap, 10–11 Jun 1975, S. Peck; 1 male, La Chiquita, dung, 7–8 Jun 1975, S.

Peck.

Etymology: The species epithet refers to the similarity to A. megacercus.

Archiceroptera pussula Paiero & Marshall, n. sp.

Figs. 6.46–6.47

Size: 1.6–2.4 mm.

Head: Ocellar triangle with 5 setae. Frons with 4 interfrontal seta and 6–7 inclinate orbital setulae.

Eye:gena ratio = 1.5:1. Gena with 11–13 smaller setae.

Thorax: Acrostichal setulae in 8 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 5–6 posterodorsal and 5–6 posteroventral setae. Mid femur anteriorly with series of 7–8 stout setae extending from base to apical 3/4; apicoventral series of 4–5 setae on apical

1/4; male, posteroventrally, with 5–7 robust setae in elongate cluster on basal 1/3. Mid tibia with 1

(rarely 2) median anterodorsal and 1 (rarely 2) median posterodorsal setae; weak predistal dorsal and strong distal posterodorsal setae present; male ventral comb with 9–11 robust setae on apical

2/3; midventral seta present only on females. Mid basitarsus with basal posteroventral setae slightly longer than distal setae.Hind tibia simple.

- 187 -

Wings: Costa with longer costagial setae not reaching humeral crossvein. C2:C3 ratio = 5:4.

Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm with CuA1 stub vein usually present.

Male Abdomen (Fig. 6.46): Sternite 4 posteromedially entire. Sternite 5 posteromedially with broad, shallow emargination; emargination ~3/4 width and 1/3 length of sternite. Transverse portion of sternite 6 weakly arcuate; right lateral portion recurved. Synsternite 6+7 with acute medially projecting process with elongate apical sclerite (closely approximated with posterior margin of transverse portion of sternite 6). Cercus triangular, elongate, with strong seta on basal ¼; apically partially twisted on apical ¼, making apex (in dorsal view) appear acute (actually broadly rounded in lateral view). Surstylus (in lateral view) weakly boot-shaped, with conical anterior lobe and sinuate distal margin; anterior lobe with small distal swelling. Postgonite with ventral margin broadly rounded on basal ¼, sinuate near mid-length; apical ½ acuminate. Distiphallus: dorsal sclerite not projecting beyond apex of acrophallus; acrophallus dorsolaterally with rounded obstuse projection near midlength of distiphallus.

Female Abdomen (Fig. 6.47): Tergite 7 posteromedially weakly sinuate; posterolaterally narrowly fused to tergite 8. Tergite 8 with lateral sclerites broadly rounded posterolaterally; medial sclerite elongate, narrow. Epiproct broadly rounded anterolaterally. Cercus elongate oval. Sternite 7 posteriorly broadly rounded. Spermatheca ovoid; sclerotized portion of duct ~1/3 length of spermatheca.

Type Material: Holotype (male, debu01085653, UASC) + 1 paratype (1 male): BOLIVIA:

Santa Cruz: Potrerillos de Guenda, 17°40'29"S, 63°27'22"W, 4–7 Apr 1998, H. & A. Howden.

Additional Paratypes: ARGENTINA: Jujuy: 1 male, Calilegua National Park, Estaca El Cero,

900m, forest, carrion trap, 19–20 Dec 1987, S. & J. Peck; Salta: 2 males, 3 females, Alto de la

Sierra, 22°44'S, 62°30'W, 1550 m, subtropical humid forest, malaise trap/FIT, 2–30 Dec 1987, S.

& J. Peck; 3 males, 5 females, Campo Quijano, 30km E Salta, in forest remnant, dung trap, 18–20

Feb 1992, S.A. Marshall; 3 males, 3 females, Campo Quijano, 30km E Salta, sweep, dung, 20 Feb

1992, S.A. Marshall; 2 males, 1 female, Rosario de Lerma, INESALT yard, 24°59'S, 65°35'W,

malaise trap, 16–28 Feb 1992, S.A. Marshall; 7 males, 10 females, El Rey Nat. Pk., Pozo Verde

trail, km 7, 1000m, Yungas forest, malaise FIT, 5–15 Dec 1987, S. & J. Peck; Tierra del Fuego: 2 - 188 -

females, Ushuaia, 3 km E, marshy area near river, pan traps, 11–14 Feb 1992, S.A. Marshall.

ECUADOR: Napo: 4 males, Jatun Sacha Reserve, 6 km E Misahuallí, 1°4'S, 77°37'W, 450 m,

varzea, dung pans, 2–7 May 2002, M. Buck (DEBU, QCAZ).

Comments: There is some minor variation in the depth of the emargination on the male sternite 5 and in the shape of the surstylus, but this appears to be intraspecific variation .

Etymology: The species epithet is the Latin for bubble, referring to the small swelling at the tip of the male surstylus.

Archiceroptera ternum Paiero & Marshall, n. sp.

Figs. 6.48–6.49

Size: 2.1–3.1 mm.

Head: Ocellar triangle with 6–7 setae. Frons with 4–5 interfrontal seta and 4–5 inclinate orbital setulae. Eye:gena ratio = 1.7–2.0:1. Gena with 20–25 smaller setae.

Thorax: Acrostichal setulae in 8 rows between prescutellar dorsocentrals. Scutellum without setulae. Anterior katepisternal seta present, weak.

Legs: Fore femur with 5–7 posterodorsal and 5–6 posteroventral setae. Mid femur anteriorly with series of 9–11 stout setae extending from base to apical 3/4; apicoventral series of 5–7 setae on apical 1/4.; male, ventrally, with 15–17 robust setae in cluster on basal 1/3 with 4–5 in linear series along posterior margin immediately distal to basal cluster. Mid tibia with 2–3 median anterodorsal and 2–4 median posterodorsal setae; weak predistal dorsal, and weak distal posterodorsal setae present; male ventral comb with 11–12 robust setae on apical 2/3; midventral seta present only in female. Mid basitarsus with basal posteroventral setae distinctly longer than distal setae. Hind tibia simple (1–2 setae sometimes weakly strenghtened).

Wings: Costa with longer costagial setae not reaching humeral crossvein. C2:C3 ratio = 3:2.

Distance between r-m and dm-cu ~5.0× dm-cu. Cell dm with small CuA1 stub vein present.

Male abdomen (Fig. 6.48): Sternite 4 posteromedially entire. Sternite 5 posteromedially with large arcuate emargination bordered by pair of irregular process; processes each with clavate portion extending from corner of emargination and distal posterior portion that encompasses clavate portion - 189 - posteriorly; anterior portion of process heavily setose, posterior portion of process with 18–24 flattened setae along posterior margin. Transverse portion of sternite 6 arcuate (usually immediately below processes; right lateral portion recurved, simple. Synsternite 6+7 with narrowly rounded, medially projecting process; process with triangular sclerite distal to apex. Cercus triangular, elongate with long seta on basal 1/3, weakly narrowed near basal 1/3 and narrowly rounded distally

(appearing acute in posterior view due to weak apical twist of cercus). Surstylus (in lateral view) elongate conical with triangular anterior process; process slightly narrowed at midlength; posterior surface hirsute with 8–11 long setae. Postgonite with ventral surface rounded on basal 1/3, weakly sinuate near midlength; apically wedge-shaped. Distiphallus: dorsal sclerite not projecting distally; acrophallus dorsolaterally with broadly acute angle (not distinctly projecting).

Female Abdomen (Fig. 6.49): Tergite 7 posteromedially weakly emarginate; posterolaterally narrowly fused to tergite 8. Tergite 8 with lateral sclerites posterolaterally broadly rounded; medial sclerite elongate rectangular, anteriorly rounded. Epiproct with lateral sclerites elongate oval.

Cercus narrow, triangular. Sternite 7 posteriorly broadly rounded. Spermatheca spherical; sclerotized portion of duct ~1/3 length of spermatheca.

Type Material: Holotype (male, debu01085079, QCAZ) and 14 Paratypes (14 males; DEBU,

QCAZ): ECUADOR: Pichincha: Rio Palenque, dung, 25 Feb 1979, S.A. Marshall. Additional

Paratypes: BOLIVIA: La Paz: 2 males, Arroyo Tuhiri W Mapiri, 15°17'27"S, 68°15'29"W, 10

Apr 2001, S.A. Marshall; 1 female, Heath River Wildlife Centre, ~21 km SSW Puerto Heath,

12°40'S, 68°42'W, rainforest, malaise, 1–11 May 2007, S.M. Paiero; 2 males, Heath River Wildlife

Centre, ~21 km SSW Puerto Heath, 12°40'S, 68°42'W, tree fall, yellow pans, 5–9 May 2007, Paiero

& Kits (UASM); BRAZIL, São Paulo: 1 male, 1 female, USP Biology Station, human dung, 5–6

Feb 1979, R. Woodruff & J. Runnacles. COSTA RICA: Alajuela: 6 males, Florencia Forest, dung tp., 28 Feb 1980, H. Howden; 11 males, 11 females, Volcán Tenorio, N slope near Bijagua

Biological Station, 700 m, rainforest, RET over Atta mound, 16–20 Jun 2000, S.A. Marshall

(DEBU, INBC); Cartago: 4 males, Turrialba Catie, 600 m, 26 Feb 1980, H. & A. Howden; 4 males, Turrialba Catie, 600 m, 28 Feb 1980, H. & A. Howden; 3 males, Turrialba, Catie, Florence

Forest, 600m, cup traps, 28 Feb 1980, H. & A. Howden (INBC); Heredia: 3 males, 10km W Puerto - 190 -

Viejo, La Selva Verde, 2–4 Mar 1991, H. & A. Howden; 2 males, Braulio Carrillo National Park, El

Ceibo Biological Station, 400–600 m, Oct 1989, Aguilar & Zumbado; 3 females, La Selva

Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 14–17 Jun 2001, S.

Chatzimanolis; 3 males, La Selva Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°1'W, 80 m,

FIT, 18–20 Jun 2001, S. Chatzimanolis (INBC); 2 females, La Selva Biological Station, 3 km S

Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 20–23 Jun 2001, S. Chatzimanolis; 3 males, 6 females,

La Selva Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 2–5 Jun 2001, S.

Chatzimanolis; 3 males, 2 females, La Selva Biological Station, 3 km S Puerto Viejo, 10°26'N,

84°1'W, 80 m, FIT, 5–8 Jun 2001, S. Chatzimanolis; 1 male, La Selva Biological Station, 3 km S

Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 8–11 Jun 2001, S. Chatzimanolis (INBC); Limón: 1 male, Tortuguero National Park, Cuatro Esquinas, 0 m, Apr 1990, J. Solano; 3 males, Tortuguero

National Park, Cuatro Esquinas, 0 m, Jun 1990, J. Solano; 1 male, Tortuguero National Park, Cuatro

Esquinas, 0 m, Jun 1990, M. Barrelier; 3 males, Tortuguero National Park, Cuatro Esquinas, 0 m,

Jun 1990, U. Chavarría; Puntarenas: 1 male, Amistad National Park, Las Mellizas Biological

Station, Finca Cafrosa, 1300 m, Jun–Jul 1990, J.C. Saborio (INBC); 2 males, Corcovado National

Park, Sirena Biological Station, 0–100 m, Jun 1990, F. Quesada. ECUADOR: Esmeraldas: 2 males, La Chiquita, 11 km SE San Lorenzo, 5 m, carrion, 9–10 Jun 1975, S. Peck (QCAZ); 3 males,

La Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, 10–11 Jun 1975, S. Peck; 14 males, La

Chiquita, 11 km SE San Lorenzo, 5 m, dung trap, day 3, 8–9 Jun 1975, S. Peck (QCAZ); 13 males,

9 females, La Chiquita, 17km SE San Lorenzo, 5m, dung, 7–8 Jun 1975, S. Peck; Manabi: 1 male,

Chone, 20km N, 300m, cacao plantation, 2 dung traps, 6–9 Jun 1976, S. Peck; Napo: 1 male, Jatun

Sacha Reserve, 6 km E Misahuallí, 1°4'S, 77°37'W, 450 m, varzea, dung pans, 2–7 May 2002, M.

Buck; 3 males, 8 females, Tiputini Biodiversity Station, vicinity Yasuní National Park, 0°38'S,

76°0'W, human dung pitfalls, 14–19 Feb 1998, D.C. Darling (ROME); 1 male, Tiputini Biodiversity

Stn., 0°36'50"S, 76°9'1"W, sweep, May 2011, S.A. Marshall; 2 males, 1 female, Tiputini

Biodiversity Stn., 0°36'50"S, 76°9'1"W, May 2011, S.A. Marshall; 5 males, Tiputini Biodiversity

Stn., vicinity Yasuni National Park, 0°38'S, 76°10'W, pitfall trap (human dung), 14–19 Feb 1998,

D.C. Darling (QCAZ); Pichincha: 1 male, Allurquin, 28km E, Chiribaga Rd., 5200', moss forest, - 191 - carrion trap, 19–27 Jun 1975, S. Peck; 39 males, Palenque, day 3 trap, 24–25 Mar 1976, S. Peck; 2 males, Rio Palenque, carrion, 27 Feb 1979, S.A. Marshall; 10 males, Rio Palenque, dung, 27 Feb

1979, S.A. Marshall; 10 males, Rio Palenque, dung trap, 22–23 Feb 1976, S. Peck; 17 males, Rio

Palenque, dung trap, 25–26 Feb 1976, S. Peck (QCAZ); 4 males, Rio Palenque, J. Glasser trap, 26

Feb 1976, S. Peck; 1 male, Rio Palenque, 22 Feb 1976, S. Peck; 4 males, 4 females, Rio Palenque,

26 Feb 1976, J. Glaser; 3 males, Rio Palenque Stn., 47 km S Santo Domingo, 160m, 1° lowland rainforest, malaise head, 30 Apr–5 May 1987, Coote & Brown; 64 males, 1 female, Río Palenque

Stn., 47 km S Santo Domingo, traps 3–5, day 1, 22–23 Feb 1976, S. Peck; 3 males, Rio Palenque

Stn., 47km S Santo Domingo, 250m, dung, 17–25 Feb 1979, S.A. Marshall; 3 males, Rio Palenque

Stn., 47km S Santo Domingo, 250m, forest dung traps, 22–27 Feb 1976, S. Peck; 7 males, Rio

Palenque Stn., 47km S Santo Domingo, 250m, 17–25 Feb 1979, S.A. Marshall (QCAZ); 2 males,

Santo Domingo, 4km SE, 500m, 3 forest dung pans, 8–11 Jun 1976, S. Peck; 1 male, Tinalandia,

1120m, wet lower montane rainforest, Malaise head, 9–13 May 1987, L.D. Coote & B.V. Brown; 1 male, Tinalandia, 16 km SE Santo Domingo, 680 m, forest, dung traps 32, 16–28 Jun 1975, S. Peck;

3 males, Tinalandia, 16 km SE Santo Domingo, 680 m, rainforest, malaise-FIT, 4 May–25 Jul 1985,

S. & J. Peck (QCAZ); 12 males, Tinalandia, 16 km SE Santo Domingo, 680 m, dung trap, 21–22

Jun 1975, S. Peck. FRENCH GUIANA: St. Laurent du Maroni: 1 male, Maripasoula, Mitaraka, near base camp & along trails, tropical moist forest, SLAM trap, 1–6 Mar 2015, Touroult & Poirier

(MHNM). GUYANA: 7 males, 9 females, Kabocalli, Iwokrama Forest Reserve, 100 m, FIT, 22–25

May 2001, Brooks & Falin; 1 male, Kabocalli, Iwokrama Forest Reserve, 60 m, FIT, 3–5 Jun 2001,

Brooks & Falin. PANAMA: Chiriquí: 1 male, Hartmann Finca, 15 km NW Hato de Volcán, dung trap, 20–25 May 1977, S. Peck; 1 male, Hartmann's Finca, 1550 m, dung trap, 31 May 1977, S.

Peck; 1 male, Hartmann's Finca, 15 km NW Hato de Volcán, 1500m, dung, 20–25 May 1977, S.

Peck; 3 males, 4 females, Hartmann's Finca, 15 km NW Hato de Volcán, 1200 m, dung trap, 20–31

May 1977, S. Peck; 3 males, Lagunas, 5km SW Hato del Volcan, 1360m, dung, 22–26 May 1977,

S. Peck; 2 males, Lagunas, 5km SW Hato del Volcan, 1360m, 22 May 1977, S. Peck. PERU:

Madre de Dios: 3 males, Pantiacolla Lodge, Alto Madre de Dios River, 12°39'18"S, 71°13'54"W,

420 m, FIT, 14–19 Nov 2007, D. Brzoska (MUSM). VENEZUELA: Bolivar: 1 male, "20km", - 192 - evergreen dry forest, FIT, Jun 1987–12 Jul 1987, S. & J. Peck (MIZA); 3 males, 22km S El Dorado, lowland rainforest, FIT, 25 Jun–12 Jul 1987, S. & J. Peck (DEBU, MIZA).

Comments: This is a widespread species that has been collected at a variety of baits. A series from

Panama has an eye:genal height ratio of 1.7:1, much lower than other members of this species (2:1), but no other morphological differences suggest that the Panamanian material is distinct.

Etymology: The species epithet is from the Latin for pinch, referring to the appearance of the posterior margin of the male sternite 5.

Archiceroptera uncinata Paiero & Marshall n. sp.

Figs. 6.50–6.51

Size: 1.6–2.2 mm.

Head: Ocellar triangle with 2–3 setae. Frons with 5 interfrontal seta and 3–4 inclinate orbital setulae. Eye:gena ratio = 1:1. Gena with 12–14 smaller setae.

Thorax: Acrostichal setulae in 6 rows between prescutellar dorsocentrals. Scutellum without setulae near basal seta. Anterior katepisternal seta present, weak.

Legs: Fore femur with 7–8 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 7–9 stout setae extending from base to apical 3/4; apicoventral series of 4–5 setae on apical

1/4.; male, ventrally, with 9–10 robust setae in elongate cluster on basal cluster half. Mid tibia with

2–3 median anterodorsal and 2 median posterodorsal setae; weak predistal dorsal setae present; male ventral comb with 9–10 robust setae on apical 2/3; midventral seta present only in females.

Mid basitarsus with basal posteroventral setae slightly longer than distal setae. Hind tibia usually simple (1–2 preapical setae sometimes weakly developed).

Wings: Costa with costagial seta not reaching humeral crossvein. C2:C3 ratio = 3:2. Distance between r-m and dm-cu ~5.0× dm-cu. Cell dm with small CuA1 stub vein usually present.

Male Abdomen (Fig. 6.50): Sternite 4 posteromedially usually entire (rarely with weak, poorly defined shallow emargination). Sternite 5 posteromedially with broad arcuate emargination; emargination ~3/4 width and 2/5 length of sternite. Transverse portion of sternite 6 strongly arcuate with broad swelling on left side of arc; right lateral portion curved, simple. Synsternite 6+7 - 193 - withmedially projecting process narrowly rounded, with small apical sclerite (sclerite closely approximated with swelling on left side of actuate transverse portion of sternite 6). Cercus triangular, elongate, with abrupt constriction and elongate seta near basal 1/4. Surstylus (in lateral view) elongate with anterobasally projecting process; posterior surface with 1–2 strong setae and 9–

12 smaller setae. Postgonite basally quadrate, with apical ½ acuminate; ventral surface weakly sinuate near middle. Distiphallus: dorsal sclerite with apical 1/8 extending beyond apex of acrophallus; acrophallus dorsolaterally with obtuse angle near midlength of distiphallus

Female Abdomen (Fig. 6.51): Tergite 7 posteromedially weakly sinuate; posterolaterally narrowly fused to tergite 8. Tergite 8 with lateral sclerites broadly rounded; medial sclerite elongate triangular. Epiproct with lateral sclerites ovoid. Cercus elongate-ovoid. Sternite 7 with posterior margin narrowly rounded. Spermatheca ovoid, slightly flattened, with swelling at duct junction; sclerotized portion of duct ~1/4 length of spermatheca.

Type Material: Holotype (male, debu01084664, MIZA) + 16 Paratypes (8 males, 8 females;

DEBU, MIZA): VENEZUELA: Bolivar: km40 Sta. Elena Icabaru Road, 100m, 4 Aug 1986, B.D.

Gill. Additional Paratypes: BOLIVIA: La Paz: 9 males, Arroyo Tuhiri W Mapiri, 15°17'27"S,

68°15'29"W, 10 Apr 2001, S.A. Marshall; 1 male, Caranavi, ca. 10 km NW, road to ENTEL tower,

1700 m, bamboo, dung pans, 13 Apr 2001, S.A. Marshall (UASC); 2 males, Heath River Wildlife

Centre, ~21 km SSW Puerto Heath, 12°40'S, 68°42'W, tree fall, yellow pans, 5–9 May 2007, Paiero

& Kits (UASC); 3 males, 6 females, San Antonio, ca. 8 km S Mapiri, 15°20'56"S, 68°13'31"W, secondary forest, dung pans, 11 Apr 2001, S.A. Marshall. COLOMBIA: Amazonas: 5 males, 3 females, Leticia, pepper farm, dung, 1 Mar 1974, V. Nealis; 2 males, 3 females, Leticia, dung traps,

28 Feb 1974, V. Nealis; 2 males, 14 females, Leticia, 28 Feb 1974, V. Nealis. ECUADOR: Napo:

1 male, Tena, 12 km SW, 500m, dung trap 30–33, day 2–5, 8–11 Jul 1976; 1 male, Yasuní National

Park, Yasuní Research Station, 0°38'S, 76°36'W, rainforest, malaise trap, 3–20 Nov 1998, Pape &

Viklund; 1 male, Tiputini Biodiversity Stn., vicinity Yasuni National Park, 0°38'S, 76°10'W, pitfall trap (human dung), 14–19 Feb 1998, D.C. Darling (ROME). FRENCH GUIANA: St. Laurent du

Maroni: 1 male, Maripasoula, Mitaraka, MIT-DZ-RBF1, 2°14'4"N, 54°27'2"W, 270m, tropical wet forest (bas fond), blue pan traps, 26 Feb–2 Mar 2015, M. Pollet (MHNM). GUYANA: Potaro- - 194 -

Siparuni Distr.: 1 male, Mount Wokomung, 5°6'35"N, 59°49'15"W, 1234m, 1° rainforest, human dung, pitfall trap, 27 Oct–1 Nov 2004, B. Hubley (ROME); 5 males, Mount Wokomung, 5°7'53"N,

59°48'31"W, 698m, 1° forest, pitfall trap (human dung), 21–26 Oct 2004, B. Hubley (ROME); 1 male, Potaro River vic., 5°9'56"N, 59°46'43"W, 680m, 1° rainforest, human dung, pitfall trap, 19–

20 Oct 2004, B. Hubley; Rupununi Distr.: 1 female, Kurupukari, E side Essequibo River, 200', 1° forest edge/field, malaise, 11–16 Oct 1990, B. Hubley; 3 males, 1 female, Kurupukari, Essequibo

River, 200', 1° forest, dung traps, 9 Oct 1990, B. Hubley; (distr. unknown): 4 males, Kabocalli,

Iwokrama Forest Reserve, 100 m, FIT, 22–25 May 2001, Brooks & Falin; 7 males, Kabocalli,

Iwokrama Forest Reserve, 60 m, FIT, 3–5 Jun 2001, Brooks & Falin. PERU: Loreto: 4 males,

Campamento San Jacinto, FIT, 12 Jul 1993, R. Leschen; 1 male, Campamento San Jacinto, 175–215 m, FIT, 5 Jul 1993, R. Leschen; 1 male, Campamento San Jacinto, 175–215 m, FIT #44, 7 Jul 1993,

R. Leschen (MUSM); 1 male, Campamento San Jacinto, 175–215 m, FIT #84, 11 Jul 1993, R.

Leschen (MUSM); 2 males, Campamento San Jacinto, 175–215 m, FIT, #67, 9 Jul 1993, R. Leschen

(MUSM); 8 males, 3 females, Teniente López, Riv. forest, FIT, #211, 26 Jul 1993, R. Leschen; 3 males, Teniente López, riverine forest, FIT, #199, 24 Jul 1993, R. Leschen; 6 males, Teniente

López, FIT, 23 Jul 1993, R. Leschen; 3 males, Teniente López, FIT, 26 Jul 1993, R. Leschen

(MUSM); 6 males, Teniente López, 1.5 km N, 2°31'S, 76°10'W, 230–305 m, FIT, 18 Jul 1993, R.

Leschen; 2 males, Teniente López, 1.5 km N, 2°31'S, 76°10'W, 230–305 m, FIT, 20 Jul 1993, R.

Leschen (MUSM); 7 males, Teniente López, 1.5 km N, 2°31'S, 76°10'W, 230–305 m, FIT, 22 Jul

1993, R. Leschen; Madre de Dios: 1 male, Amazonas Lodge, N of Atalaya, 12°52'12"S,

71°22'36"W, 480 m, FIT, 10–13 Nov 2007, D. Brzoska; 2 males, CIRCA Field Stn., trail 6, research plot, 12°33'7"S, 70°6'35"W, 295m, Malaise trap, 9–11 Jun 2011; 2 males, Zona Reserva Manu,

Pakitza, 11°57'S, 71°17'W, 400 m, malaise trap 3, 18–23 Feb 1992, B. Brown & D. Feener; 2 males,

Zona Reserva Manu, Pakitza, 11°57'S, 71°17'W, 400 m, Malaise trap 3, 4–9 Mar 1992, Brown &

Feener. TRINIDAD & TOBAGO: Trinidad: 13 males, 8 females, St. George Co., Arima Ward,

NY Zoological Society Station, pig dung trap, 11 Jun 1927, R. Woodruff. VENEZUELA: Bolivar:

1 female, 105km S El Dorado, 350m, 17 Jul–7 Aug 1986, B. Gill; 15 males, 19 females, 10km S El

Dorado, 200m, 17 Jul–7 Aug 1986, B.D. Gill; 2 males, 12 females, 20 km S El Dorado, 220 m, 20– - 195 -

23 Jul 1986, B. Gill (MIZA); 26 males, 22km S El Dorado, lowland rainforest, FIT, 25 Jun–12 Jul

1987, S. & J. Peck; 14 males, 18 females, 33km S El Dorado, 220m, 2–7 Aug 1986, B.D. Gill; 14 males, 4 females, km40 Sta. Elena Icabaru Rd., 220m, 4–6 Aug 1986, B.D. Gill; 6 males, 1 female, km40 Sta. Elena Icabaru Road, 1000m, 4–6 Aug 1986, B.D. Gill;

Dubious Record: U.S.A.: SC: 1 male, Colleton Co., Colleton State Park, pig dung trap, 27 Sep–5

Oct 1983, R.E. Woodruff.

Etymology: The species epithet is the Latin for barb, in reference to the basally projecting process on the male surstylus.

6.7.4 Archiceroptera brevivilla species subgroup

The A. brevivilla species subgroup includes five newly described species that form a derived group within the A. ternum species group. Members of this group can be recognized within Archiceroptera by their reduced mid tibial chaetotaxy; the mid tibia has only 1 basal anterodorsal seta and 1 anterodorsal seta near the midlength (no posterodorsal setae are present on the basal 2/3).

Description:

Head: Frons with 4 pairs of interfrontal setae. Gena with 1–2 strong anterior seta(e) and additional smaller posterior setae.

Thorax: Scutum with 1 prescutellar dorsocentral seta. Acrostichal setae in 4–6 rows between prescutellar dorsocentral setae. Anterior katepisternal seta weak (< 1/3 posterior katepisternal seta).

Scutellum without setulae.

Legs: Mid tibia with 1 anterodorsal seta on basal 1/3, 1 anterodorsal seta near midlength, strong distal anterodorsal and distal dorsal setae; predistal dorsal seta sometimes present; male with distinct ventral comb of dark setae on apical 1/3-1/2; midventral seta present only in female. Mid basitarsus with basal posteroventral seta longer than distal setae.Hind tibia without distinct dorsal setae.

Wings: Costa with 1 strong costagial seta. Cell dm with CuA1 stub vein present or absent.

Male abdomen: Sternite 4 posteriorly simple. Sternite 5 posterior margin variable (usually tab-like); posteromedially with cluster(s) of setae. Transverse portion of sternite 6 straight. Surstylus - 196 - anteriorly with narrow process. Cercus basally ovoid or triangular in basal ¼–1/3 , apical 2/3-3/4 narrower; basal part with long seta. Distiphallus with well-developed membranous acrophallus.

Female abdomen: Tergite 7 posteromedially entire; posterolaterally fused to anterolateral corner tergite 8. Tergite 8 tripartite with medial sclerite elongate, rectangular and strongly sclerotized.

Epiproct medially desclerotized into pair of sclerites; with scattered setulae. Cercus ovoid/droplet- shaped with 1 strong apical flattened seta; preapical seta (weaker and shorter than apical seta) adjacent or appressed to apical seta. Sternite 8 divided into pair of lateral triangular sclerites; sclerites narrowly meeting medially. Spermatheca shape variable; paired spermathecae with stems short (< 1/4 spermathecal length)

Archiceroptera braziliensis Paiero & Marshall, n. sp.

Figs. 6.52–6.53

Size: 1.7–1.9 mm.

Head: Frons with 1–2 inclinate orbital setulae. Eye:gena ratio = 20:9. Gena with 2 enlarged setae and 7–8 smaller setae.

Thorax: Acrostichal setae in 4 rows between prescutellar dorsocentrals.

Legs: Fore femur with 5–6 posterodorsal and 3–4 posteroventral setae. Mid femur anteriorly with series of 5–6 strong setae extending from base to apical 3/4; apicoventral series of 3 setae on apical

1/4.; male, ventrally with 8–10 robust setae in elongate cluster on basal 1/4. Mid tibia with weak predistal dorsal seta;ale ventral comb with series of 13–15 robust setae on apical ¾.

Wings: Costa with costagial seta not reaching humeral crossvein. C2:C3 ratio = 3:2. Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm with weak CuA1 stub vein, or absent.

Male Abdomen (Fig. 6.52): Sternite 5 posteromedially entire, with 8–10 dark, robust setae. Cercus basal ¼ ovoid, apical 3/4 narrow and acute. Surstylus (in lateral view) anterior process elongate, curved near apical 1/3, and with acute distal angle; surface setose with 3–4 setae on posterior surface. Postgonite with basal ½ quadrate (ventral surface somewhat rounded) and apical half acute, narrow. Distiphallus:dorsal sclerite elongate, extending beyond apex of acrophallus; lateral acrophallus projecting, reflexed and apically rounded. - 197 -

Female Abdomen (Fig. 6.53): Tergite 7 posterolaterally broadly fused with tergite 8. Tergite 8 with lateral sclerites posteriorly roundedEpiproct with lateral sclerites transverse. Sternite 7 posteromedially broadly emarginate, with pair of teeth near middle. Spermatheca barrel-shaped with short narrow invaginations on both ends; swollen near junction with duct; sclerotized portion of duct short, < 1/2 spermathecal length; paired spermathecae with stems abbreviated, short.

Type Material: Holotype (male, debu001088408, MZSP) + 2 Paratypes (2 males; MZSP):

BRAZIL: Paraná: Curitiba, 30 km SE, BR 277, dung traps, 6–9 Feb 1990, S.A. Marshall.

Additional Paratypes: BRAZIL: São Paulo: 2 males 7 females, USP Biology Station, human dung, 5–6 Feb 1979, R. Woodruff & J. Runnacles (DEBU, MZSP).

Comments: This species is currently known only from the Atlantic Forest of Brazil.

Etymology: The species epithet refers to the only country from which this species is known.

Archiceroptera brevivilla Paiero & Marshall, n. sp.

Figs. 6.54–6.55

Size: 1.5–1.9 mm.

Head: Frons with 2–3 inclinate orbital setulae. Eye:gena ratio = 2:1. Gena with 2 enlarged setae and

8–9 smaller setae.

Thorax: Acrostichal setae in 6 rows between prescutellar dorsocentrals.

Legs: Fore femur with 2–4 posterodorsal and 3–5 posteroventral setae. Mid femur anteriorly with series of 7–8 stout setae extending from base to apical 3/4; apicoventral series of 2–3 setae on apical

1/4; male, ventrally, with 5–7 robust setae in elongate cluster on basal 1/3. Mid tibia without predistal dorsal seta; male ventral comb with 7–8 robust setae on apical ½.

Wings: Costa with costagial seta not reaching humeral crossvein. C2:C3 ratio = 2:1. Distance between r-m and dm-cu ~3.0× dm-cu. Cell dm usually with small CuA1 stub vein present.

Male Abdomen (Fig. 6.54): Sternite 5 posteromedially with broad tab (~1/3 sternite width); tab with 21–25 pale flattened setae apically and with 2 dense clusters of 20–25 setae extending from tab to disc of sternite. Cercus basal 1\3 ovoid, apical 3/4 narrow and acute. Surstylus (in lateral view) with short, finger-like anterior process; apically broadly angled; surface largely setose with 4–5 - 198 - setae on posterior surface. Postgonite with basal ½ quadrate (ventral surface somewhat rounded) and apical half acute, narrow. Distiphallus: dorsal sclerite long, with apical 1/3 extending beyond apex; acrophallus dorsally with rounded, reflexed process and ventrally with rounded elongate projection.

Female Abdomen (Fig. 6.55): Tergite 7 posterolaterally broadly fused with tergite 8. Tergite 8 with lateral sclerites with distinct posterolateral process (~1/3 length of tergite). Epiproct with lateral sclerites triangular. Sternite 7 posteromedially acutely rounded. Spermatheca flattened ovoid, with small swelling near duct junction; sclerotized portion of duct < 1/4 spermathecal length); paired spermathecae with stems abbreviated, short (< 1/4 spermathecal length).

Type Material: Holotype (male, debu01088247, FMNH) + 67 Paratypes (39 males, 28 females;

DEBU, FMNH, UNAM): MEXICO: Hidalgo: Tlanchinol, 2.5mi N, 5200', cloud forest, dung, 6–

11 Jul 1973, A. Newton. Additional Paratypes: GUATEMALA: Baja Verapaz: 1 female,

Purulhá, 7.4 km S, 1650 m, FIT, #176, 2 Jul 1993, Ashe & Brooks; 1 male, Purulhá, 7.4 km S, 1650 m, FIT, #189, 2–3 Jul 1993, Ashe & Brooks; Izabal: 1 female, Izabal, 350 m, malaise trap, 14 Dec

1986, M.J. Sharkey (UVGC); Sacatepéq: 1 female, Antigua, 5 km SE, 14°31'43"N, 90°41'20"W,

2330 m, oak forest, malaise trap, 10–13 Jun 2009 (UVGC). MEXICO: Chiapas: 1 female, Lagunas de Montebello Parque Nacional, Aqua Tinta, 4900', oak-pine, human dung, 21–24 Aug 1971, A.

Newton; 1 female, Teopisca, 5 mi NW, 16°34'48"N, 092°31'12"W, 6600', oak-pine-juniper woodland, human dung, 21–24 Aug 1971, A. Newton; 1 male, San Cristóbal de las Casas, dung traps, 26–28 May 1990, B. Gill; 2 males, Bochil, 22mi. N, 5600 ft, pine, oak, liquidamber, human dung, 18–24 Aug 1971, A. Newton; Hidalgo: 1 male, Rancho Viejo, 10mi. NE, 5100', dung, 23–29

Jun 1971, A. Newton; 2 males, 6 females, Tenango de Doria, 7mi SW, 7000', cloud oak forest, human dung, 2–6 Jul 1971, A. Newton; 52 males, 56 females, Tlanchinol, 3.5mi N, 5100', cloud forest, human dung, 6–11 Jul 1973, A. Newton (DEBU, FMNH, UNAM); México: 1 female, Santa

Marta, 5 mi E, km 8.5, 10100 ft, fir forest, human dung, 29 Aug–4 Sep 1971, A. Newton; Puebla:

21 males, 20 females, Huanchinango, 5mi W, 6000', hardwood-pine, human dung, 3–7 Jul 1971, A.

Newton (DEBU, FMNH, UNAM); 8 males, 9 females, Teziutlan, 4.5mi E, 5000', cloud forest, human dung, 10–14 Jul 1971, A. Newton; San Luis Potosi: 1 female, Xilitla, 40km W, 1700m, - 199 - pine-oak forest, FIT, 12 Jun–6 Aug 1983, S. & J. Peck; 7 males, 14 females, Xilitla, 14mi. W, 1800',

Liquidambar forest, dung, 20–28 Jun 1971, A. Newton; Veracruz: 1 female, Catemaco, 6mi. NE,

1500', rainforest, human dung, 30 Jul–8 Aug 1970, A. Newton; 1 female, Teocelo, 10mi SW, 4400', wet oak forest, human dung, 11–16 Jul 1971, A. Newton; 5 males, 8 females, Teocelo, 10mi SW,

4400', oak, wet, human dung, 11 Jul 1971, A. Newton

Etymology: The species epithet refers to the short finger-like projection on the male surstylus.

Archiceroptera curvavilla Paiero & Marshall, n. sp.

Figs. 6.56–6.57

Size: 1.2–1.9 mm.

Head: Frons with 3–4 inclinate orbital setulae. Eye:gena ratio = 2:1. Gena with 1 long setae seta and 6–7 smaller setae.

Thorax: Acrostichal setae in 4 rows between prescutellar dorsocentral setae.

Legs: Fore femur with 5 posterodorsal and 2–3 posteroventral setae. Mid femur anteriorly with series of 10–11 robust setae extending from base to apical 3/4; apicoventral series of 3 setae on apical 1/4; male, ventrally, with 8–10 robust setae in elongate cluster on basal 1/4. Mid tibia without predistal dorsal setae; male ventral comb with 8–11 robust setae on apical ½.

Wings: Costa with costagial seta extending to humeral crossvein. C2:C3 ratio = 4:3. Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm with CuA1 stub vein usually present.

Male Abdomen (Fig. 6.56): Sternite 5 posteromedially produced on medial 1/3 as pair of broad rounded tabs, each with 10–12 pale flattened setae on margin. Cercus with basal 1\3 triangular- ovoid, narrowing to acute apex. Surstylus (in lateral view) with a short, lateraly curved process on anterior surface; lateral surface setose with 10–12 setae present. Postgonite with basal ½ quadrate

(ventral surface somewhat rounded) and apical half acute, narrow. Distiphallus: dorsal sclerite long, with apical 1/3 extending beyond apex; acrophallus dorsally with narrowly rounded process, ventrally with rounded elongate process.

Female Abdomen (Fig. 6.57): Tergite 7 posterolaterally broadly fused with tergite 8. Tergite 8 with lateral sclerites posterolaterally acute. Epiproct with lateral sclerites transverse, anteriorly rounded. - 200 -

Posterior magin of sternite 7 acutely rounded. Spermatheca ovoid with swelling near duct junction; sclerotized portion of duct < 1/3 length of spermatheca); paired spermathecae with stems obsolete or extremely abbreviated.

Type Material: Holotype (male, debu01088461, QCAZ) + 5 Paratypes (3 males, 2 females):

ECUADOR: Pichincha: Rio Palenque, dung trap, 22–23 Feb 1976, S. Peck. Additional

Paratypes: COSTA RICA: Limon, 1 male, Guapiles, 16km W, 400m, 19 Apr 1989, P. Hanson;

Puntarenas, 1 male, Las Tablas, ENE Las Mellizas, 15km ENE San Vito, 28 May 1987, A.

Norrbom; San José, 1 male, Braulio Carillo National Park, 9.5km E tunnel, 1000 m, Malaise trap,

Apr 1989, P. Hanson (INBC). ECUADOR: Emeraldas, 1 male, La Chiquita, 11 km SE San

Lorenzo, 5 m, carrion, 9–10 Jun 1975, S. Peck; Pichincha, 1 female, Río Palenque Stn., 27 km S

Santo Domingo, 250 m, 17–25 Feb 1979; 1 male, Río Palenque Stn., 47 km S Santo Domingo, 26–

27 May 1975, S. Peck; 1 male, 1 female, Rio Palenque, dung, 25 Feb 1979, S.A. Marshall (QCAZ);

1 male, 1 female, Rio Palenque Stn., 47 km S Santo Domingo, 160m, 1° lowland rainforest, malaise head, 30 Apr–5 May 1987, Coote & Brown; 1 male, 1 female, Río Palenque Stn., 47 km S Santo

Domingo, traps 3–5, day 1, 22–23 Feb 1976, S. Peck (QCAZ); 2 males, Rio Palenque, J. Glasser trap, 26 Feb 1976, S. Peck; 2 males, 2 females, Rio Palenque, dung trap, 25–26 Feb 1976, S. Peck

(CNCI); 4 males, Rio Palenque Stn., 47km S Santo Domingo, 250m, 17–25 Feb 1979, S.A.

Marshall (QCAZ); 5 males, 6 females, Tinalandia, 1120m, wet lower montane rainforest, Malaise head, 9–13 May 1987, L.D. Coote & B.V. Brown; 7 females, Tinalandia, 680m, dung, 22–28 Jun

1975, S. Peck. PANAMA: Chiriqui, 1 female, Cerro Punta, 27km W, 1260m, 2 Jun 1977, S. Peck;

21 males, Clara, 2km NS(?), 1500m, 20–25 May 1977, S. Peck; 1 male, Hartmann's Finca, 15 km

NW Hato de Volcán, 1500m, dung, 20–25 May 1977, S. Peck; 2 males, Cerro Punta, 27 km W,

1700 m, carrion, 5 Jun 1977, S. Peck; 2 males, Hartmann's Finca, 1700 m, 28 Jun–3 Jul 1981, B.

Gill; 4 males, 1 female, Hartmann's Finca, 15 km NW Hato de Volcán, 1200 m, dung trap, 20–31

May 1977, S. Peck; 9 males, 1 female, Hartmann's Finca, 15 km NW Hato de Volcán, 1500m, carrion trap, 20–31 May 1977, S. Peck.

Etymology: The species epithet refers to the curved finger-like process on the anterior surface of the male surstylus. - 201 -

Archiceroptera llama Paiero & Marshall, n. sp.

Figs. 6.58–6.59

Size: 1.5–2.2 mm.

Head: Frons with 3–4 inclinate orbital setulae. Eye:gena ratio = 2:1. Gena with 2 enlarged setae and

9–10 smaller setae present.

Thorax: Acrostichal setae in 6 rows between prescutellar dorsocentral setae.

Legs: Fore femur with 3–5 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 7–8 stout setae extending from base to apical 3/4; apicoventral series of 2–3 setae on apical

1/4.; male, ventrally, with 6–8 robust setae along posterior margin, extending to midlength. Mid tibia without predistal dorsal seta; male ventral comb with 11–13 robust dark setae on distal ¾.

Wings: Costa with costagial seta extending to humeral crossvein. C2:C3 ratio = 2:1. Distance between r-m and dm-cu ~5.0× dm-cu. Cell dm with short CuA1 stub vein usually present.

Male Abdomen (Fig. 6.58): Sternite 5 posteromedially with broad rectangular emargination; emargination with 2 poorly sclerotized, lateral ovoid ‘pads’ extending out from sides; ‘pads’ with small pale setae along posterior margin. Cercus base triangular, narrowing slightly at basal 1/3 and compressed laterally; . Surstylus (in lateral view) with 2 small finger-like anterior processes; posterior half setose with numerous long setae present. Postgonite with basal ½ quadrate (ventral surface somewhat rounded) and apical half acute, narrow. Distiphallus: dorsal sclerite not projecting beyond apex of acrophallus; lateral acrophallus with narrowly rounded projections curved dorsally.

Female Abdomen (Fig. 6.59): Tergite 7 posterolaterally narrowly fused with tergite 8. Tergite 8 with lateral sclerites posterolaterally narrowly rounded, almost acutely angled Epiproct with lateral sclerites broadly rounded anterolaterally. Sternite 7 posteromedially acutely rounded. Spermathecae triangular, slightly flattened, with distinct swelling at duct junction; sclerotized portions of duct ~1/2 length of spermatheca; paired spermathecae with stems greatly abbreviated or obsolete.

Type Material: Holotype (male, debu00386753, UVGC) + 14 Paratypes (3 males, 11 females;

DEBU, UVGC): GUATEMALA: Sacatepéquez: Volcán Atitlán, Ref. Quetzal, 14°33'2"N,

91°11'32"W, 1670m, cloud forest, FIT, 13–16 Jun 2015, Falin & Carrillo. Additional Paratypes: - 202 -

GUATEMALA: Sacatepéquez: 10 males, 25 females, Volcán Atitlán, Ref. Quetzal, 14°33'2"N,

91°11'32"W, 1670m, cloud forest, FIT, 3–6 Jun 2015, Falin & Carrillo (DEBU, UVGC).

MEXICO: Chiapas: 1 male, Custepac, 2 km SE, 15°43'15"N, 92°57'2"W, 1510 m, mesophil forest, malaise trap, 17 May 2008, (no collector indicated) (UNAM).

Etymology: The species epithet is from the acronym for the Leaf Litter Arthropods of Meso-

America (LLAMA) project, which yielded all of the type material from Guatemala.

Archiceroptera megavilla Paiero & Marshall, n. sp.

Figs. 6.60–6.61

Size: 1.5–2.1 mm.

Head: Frons with 2–3 inclinate orbital setulae. Eye:gena ratio = 2:1. Gena with 2 enlarged setae and

6–7 smaller setae.

Thorax: Acrostichal setae in 4 rows between prescutellar dorsocentral setae.

Legs: Fore femur with 3–5 posterodorsal and 4–5 posteroventral setae. Mid femur anteriorly with series of 7–8 stout setae extending from base to apical 3/4; apicoventral series of 2–3 setae on apical

1/4.; male, ventrally, with 6–8 robust setae posteriorly on basal 1/2. Mid tibia without predistal dorsal seta; male ventral comb with 13–14 dark, robust setae on apical ¾.

Wings: Costa with costagial seta extending to humeral crossvein. C2:C3 ratio = 5:3.5. Distance between r-m and dm-cu ~4.0× dm-cu. Cell dm with distinct CuA1 stub vein present.

Male Abdomen (Fig. 6.60): Sternite 5 posteromedially sinuate, with 16–18 pale setae on posterior margin and discal setation denser anterobasal to medial emargination. Cercus basal ¼ ovoid, apical

3/4 narrow and acute. Surstylus (in lateral view) with long, finger-like anterior process; apically broadly angled; surface largely setose with 3–5 long setae on posterior surface. Postgonite with basal ½ quadrate (ventral surface somewhat rounded) and apical half acute, narrow. Distiphallus: dorsal sclerite long, with apical 1/3 extending beyon apex; acrophallus dorsally with rounded, reflexed process and ventrally with rounded elongate projection.

Female Abdomen (Fig. 6.61): Tergite 7 posterolaterally broadly fused to tergite 8. Tergite 8 with lateral sclerites posterolaterally projecting, narrowly rounded posteriorly; Epiproct with sides - 203 - transverse. Sternite 7 with posterior margin acutely rounded. Spermathecae ovoid, with small swelling at junction with duct; sclerotized portion of duct < 1/4 length of spermatheca; paired spermathecae with stems obsolete or extremely abbreviated.

Type Material: Holotype (male, debu01088606) + 18 Paratypes (18 males): PANAMA:

Chiriquí: Cerro Punta, 2 km W, 1760 m, dung, 5 Jun 1977, S. Peck. Additional Paratypes:

ARGENTINA: Salta: 1 male, Camino de la Cornisa, 40 km N Salta, roadside forest, sweep, 29 Feb

1992, S.A. Marshall; 6 males, 9 females, Campo Quijano, 30km E Salta, dung trap in forest remnant, 18–20 Feb 1992, S.A. Marshall; 1 female, Campo Quijano, 30km E Salta, El Alisa forest remnant, FIT, 18–20 Feb 1992, S.A. Marshall; 2 female, Cañada La Gotera, 15 km W Chicoana, sweeping wet litter, 19 Feb 1992, S.A. Marshall; 1 male, El Ucumar, 22km N La Caldera, 1550m, subtropical humid forest, malaise FIT, 2–30 Dec 1987, S. & J. Peck. COSTA RICA: Cartago: 4 males, 2 females, Tapantí National Park, Ranger Stn., 1200 m, human dung, hand & traps, 9–12 Oct

1999, Buck & Marshall (INBC); Guanacaste: 1 male, 4 females, Cacao Field Station, 1250m, dung trap, 12–15 Feb 1996, S.A. Marshall; Heredia: 1 male, La Selva Biological Station, 3 km S Puerto

Viejo, 10°26'N, 84°1'W, 80 m, FIT, 14–17 Jun 2001, S. Chatzimanolis (INBC); 1 female, La Selva

Biological Station, 3 km S Puerto Viejo, 10°26'N, 84°1'W, 80 m, FIT, 20–23 Jun 2001, S.

Chatzimanolis (INBC); Puntarenas: 5 males, 3 females, (probably from Las Alturas), 1700m, dung trap, 12–13 Aug 1995, S.A. Marshall; 1 female, Coto Brus, Z.P. Las Tablas, Estacion Biologica Las

Alturas, 8°57'7”N, 82°50'4”W, 1500–1600 m, ZADBI-1271, #107950, Malaise trap, 13–20 Aug

2013, ZADBI (INBC); 2 males, 3 females, Las Alturas, 8°57'N, 82°58'W, 1600 m, ground, Eciton raid, 15 Aug 1995, S.A. Marshall; 3 males, Las Alturas, 2000 m, tree fall, dung piles, 12–15 Aug

1995, S.A. Marshall; 1 male, 1 female, Las Alturas Biological Station, 1550 m, 17 Aug 1995, T.

Pape; 1 male, Monteverde, 1500m, cloud forest, dung traps, 19–25 Aug 1993, E.R. Barr; 5 males,

Monteverde, 1520 m, FIT, 11–18 Jun 1983, D.H. Lindeman; 2 males, 2 females, Monteverde, 1520 m, FIT, 15–23 Jul 1983, D.H. Lindeman (INBC); 1 male, 2 females, Monteverde, 1520 m, FIT, 23–

30 Jul 1983, D.H. Lindeman; 6 females, Monteverde, 1560 m, dung trap, 11–18 Jun 1983, D.H.

Lindeman; 1 female, Monteverde, 1700–1800 m, yellow pan trap, 23–27 Feb 1991, B.J. Sinclair; 3 males, 3 females, Monteverde, 3 dung traps, 27 Feb 1991, H. & A. Howden; 1 female, Monteverde - 204 -

Biological Reserve, 1500 m, cloud forest, sweeping, 11 Jun 2000, M. Buck; 1 male, 1 female,

Monteverde Biological Reserve, 1500 m, cloud forest, 11–13 Jun 2000, S.A. Marshall; 1 female,

Osa Peninsula, Rincón, 2.5 km S, 8°42'1”N, 83°30'50”W, ~50 m, secondary forest, fish pitfalls, 10–

11 Aug 2001, ; San Jose: 1 male, Zurqui de Moravia, 10°2'58”N, 84°0'57”W, 1600m, Creek 2 north, Malaise trap, ZADBI-711 #106716, 29 Nov–7 Dec 2012, ZADBI (INBC); 2 females, Zurqui de Moravia, 10°2'58”N, 84°0'57”W, 1600m, Tower path, ZADBI-1136, #107741, Malaise trap #1,

6–13 Sep 2013, ZADBI (INBC); 1 female, Zurqui de Moravia, 10°2'58”N, 84°0'57”W, 1600m,

Tower path, ZADBI-1280 #107959, Malaise trap #1, 11–18 Oct 2013, ZADBI (INBC); 1 female,

Zurqui de Moravia, 10°2'58”N, 84°0'57”W, 1600m, Tower path, ZADBI-785, #106841, Malaise trap #1, 24–30 May 2013, ZADBI (INBC); 1 female, Zurqui de Moravia, 10°2'58”N, 84°0'57”W,

1600m, Tower path, ZADBI-894, #107115, Malaise trap #1, 24–31 May 2013, ZADBI (INBC); 1 female, Zurqui de Moravia, 10°2'58”N, 84°0'57”W, 1600m, Tower path, ZADBI-949, #107268,

Malaise trap #1, 12–19 Jul 2013, ZADBI; 1 female, Zurqui de Moravia, 10°2'58”N, 84°0'57”W,

1600m, ZADBI-5, #104990, Malaise trap #2, 12 Sep 2012, ZADBI; 1 male, Zurqui de Moravia,

10°2'58”N, 84°0'57”W, 1600m, Malaise trap #1, ZADBI-3 #104975, 12 Sep 2012, ZADBI (INBC);

1 male, Zurqui de Moravia, 10°2'58”N, 84°0'57”W, 1600m, Malaise trap #2, ZADBI-176, #105285,

9–19 Oct 2012, ZADBI. ECUADOR: Emeraldas: 1 female, La Chiquita, 11 km SE San Lorenzo,

5 m, dung, 9–10 Jun 1975, S. Peck; Napo: 2 males, 2 females, Baeza, 5 Mar 1979, S.A. Marshall; 1 male, 1 female, Baeza, 17 km NE, 1400 m, carrion trap, 3–6 Mar 1976, S. Peck; Pichincha: 17 males, 17 females, Alluriquin, 23km E, Chiriboyo Ret., 4600', dung, 19–27 Jun 1975, S. Peck

(CNCI, DEBU, QCAZ); 2 males, 1 female, Allurquin, 28km E, Chiribaga Rd., 5200', moss forest, carrion trap, 19–27 Jun 1975, S. Peck; 2 females, Nanegalito, 7 km SE, trout farm 'San José',

0°3'54”S, 78°40'36”W, 1500 m, river edge, pan traps, 30–31 Oct 1999, S.A. Marshall; 1 female,

Tandapi, 22.7km E, 8000 ft, mossy forest, dung trap, 24–29 Jun 1975, S. Peck; 1 male, Tinalandia,

16 km SE Santo Domingo, 680 m, dung trap, 21–22 Jun 1975, S. Peck. MEXICO: Chiapas: 1 female, Custepec, 3 km SE, 15°42'52”N, 92°56'17”W, 50 m, 1740 m, 2° mesophil forest, malaise trap, 17–20 May 2008 (UNAM). PANAMA: Chiriquí: 1 male, Cerro Punta, 27km W, 1260m, 2

Jun 1977, S. Peck; 3 females, Clara, 2km NS(?), 1500m, 20–25 May 1977, S. Peck; 7 males, 1 - 205 - female, Cerro Punta, 2 km E, 1760 m, Baldwin forest, carrion, 30 May–8 Jun 1977, S. Peck; 2 females, Cerro Punta, 2 km E, 1760 m, Baldwin Forest, dung, 3 Jun 1977, S. Peck; 10 males, 8 females, Cerro Punta, 2 km E, 1760 m, Baldwin forest, dung traps, 30 May–8 Jun 1977, S. Peck; 2 females, Cerro Punta, 2 km E, 2200 m, forest, carrion traps, 1–4 Jun 1977, S. Peck; 1 male, 1 female, Cerro Punta, 2 km W, 1760 m, Baldwin Forest, carrion, 30 May–2 Jun 1977, S. Peck; 6 males, 24 females, Cerro Punta, 2 km W, 1760 m, Baldwin forest, dung, 5 Jun 1977, S. Peck; 25 males, 37 females, Cerro Punta, 2 km W, 1760 m, Baldwin Forest, dung trap, 30 May–2 Jun 1977,

S. Peck; 1 female, Cerro Punta, 4.5 km E, 2500 m, carrion, 23–28 May 1977, S. Peck; 13 males, 4 females, Hartmann's Finca, 1550 m, dung trap, 31 May 1977, S. Peck; 1 female, Hartmann's Finca,

15 km NW Hato de Volcán, 1200 m, carrion trap, 20–31 May 1977, S. Peck; 1 male, Hartmann's

Finca, 15 km NW Hato de Volcán, 1200 m, dung trap, 20–25 May 1977, S. Peck; 7 males, 6 females, Hartmann's Finca, 15 km NW Hato de Volcán, 1200 m, dung trap, 20–31 May 1977, S.

Peck; 2 females, Hartmann's Finca, 15 km NW Hato de Volcán, 1500m, carrion trap, 20–31 May

1977, S. Peck; 5 males, 11 females, Hartmann's Finca, 15 km NW Hato de Volcán, 1500m, dung,

20–25 May 1977, S. Peck; 2 males 3 females, Lagunas, 5km SW Hato del Volcan, 1360m, dung,

22–26 May 1977, S. Peck. PERU: Loreto: 1 male, Teniente López, 1.5 km N, 0°31'S, 76°10'W,

230–305 m, FIT, 22 Jul 1993, R. Leschen (MUSM). TRINIDAD & TOBAGO: Tobago: 1 female,

Charlotteville, Man-O-War Bay cottages, littoral rainforest, UV light, 26–30 Jun 1993, S. & J. Peck.

VENEZUELA: Aragua: 1 female, Henri Pittier National Park, Maracay-Choroni highway, km 19,

1330 m, creek, 15 Apr 1994, L. Masner; 11 males, 8 females, Henri Pittier National Park, Rancho

Grande Biological Station, 10°21'N, 67°41'W, 1250 m, May 1998, Ashe, Brooks & Hanley (DEBU,

MIZA); 2 females, Rancho Grande Biological Station, 10°21'N, 67°41'W, 1200 m, flight intercept trap, 14 May 1998, Ashe, Brooks & Hanley; Bolivar: 1 female, km40 Sta. Elena Icabaru Road,

100m, 4 Aug 1986, B.D. Gill (CNCI); 1 male, 3 females, Yacambu, 1200m, cloud forest, 7 May

1981, H.K. Townes; 1 female, Yacambu, 1200m, 10 May 1981, H.K. Townes; Mérida: 3 males, 4 females, Jaji, dung traps, 27 Apr–3 May 1988, S.A. Marshall; 1 male, Los Chorros, ~2300 m, decayed vegetation, 23 Apr 1989, S.A. Marshall (MIZA); 1 female, Los Chorros, ~2300 m, FIT,

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23–30 Apr 1988, S.A. Marshall (MIZA); 2 males, Los Chorros, dung traps, 1–5 May 1988, S.A.

Marshall.

Comments: The surstylus of A. megavilla is similar to both A. cobolorum, from which it can be easily separated by the mid tibial chaetotaxy, and A. braziliensis, which has a distinctly different sternite 5.

Etymology: The species epithet refers to the large finger-like projection on the male surstylus.

6.8 Discussion

Archiceroptera is almost entirely Neotropical, with a handful of collections from the

Nearctic. Most study material came from Central America, Ecuador, and Venezuela, where there have been numerous directed efforts to sample dung associated insects. The distribution of all study material of Archiceroptera (Fig. 6.64D) shows a tropical distribution, with northern Argentina the southern-most limit of the genus. The absence of Archiceroptera in material from numerous collecting efforts from Chile and Argentina in DEBU, including those collected with dung traps, supports this tropical distribution. Limited records from west of the Andes and Nearctic parts of

Mexico also suggests that mountains are significant barriers for this group, with only a few species occurring at high elevation sites. The morphologically unique clade formed by A.bisetosus, A. basilia and A. bilobata has a unique distribution within Archiceroptera as all three species are associated with high elevation sites in Ecuador and Peru. Although the major diversity of

Archiceroptera now appears to be documented, there may still be remaining species to be discovered, especially within the clades that occur in higher elevations (e.g., the A. basilia clade).

The most significant contribution that might be made to Archiceroptera is additional material from eastern South America. Based on the novelty rate of material collected from French Guiana for general unbaited pan traps, it is possible another 5-10 undescribed species remain from this area.

Contributions can also be made with adequately preserved material for molecular analysis.

Additional sequences of other Archiceroptera species, especially of the clades of the A. ternum group that were not represented here, may help ellucidate the relationships and determine if the differences from the morphological analysis are sampling artifacts. - 207 -

6.9 Chapter References

Buck, M. and S.A. Marshall. 2009. Revision of New World Leptocera Olivier (Diptera,

Sphaeroceridae). Zootaxa, 2039: 1–139.

Colwell, R.K. 2012. Biota 3: The biodiversity database manager. Software and 880 p. Manual.

Available online at: http://viceroy.eeb.uconn.edu/Biota.

Cumming, J.M., and D.M. Wood. 2010. Adult morphology and terminology. Pp.9–63 In B.V.

Brown (ed.) Manual of Central American Diptera Volume 1. NRC Research Press.

Goloboff, P.A., J. Farris and K. Nixon. 2008. TNT: a free program for phylogenetic analysis.

Cladistics, 24:774–786.

Guindon S., J.F. Dufayard, V. Lefort, M. Anisimova, W. Hordijk, and O. Gascuel. 2010. New

algorithms and methods to estimate Maximum-Likelihood phylogenies: Assessing the

performance of PhyML 3.0. Systematic Biology, 59(3): 307–321, 2010.

Maddison, W.P. and D.R. Maddison. 2017. Mesquite: a modular system for evolutionary analysis.

Version 3.2 http:mesquiteproject.org

Marshall, S.A. and M. Buck. 2010. Sphaeroceridae (Small dung flies). Pp1165–1187 in Manual of

Central American Diptera. Eds. B.V. Brown, A. Borkent, J.M. Cumming, D.M. Wood,

N.E. Woodley and M.A. Zumbado. NRC Research Press, Ottawa, Ontario. 1442 p.

Marshall, S.A. and Y. Cui. 2005. Systematics of Robustagramma, a new genus of New World

Sphaeroceridae (Diptera). Zootaxa, 1026: 1–122.

Nixon, K. C. 2002. WinClada ver. 1.00.08 Published by the author, Ithaca, NY, USA.

Papp, L. 1977. A contribution to the knowledge of species of the subfamily Ceropterinae (Diptera:

Sphaeroceridae). Acta Zoologica Academiae Scientiarum Hungaricae, 23(3–4): 371–385.

Richards, O.W. 1963. Sphaerocerid flies from South and Central America in the collection of the

California Academy of Sciences (Diptera). The Pan-Pacific Entomologist, 39: 231–246.

Richards, O.W. 1967. On a collection of Sphaeroceridae (Diptera) from the Galapagos Islands.

Annals and Magazine of Natural History Series 13, 9: 531–535.

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Shorthouse, D.P. 2010. SimpleMappr, an online tool to produce publication-quality point maps.

[Retrieved from http://www.simplemappr.net. Accessed January 29, 2015].

Smith, I.P., and S.A. Marshall. 2004. A review of the New World genus Pterogramma Spuler and a

revision of the Pterogramma sublugubrinum group (Diptera: Sphaeroceridae:

Limosininae). Contributions in Science, 499: 1–163.

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1 6.10 Archiceroptera Tables and Figures

2 Table 6.1. Morphological character states for phylogenetic study of Archiceroptera. Character 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 29 30 31 32 33 34 35 36 37 38 39 40 41 42 43 44 45 46 47 48 49 50 51 52 53 54 55 56 57 58 Thoracochaeta 0 0 1 0 0 0 0 01 01 0 0 2 1 0 0 1 0 ? 0 ? 0 0 ? 0 ? 0 0 0 1 0 0 0 0 0 0 0 0 0 0 0 0 ? 0 0 2 ? 0 0 0 0 0 0 0 0 0 0 0 0 Pectinosina 0 2 1 0 0 0 0 0 0 0 0 0 0 0 0 1 2 1 0 0 0 0 0 0 0 0 0 0 0 0 0 1 0 0 0 0 0 0 0 0 0 ? 0 0 2 ? 0 2 0 0 0 0 0 0 0 0 0 ? A. addenda 0 2 0 0 1 1 1 0 0 0 0 0 1 0 0 1 2 0 1 1 0 0 1 0 0 1 1 0 1 0 0 0 0 1 1 1 1 1 0 0 0 ? 0 0 1 0 1 2 0 0 1 1 1 1 0 0 0 0 A. crenulata 0 1 0 0 1 1 1 1 1 0 0 0 0 1 0 1 2 0 1 2 0 0 1 1 0 1 0 0 0 0 1 0 0 1 0 1 1 0 0 0 0 ? 0 0 2 - 0 2 0 0 1 0 1 1 0 0 2 0 A. triclavus 0 2 0 0 1 1 1 0 0 0 0 0 1 0 0 1 2 0 1 1 0 0 1 0 0 0 0 0 1 0 0 0 0 0 1 0 1 0 0 0 0 ? 0 0 2 - 0 2 0 0 1 1 1 1 0 0 0 0 A. browni 2 0 2 1 0 0 1 0 0 0 1 0 0 1 1 1 1 - 0 - 0 0 0 0 2 0 0 0 0 0 0 1 0 1 0 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? A. mahukani 0 0 2 1 0 0 1 0 0 0 1 1 ? 1 0 1 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? 0 0 0 1 0 0 0 0 0 1 2 0 0 1 0 1 0 1 0 0 0 A. venezolana 1 0 2 1 0 0 1 0 0 0 0 0 0 1 0 1 1 - 0 - 0 1 0 0 2 0 0 0 0 0 0 1 1 1 0 1 0 0 0 0 1 0 1 0 0 0 1 0 0 0 1 0 1 0 1 0 0 0 A. adamas 0 2 1 0 0 0 1 0 0 0 0 0 0 1 0 0 0 1 0 1 0 2 1 0 1 0 1 0 0 0 0 1 0 1 0 1 0 1 0 0 1 1 1 0 0 1 1 0 0 1 1 1 1 0 1 1 2 1 A. barberi 0 2 1 0 0 0 1 0 01 1 0 0 0 1 0 01 2 1 0 1 0 0 0 0 2 1 1 0 0 0 0 1 0 1 1 0 0 1 0 1 1 1 1 0 0 1 1 0 0 1 1 1 1 0 1 1 1 1 A. basilia 0 2 1 0 0 1 1 1 1 0 0 1 0 0 1 0 2 1 0 0 1 1 1 0 1 0 0 0 0 0 0 1 1 1 0 0 0 0 1 0 1 1 0 1 0 0 1 3 1 1 1 1 1 0 1 1 2 0 A. bilobata 0 2 1 1 0 1 1 1 1 0 0 1 0 0 0 1 2 0 0 2 0 1 1 0 1 0 1 1 0 0 0 1 0 1 0 0 0 0 1 0 1 1 0 1 0 0 1 3 1 1 1 1 1 0 1 1 0 0 A. bisetosus 0 2 2 0 0 0 0 01 01 1 1 1 1 0 0 0 2 0 0 0 0 1 1 0 1 0 0 1 0 1 0 1 1 1 0 1 0 1 1 0 1 1 0 1 0 1 1 3 1 1 1 1 1 0 1 1 0 0 A. caliga 0 2 1 0 0 0 1 0 0 0 1 0 0 1 0 01 2 0 0 2 0 2 1 0 1 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 1 1 1 0 0 0 1 0 0 1 1 1 1 0 1 1 3 0 A. calligrapha 0 2 1 0 0 0 1 0 0 0 0 0 0 1 0 01 2 0 0 1 0 1 1 0 2 0 0 1 0 0 0 1 0 0 1 0 0 1 0 0 1 1 1 0 0 1 1 1 0 1 1 1 1 0 1 1 0 1 A. cobolorum 0 2 2 0 0 1 1 0 0 0 0 0 0 1 0 1 1 0 0 - 0 0 0 0 2 0 2 1 1 0 0 1 0 1 1 1 0 0 0 1 1 1 0 0 0 0 1 2 0 1 1 1 1 0 1 1 0 A. dolabra 0 2 1 0 0 0 1 0 0 0 0 0 0 0 0 0 2 0 0 2 0 1 1 0 1 0 0 0 0 1 0 1 0 0 1 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? A. maniba 0 2 2 0 0 0 1 0 0 0 0 0 0 1 1 0 2 1 0 1 0 2 1 0 2 0 1 0 0 0 0 1 0 0 0 0 0 1 0 0 1 1 0 0 0 1 1 0 0 1 1 1 1 0 1 1 2 1 A. masoni 0 2 1 0 0 0 1 0 1 0 1 1 0 0 1 0 2 0 0 1 0 2 1 0 1 1 1 0 0 0 0 1 1 1 0 0 0 1 0 0 1 1 0 0 0 1 1 0 0 1 1 1 1 0 1 1 0 0 A. megacerca 0 2 1 0 0 1 1 01 01 0 0 0 0 1 01 1 2 1 0 1 0 0 0 0 2 0 0 0 0 2 0 0 0 0 0 1 0 1 0 0 1 1 0 0 0 0 1 3 0 1 1 1 1 0 1 1 2 1 A. mexicorona 0 2 1 0 0 0 1 01 01 0 1 1 0 1 0 0 2 1 0 1 1 0 1 0 2 0 0 0 0 0 1 1 0 1 0 0 0 1 0 1 1 1 1 0 0 0 1 0 0 1 1 1 1 0 1 1 2 1 A. mitarakai 0 2 1 0 0 0 1 0 0 0 1 0 0 1 0 0 2 0 0 1 0 2 1 0 1 1 0 0 0 0 0 1 0 0 0 0 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? A. paracerca 0 2 1 0 0 1 1 01 01 0 2 0 0 1 1 1 2 0 0 0 0 1 0 0 2 0 0 0 0 2 0 0 0 0 0 1 0 ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? ? A. pussula 0 2 1 0 0 0 1 01 01 0 1 0 0 1 0 0 2 0 0 2 0 1 1 0 1 0 0 0 0 0 0 1 1 0 0 0 0 1 0 0 1 1 1 0 0 1 1 0 0 1 1 1 1 0 1 1 2 0 A. ternum 0 2 1 0 0 0 1 0 0 0 1 1 0 1 0 0 2 1 0 1 0 1 1 0 1 0 1 0 0 0 0 0 1 0 0 0 0 1 0 0 1 1 0 0 0 0 1 0 0 1 1 1 1 0 1 1 0 0 A. uncinata 0 2 1 0 0 0 1 0 0 1 1 2 0 1 0 01 2 0 0 2 0 2 1 0 1 0 1 0 0 0 0 1 1 0 0 0 0 1 0 0 1 1 0 0 0 1 1 0 0 1 1 1 1 0 1 1 2 1 A. braziliensis 0 2 1 0 1 0 0 1 2 2 2 2 0 1 0 0 0 0 0 - 0 0 0 0 2 0 2 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 0 1 0 0 1 2 0 1 1 1 1 0 1 1 0 1 A. brevivilla 0 2 1 0 1 0 0 1 2 2 2 2 0 1 0 0 2 0 0 1 0 0 0 0 2 1 1 0 0 0 0 1 1 1 0 0 0 0 0 0 1 0 1 1 0 0 1 3 0 1 1 1 1 0 1 1 2 1 A. curvivilla 0 2 1 0 1 1 0 1 2 2 2 2 0 1 0 0 0 1 0 - 0 0 0 0 2 1 1 0 0 0 0 1 1 1 0 1 0 0 0 0 1 0 0 1 0 0 1 3 0 1 1 1 1 0 1 1 2 0 A. llama 0 2 1 0 1 1 0 1 1 2 2 2 0 1 0 0 2 0 0 1 0 0 0 0 1 1 1 0 0 0 0 1 0 0 1 0 0 0 0 1 1 1 1 1 0 0 1 0 0 1 1 1 1 0 1 1 3 1 A. megavilla 0 2 1 0 1 1 0 1 2 2 2 2 0 1 0 0 0 0 0 - 0 0 0 0 2 0 2 0 0 0 0 1 0 0 1 0 0 0 0 0 1 0 1 1 0 0 1 23 0 1 1 1 1 0 1 1 2 0

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Figure 6.1. Strict consensus tree for Archiceroptera of 70 retained trees from Traditional Search (TNT).

Bootstrap and Jackknife values > 50 are given above and below (respectively).

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Figure 6.2. Majority Rules tree for Archiceroptera from 70 trees from Traditional Search (TNT).

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Figure 6.3. Phylogeny of Archiceroptera. Tree selected from 70 equally parsimonious trees (Length = 225, Ci =

34, Ri = 62).

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Figure 6.4. Maximum likelihood analysis of cytochrome c oxidase subunit I (COI) 5P for Archiceroptera with

3rd codon included (A) and excluded (B). Numbers at nodes are aBayes values. The leading codes for each specimen are the unique identifiers within the BOLD database.

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Figure 6.5. Archiceroptera venezolana male terminalia: A) abdomen, ventral view; B) sternite 5, ventral view;

C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, close up lateral; F) postgonite; lateral view; G) phallus, dorsal view; H) same, dorsolateral view; I) same, lateral view. A-I) from debu00373840.

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Figure 6.6. Archiceroptera venezolana phallus. A) dorsal view; B) dorsolateral view; and C) lateral view. A-C) from debu00373840.

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Figure 6.7. Archiceroptera venezolana female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00373801.

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6.10.1 Archiceroptera addenda species group

Figure 6.8. Archiceroptera addenda male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior half of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) distiphallus, postgonite and phallapodeme, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-

H) from debu00260830.

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Figure 6.9. Archiceroptera addenda female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084477.

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Figure 6.10. Archiceroptera crenulata male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) left surstylus; lateral view; F) distiphallus and postgonite, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-E) from debu01084494; F-H) from debu01084488.

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Figure 6.11. Archiceroptera crenulata female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084493.

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Figure 6.12. Archiceroptera triclavus male terminalia: A) abdomen, ventral view; B) posterior margin of sternite 5, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus; lateral view; F) distiphallus and postgonite, dorsal view; G) distiphallus, phallapodeme (in part) and postgonite, dorsolateral view; H) same, lateral view. A-H) from debu01084483.

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Figure 6.13. Archiceroptera triclavus female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00378580.

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6.10.2 Archiceroptera mahukani species group

Figure 6.14. Archiceroptera browni (holotype; debu01077561). A) habitus, lateral view; B) habitus, dorsal view; C) head, dorsolateral view.

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Figure 6.15. Archiceroptera browni male terminalia: A) abdomen, ventral view; B) sternites 5–8, ventral view;

C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, close up lateral; F) postgonite; lateral view; G) phallus, postgonite and phallapodeme, dorsal view; H) phallus, dorsolateral view;

I) same, lateral view. A-I) from holotype (debu01077561).

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Figure 6.16. Archiceroptera mahukani (holotype). A) habitus, dorsal view; B) habitus, lateral view; C) head, dorsolateral view.

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Figure 6.17. Archiceroptera mahukani female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00295088.

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Figure 6.18. Archiceroptera venezolana (holotype). A) head, anterior view; B) habitus, lateral view; C) habitus, dorsal view.

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6.10.3 Archiceroptera ‘ternum species group’

Figure 6.19. Archiceroptera adamas male terminalia: A) abdomen, ventral view; B) sternite 5 (with fungal thalli) and synsternite 6+7 ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral

- 229 - view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view.

A-H) from debu00275292.

Figure 6.20. Archiceroptera adamas female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00379812.

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Figure 6.21. Archiceroptera barberi male terminalia: A) abdomen, ventral view; B) posterior part of sternite 4, sternite 5, and transverse portion of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view;

H) same, lateral view. A-H) from debu00107565.

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Figure 6.22. Archiceroptera barberi female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00107520.

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Figure 6.23. Archiceroptera basilia male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior parts of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) phallus, postgonite and phallapodeme, dorsal view; F) same, dorsolateral view; G) same, lateral view. A-G) from debu00140620.

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Figure 6.24. Archiceroptera basilia female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00140638.

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Figure 6.25. Archiceroptera bilobata male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior portion of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view.

A-H from debu01084831.

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Figure 6.26. Archiceroptera bilobata female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084833.

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Figure 6.27. Archiceroptera bisetosus male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view;

E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-D) from debu00189533; E-H) from debu00189536.

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Figure 6.28. Archiceroptera bisetosus female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00189555.

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Figure 6.29. Archiceroptera caliga male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite

6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01085358;

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Figure 6.30. Archiceroptera caliga female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085364.

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Figure 6.31. Archiceroptera calligraphia male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01085530.

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Figure 6.32. Archiceroptera calligraphia female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085517.

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Figure 6.33. Archiceroptera cobolorum male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view;

E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01086488.

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Figure 6.34. Archiceroptera cobolorum female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01086522.

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Figure 6.35. Archiceroptera dolabra male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite

6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00385443.

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Figure 6.36. Archiceroptera maniba male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view;

E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00378585.

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Figure 6.37. Archiceroptera maniba female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00378572.

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Figure 6.38. Archiceroptera masoni male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view;

E) postgonite; lateral view; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01085698.

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Figure 6.39. Archiceroptera masoni female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085701.

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Figure 6.40. Archiceroptera megacercus male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view;

E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-

H) from debu01084910.

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Figure 6.41. Archiceroptera megacercus female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084919.

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Figure 6.42. Archiceroptera mexicorona: A) epandrium, surstylus and cercus, caudal view; B) same, lateral view; C) sternite 5 and anterior part of synsternite 6+7, ventral view; D) left surstylus, lateral view; E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01085710.

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Figure 6.43. Archiceroptera mexicorona female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085713.

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Figure 6.44. Archiceroptera mitarakai male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view;

E) distiphallus and postgonite, dorsal view; F) same, dorsolateral view; G) same, lateral view. A-G) from debu00394580.

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Figure 6.45. Archiceroptera paracercus, male terminalia male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) surstylus, posterolateral view; F) postgonite; lateral view; G) distiphallus, dorsal view;

H) same, dorsolateral view; I) same, lateral view. A-I) from debu00260795.

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Figure 6.46. Archiceroptera pussula male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite

6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A) from debu00196122, B-H) from debu01085674.

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Figure 6.47. Archiceroptera pussula female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01085659.

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Figure 6.48. Archiceroptera ternum male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite

6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu00132380.

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Figure 6.49. Archiceroptera ternum female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-C) from debu00260557; D) from debu00132367.

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Figure 6.50. Archiceroptera uncinata male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior part of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view;

E) postgonite; lateral view; F) distiphallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-

H) from debu00382406.

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Figure 6.51. Archiceroptera uncinata female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01084712.

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6.10.4 Archiceroptera brevivilla species subgroup

Figure 6.52. Archiceroptera braziliensis male terminalia: A) abdomen, ventral view; B) sternite 5 and synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite, close up lateral; F) phallus,dorsal view; G) phallus, dorsolateral view; H) phallus, lateral view. A-

H) from debu01088400.

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Figure 6.53. Archiceroptera braziliensis female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01088403.

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Figure 6.54. Archiceroptera brevivilla male terminalia: A) abdomen, ventral view; B) sternite 5 and anterior portion of synsternite 6+7, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite, close up lateral; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01088231.

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Figure 6.55. Archiceroptera brevivilla female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01086593.

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Figure 6.56. Archiceroptera curvivilla male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite, close up lateral; F) phallus,dorsal view; G) same, dorsolateral view; H) same, lateral view. A-H) from debu01088418.

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Figure 6.57. Archiceroptera curvivilla female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu01088446.

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Figure 6.58. Archiceroptera llama male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) phallus and postgonite, dorsal view; G) same, dorsolateral view; H) same, lateral view. A-G) from debu00386692.

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Figure 6.59. Archiceroptera llama female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. A-D) from debu00386665.

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Figure 6.60. Archiceroptera megavilla male terminalia: A) abdomen, ventral view; B) sternite 5 and transverse part of sternite 6, ventral view; C) epandrium, surstylus and cercus, caudal view; D) same, lateral view; E) postgonite, close up lateral; F) phallus, dorsal view; G) same, dorsolateral view; H) same, lateral view. (A and

C-H from debu01088807, B from debu01086594).

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Figure 6.61. Archiceroptera megavilla female terminalia and spermathecae: A) terminal abdominal segments, dorsal view; B) terminal abdominal segments, lateral view; C) terminal abdominal segments, ventral view; D) spermathecae. (A and C from debu01086596; B and D from debu01086595).

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6.10.5 Distribution maps

Figure 6.62. Distribution of Archiceroptera species: A) the addenda species group (A. crenulata, A. addenda, and A. triclavus); B) Archiceroptera s. str. (A. browni, A. mahukani, and A. venezolana); C) A. maniba, A. adamas, and A. calligraphia; and D) A. masoni, A. mexicorona, A. uncinata, and A. cobolorum.

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Figure 6.63. Distribution of Archiceroptera species: A) A. megacercus and A. paracercus; B) A. caliga, A. pussula, A. mitarakai, and A. dolabra; C) A. basilia, A. bisetosus, and A. bilobata; and D) A. ternum.

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Figure 6.64. Distribution of Archiceroptera species: A) A. barberi; B) A. brevivilla, A. curvivilla, and A. llama;

C) A. braziliensis and A. megavilla; and D) all Archiceroptera specimens.

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CHAPTER 7 – SUMMARY

A preliminary estimate of the relationships within the EPG is provided on the basis of the morphological phylogeny. The Archiceroptera genus complex was not recovered as monophyletic in the morphological analysis, instead resolving into four distinct clades within the EPG. The EPG is maintained here as a monophyletic group, based on the synapomorphic epandrial process and mid tibial chaetotaxy, but further morphological and molecular studies are needed to examine the relationships with other Limosininae genera. The molecular analysis of the

Limosininae did not provide any useful resolution to potential outgroups for the EPG or strong support for any relationships between EPG genera. A narrower analysis of only Archiceroptera sequences supports the morphological treatment of brevivilla and ternum groups within Archiceroptera, along with the treatment of the extension (addenda) group as a basal clade to the rest of Archiceroptera.

Archiceroptera and Rudolfina are redefined, Pectinosina is described, and 39 new species are described in these three genera. The revisions of Archiceroptera, Rudolfina and Pectinosina provide keys, descriptions and illustrations of all the described species. Archiceroptera is primarily Neotropical, now contains 29 described species and is treated in a clade that also includes Pectinosina and Robustagramma. Pectinosina is also largely Neotropical, and includes only two described species. Rudolfina is largely Nearctic, now contains 13 described species, and is treated in a clade that also includes Pterogramma, Aptilotella, Bromeloecia, and Bitheca, along with the sororcula and enigmata groups. The sororcula/enigmata clade remains undescribed, but preliminary work on the group suggests there are ~10 species.

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CHAPTER 8 – REFERENCES

The following references are for the thesis as a whole. Several chapters provide an annotated list of references for

that chapter.

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(southeastern Mexico). Tectonophysics, 287, 77–96

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with a description of 8 new species. Zootaxa, 3953: 1–157.

Buck, M. and S.A. Marshall. 2009. Revision of New World Leptocera Olivier (Diptera, Sphaeroceridae). Zootaxa,

2039: 1–139.

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Systematics, 27: 282–293

Kits, J.H. and S.A. Marshall. 2011. A revision of Frutillaria Richards and Penola Richards (Diptera:

Sphaeroceridae: Archiborborinae). Zootaxa 2863: 1–34.

Kits, J.H., S.A. Marshall and J.H. Skevington. 2012. Phylogeny of the Archiborborinae (Diptera: Sphaeroceridae)

based on combined morphological and molecular analysis. PLoS ONE 8(1): e51190.

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Maddison, W.P. and D.R. Maddison. 2017. Mesquite: a modular system for evolutionary analysis. Version 3.2

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(PhD Thesis). University of Guelph, Guelph, Ontario. 320p.

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Marshall, S.A. 1983b. The genus Bromeloecia Spuler in North America (Diptera: Sphaeroceridae: Limosininae).

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Marshall, S.A. 1983c. Ceroptera sivinskii, a new species of Sphaeroceridae (Diptera) in a genus new to North

America, associated with scarab beetles in the southwestern United States. Proceedings of the

Entomological Society of Washington, 85: 139–143.

Marshall, S.A. 1985. The genera Xenolimosina and Terrilimosina (Diptera: Sphaeroceridae: Limosininae) in North

America. Proceedings of the Entomological Society of Washington, 87: 759–769.

Marshall, S.A. 1991. Rudolfina digitata sp. nov., a new Nearctic sphaerocerid with a disjunct alpine-arctic

distribution. Canadian Entomologist, 123: 621–626.

Marshall, S.A. 1995. Sclerocoelus and Druciatus, new genera of New World Sphaeroceridae (Diptera;

Sphaeroceridae; Limosininae). Insecta Mundi, 9:283–289. - 277 -

Marshall, S.A. 1996. Tucma fritzi, a new species in the enigmatic genus Tucma Mourgues-Schurter (Diptera;

Sphaeroceridae; Tucminae, new subfamily). Studia Dipterlogica 3: 283–288.

Marshall, S.A. 1997. Limomyza, a new genus of primitive Limosininae (Diptera: Sphaeroceridae), with five new

species from United States, Mexico, and Central America. Proceedings of the Entomological Society of

Washington, 99: 279–280.

Marshall, S.A. 1998. A revision of the Archileptocera group, including Anomioptera Schiner, Palaeocoprina Duda

and Archileptocera Duda, with a key to sphaerocerid genera with similar wing venation and a description

of a new species of Palaeoceroptera Duda (Diptera Sphaeroceridae). Journal of Natural History, 32: 173–

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Dipterlogica, 7: 109–144.

Marshall, S.A. 2000b. Chespiritos, a new genus of Limosininae (Diptera: Sphaeroceridae) from Costa Rica.

Proceeding of the Entomological Society of Washington, 102: 609–612.

Marshall, S.A. 2001. A review of the southern South American genus Gyretria Enderlein (Diptera: Sphaeroceridae:

Limosininae). Proceedings of the Entomological Society of Washington, 103: 282–290.

Marshall, S.A. 2013. Bregmosina, a new neotropical genus of Limosininae (Diptera: Sphaeroceridae). Zootaxa,

3641(3): 260–270.

Marshall, S.A. 2014. Albostyla, a new genus of neotropical Limosininae (Diptera: Sphaeroceridae). Zootaxa, 3793:

257–264.

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Diptera. Eds. B.V. Brown, A. Borkent, J.M. Cumming, D.M. Wood, N.E. Woodley and M.A. Zumbado.

NRC Research Press, Ottawa, Ontario. 1442 p.

Marshall, S.A. and Y. Cui. 2005. Systematics of Robustagramma, a new genus of New World Sphaeroceridae

(Diptera). Zootaxa, 1026: 1–122.

Marshall, S.A. and H. Dong. 2008. Parasclerocoelus, a new south temperate genus of Limosininae (Diptera:

Sphaeroceridae). Studia Dipterologica, 14:223–230.

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(Diptera) from Colorado and Arizona. Proceedings of the Entomological Society of Washington, 99: 641–

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Sphaeroceridae, Limosininae). Contributions in Science, 474: 1–27.

Marshall, S.A., R. Langstaff and D. Grimaldi. 1999. Taxonomic names, in new species of Sphaeroceridae (Insecta:

Diptera) in Dominican amber. Studia Dipterologica, 6:295–304.

Marshall, S.A. and O.W. Richards. 1987. Sphaeroceridae. Pp 601 – 685 In J.F. McAlpine, B.V. Peterson, G.E.

Shewell, H.J. Teskey, J.R. Vockeroth, D.M. Wood (eds), Manual of the Nearctic Diptera, Part 2.

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world catalog update covering the years 2000–2010, with new generic synonymy, new combinations, and

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Nixon, K. C. 2002. WinClada ver. 1.00.08 Published by the author, Ithaca, NY, USA.

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APPENDIX 1 – SYNOPSIS OF NEWLY DESCRIBED SPHAEROCERDIAE SPECIES SINCE THE LAST CATALOG UPDATE.

The following is a list of newly described Sphaeroceridae species since the last catalog update (Marshall et al.

2010). In total 115 Limosininae, 1 Sphaerocerinae, 10 Copromyzinae and 83 Archiborborinae (previously treated as part of the Copromyzinae) species have been newly described since 2010; 12 new Limosininae and 5 new

Archiborborinae genera have also been described since. Style follows Marshall et al. (2010). New distributional data for previously described species is not incorporated here. A preliminary list of manuscript names is also included, representing the work of this thesis and that of Yau (M.Sc.); a total of 68new species are represented.

The follow are the acronyms for the depositories of holotypes:

BMSA Entomology Department, National Mseum, Bloemfontein, South Africa

CBFC Museo Nacional de Historia Natural, La Paz, Bolivia

CNCI Canadian National Collection of Insects, Agriculture and Agri-Food Canada, Ottawa, Ontario, Canada.

DEBU Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada.

DZUP Universidade Federal do Paraná, Curitibia, Brazil

FMNH Field Museum of Natural History, Chicago, USA

HMNH Hugarian Natural History Museum, Budapest

HNHM Hungarian Natural History Museum, Budapest, Hungary.

IAVH Institutio Alexander von Humboldt, Villa de Leyva, Colombia

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

IRSN Institute Royal des Sciences Naturelles de Belgique, Brussels

SACS Liaoning Key Laboratory of Urban Integrated Pest Management and Ecological Security, Shenyang

University, China

MNHN Entomologie, Museum National d'Histoire Naturelle, Paris, France.

MNNC Museo Nacional de Historia Natural, Santiago, Chile

MUSM Museo de Historia Natural, Universidad Nacional Mayor de San Marcos, Lima, Peru

MZLU Lund University, Lund, Sweden

MZSP Museu de Zoologia, Universidade de São Paulo, São Paulo, São Paulo, Brazil

NMSA Departement of Natural Sciences, KwaZulu-Natal Museum, Pietermaritzburg, South Africa - 283 -

NZAC New Zealand Arthropod Collection, Auckland

NMNZ Entomological Collection, Museum of New Zealand, Wellington.

MIZA Museo del Instituto de Zoología Agrícola, Maracay, Venezuela

QCAZ Departamento de Biología, Pontífica Universidad Católica del Ecuador, Quito, Ecuador

ROME Royal Ontario Museum, Toronto, Ontario, Canada

SMOC Slezské zemské muzeum, Opava, Czech Republic

UASC Museo de Historia Natural Noel Kempff Mercado, Santa Cruz de la Sierra, Bolivia

UNAM Universidad Nacional Autónoma de México, Mexico City, Mexico

USNM United States National Museum of Natural History, Smithsonian Institution, Washington, District of

Columbia, USA

UTPL Universidad Técnica Particular de Loja, Loja, Ecuador (held at Institut Royal des Sciences Naturelles de

Belgique, Brussels, Belgium)

A1.1 DESCRIBED LIMOSININAE SINCE LAST CATALOG UPDATE

Genus Afrolimosina Papp 2014b

Afrolimosina, 2014b: 102 (feminine). Type species: Afrolimosina albitarsis, original designation. – Papp (2014b): 102–104 [diagnosis, description, illustr.].

Afrolimosina albitarsis Papp, 2014b. Distr.: Afrotropical: Burundi.

Afrolimosina albitarsis Papp 2014b: 103 [both sexes, illustr.]. Type locality: Burundi, Bururi Province, Reserve Naturelle de Rumonge. HT male

(BMSA)

Genus Albistyla Marshall, 2014

Albistyla Marshall, 2014: 257 (feminine). Type species: Albistyla spatulisterna, original designation. – Marshall (2014): 257 [diagnosis, description, key, illustr.]. - 284 -

Albistyla fimbriata Marshall 2014. Distr.: Neotropical: Costa Rica, Ecuador, Venezuela.

Albistyla fimbriata Marshall 2014: 259 [males, illustr.]. Type locality: Venezuela, Maracay. Rancho Grande Biological Station. HT male (MIZA).

Albistyla occulta Marshall 2014. Distr.: Neotropical: Ecuador.

Albistyla occulta Marshall 2014: 261 [both sexes, illustr.]. Type locality: Ecuador, Napo Prov., Baeza. HT male (ROME).

Albistyla spatulisterna Marshall 2014. Distr.: Neotropical: Costa Rica.

Albistyla spatulisterna Marshall 2014: 261 [both sexes, illustr.]. Type locality: Costa Rica, Cartago, Tapantí National Park. HT male (INBC).

Genus Aptilotella Duda, 1924

Aptilotella Duda, 1924c: 74 (feminine). Type species: Aptilotella borgmeieri Duda, 1924, monotypy. - Richards, 1951a: 847, 849 [taxonomic notes, key]; Richards, 1965a: 459

[diagnosis in key]; Richards, 1967b: 7 [Neotropical catalog]; Hackman, 1969a: 207 [list, biogeography]; Luk & Marshall, 2014 [diagnosis, rediscription, revision, key, phylogeny, illustr.]

Aptilotella caerulea Luk & Marshall 2014. Distr.: Neotropical: Dominican Republic.

Aptilotella caerulea Luk & Marshall 2014: 18 [both sexes, illustr.]. Type locality: Dominican Republic, Independencia, 32 km NW La

Descrubierta Sabana Real. HT male (DEBU).

Aptilotella germana Luk & Marshall 2014. Distr.: Neotropical: Mexico.

Aptilotella germana Luk & Marshall 2014: 19 [both sexes, illustr.]. Type locality: Mexico, Chiapas, Cerro El Calvario, near Tapalapa. HT male

(UNAM).

Aptilotella pyropanda Luk & Marshall 2014. Distr.: Neotropical: Mexico.

Aptilotella pyropanda Luk & Marshall 2014: 20 [both sexes, illustr.]. Type locality: Mexico, Huixtán, Bazóm. HT male (UNAM).

- 285 -

Aptilotella gracilis Luk & Marshall 2014. Distr.: Nearctic: Mexico.

Aptilotella gracilis Luk & Marshall 2014: 21 [both sexes, illustr.]. Type locality: Mexico, Tamaulipas, Joya de Manantiales. HT male (UNAM).

Aptilotella gladia Luk & Marshall 2014. Distr.: Nearctic: Mexico.

Aptilotella gladia Luk & Marshall 2014: 23 [both sexes, illustr.]. Type locality: Mexico, Oaxaca, 5.1 km S Suchixtepec. HT male (UNAM).

Aptilotella hamata Luk & Marshall 2014. Distr.: Neotropical: Guatemala.

Aptilotella hamata Luk & Marshall 2014: 24 [both sexes, illustr.]. Type locality: Guatemala, Izabal, Firmeza. HT male (DEBU).

Aptilotella erinacea Luk & Marshall 2014. Distr.: Neotropical: Honduras.

Aptilotella erinacea Luk & Marshall 2014: 24 [both sexes, illustr.]. Type locality: Honduras, Cortés, Parque Nacional Cusuco, 18.7 km N

Cofradía, 5.4 km W Buenos Aires, Cerro Jilinco. HT male (DEBU).

Aptilotella diffisa Luk & Marshall 2014. Distr.: Neotropical: Costa Rica.

Aptilotella diffisa Luk & Marshall 2014: 25 [both sexes, illustr.]. Type locality: Costa Rica, Cartago, Llano Bonito, trail to Cerro Chirripó. HT male (INBC).

Aptilotella involucris Luk & Marshall 2014. Distr.: Neotropical: Costa Rica.

Aptilotella involucris Luk & Marshall 2014: 27 [both sexes, illustr.]. Type locality: Costa Rica, Cartago, Tapanti—Macizo de la Muerte National

Park, N of La Esperanza del Guarco. HT male (INBC).

Aptilotella sphyra Luk & Marshall 2014. Distr.: Neotropical: El Salvador, Honduras.

Aptilotella sphyra Luk & Marshall 2014: 28 [both sexes, illustr.]. Type locality: Honduras, Guisayote, 20.5 km E Ocotepeque. HT male (DEBU).

Aptilotella pinnifera Luk & Marshall 2014. Distr.: Neotropical: Guatemala.

Aptilotella pinnifera Luk & Marshall 2014: 29 [both sexes, illustr.]. Type locality: Guatemala, El Progreso, Cerro Pinalón. HT male (DEBU).

Aptilotella corona Luk & Marshall 2014. Distr.: Neotropical: Guatemala.

Aptilotella corona Luk & Marshall 2014: pp [both sexes, illustr.]. Type locality: Guatemala, 7 km N San Lorenzo. HT male (DEBU).

- 286 -

Aptilotella andersoni Luk & Marshall 2014. Distr.: Neotropical: Mexico.

Aptilotella andersoni Luk & Marshall 2014: 32 [both sexes, illustr.]. Type locality: Mexico, Chiapas, El Porvenir. HT male (UNAM).

Aptilotella quatuorchela Luk & Marshall 2014. Distr.: Neotropical: Mexico.

Aptilotella quatuorchela Luk & Marshall 2014: 33 [both sexes, illustr.]. Type locality: Mexico, Chiapas, El Triunfo Reserve, Pico El Triunfo. HT male (UNAM).

Aptilotella gloriosa Luk & Marshall 2014. Distr.: Neotropical: Mexico.

Aptilotella gloriosa Luk & Marshall 2014: 34 [both sexes, illustr.]. Type locality: Mexico, Chiapas, El Triunfo Reserve, Pico El Triunfo. HT male

(UNAM).

Aptilotella simplex Luk & Marshall 2014. Distr.: Neotropical: Costa Rica.

Aptilotella simplex Luk & Marshall 2014: 35 [both sexes, illustr.]. Type locality: Costa Rica, Guanacaste, Santa Elena Cloud Forest Reserve. HT male (INBC).

Aptilotella solaria Luk & Marshall 2014. Distr.: Neotropical: Guatemala.

Aptilotella solaria Luk & Marshall 2014: 36 [both sexes, illustr.]. Type locality: Guatemala, El Progreso, Cerro Pinalón, Peak. HT male (DEBU).

Aptilotella radians Luk & Marshall 2014. Distr.: Nearctic: Mexico.

Aptilotella radians Luk & Marshall 2014: 37 [both sexes, illustr.]. Type locality: Mexico, Oaxaca, Valle Nacional, 47.5 km SW, km 100.5. HT male (QCAZ).

Aptilotella ebenea Luk & Marshall 2014. Distr.: Neotropical: Ecuador.

Aptilotella ebenea Luk & Marshall 2014: 38 [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Bellavista Cloud Forest Reserve, 12 km S

Nanegalito. HT male (QCAZ).

Aptilotella gemmula Luk & Marshall 2014. Distr.: Neotropical: Ecuador.

Aptilotella gemmula Luk & Marshall 2014: 39 [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Bellavista Cloud Forest Reserve, 12 km S

Nanegalito. HT male (QCAZ).

Aptilotella quadrata Luk & Marshall 2014. Distr.: Neotropical: Costa Rica. - 287 -

Aptilotella quadrata Luk & Marshall 2014: 40 [both sexes, illustr.]. Type locality: Costa Rica, Highway 2, km 96. HT male (INBC).

Aptilotella umbracatus Luk & Marshall 2014. Distr.: Neotropical: Costa Rica, Panama.

Aptilotella umbracatus Luk & Marshall 2014: 42 [both sexes, illustr.]. Type locality: Costa Rica, Chiriquí, 4.5 km E Cerro Punta. HT male

(DEBU).

Aptilotella angela Luk & Marshall 2014. Distr.: Neotropical: Ecuador.

Aptilotella angela Luk & Marshall 2014: 43 [both sexes, illustr.]. Type locality:Ecuador, Carchi, Páramo El Ángel, 14.1 km NW El Ángel. HT male (QCAZ).

Aptilotella pichinchensis Luk & Marshall 2014. Distr.: Neotropical: Ecuador.

Aptilotella pichinchensis Luk & Marshall 2014: 45 [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Campamento Pichan, ~27.5 km NW

Quito. HT male (QCAZ).

Aptilotella viva Luk & Marshall 2014. Distr.: Neotropical: Venezuela.

Aptilotella viva Luk & Marshall 2014: 45 [both sexes, illustr.]. Type locality: Venezuela, Mérida, Sierra Nevada National Park, La Mucuy, 7 km

E Tabay. HT male (MIZA).

Aptilotella macula Luk & Marshall 2014. Distr.: Neotropical: Bolivia.

Aptilotella macula Luk & Marshall 2014: 46 [both sexes, illustr.]. Type locality: Bolivia, La Paz, Coroico, Cerro Uchumachi. HT male (CBFC).

Aptilotella macta Luk & Marshall 2014. Distr.: Neotropical: Bolivia.

Aptilotella macta Luk & Marshall 2014: 48 [males, illustr.]. Type locality: Bolivia, La Paz, Caranavi, ca. 10 km NW, road to ENTEL tower. HT male (CBFC).

Genus Biphallapodema Papp 2014b

Biphallapodema, Papp 2014b: 102 (feminine). Type species: Biphallapodema polydentata, original designation. – Papp (2014b): 105–107

[diagnosis, description, illustr.].

- 288 -

Biphallapodema oligodentata Papp, 2014b. Distr.: Afrotropical: Congo.

Biphallapodema oligodentata Papp 2014b: 107 [both sexes, illustr.]. Type locality: Congo, Brazzaville, ORSTOM park. HT male (HNHM).

Biphallapodema polydentata Papp, 2014b. Distr.: Afrotropical: Congo.

Biphallapodema oligodentata Papp 2014b: 109 [both sexes, illustr.]. Type locality: Congo, Brazzaville, ORSTOM park. HT male (HNHM).

Genus Chelilimosina Papp 2014b

Chelilimosina, Papp 2014b: 102 (feminine). Type species: Chelilimosina baloghi, original designation. – Papp (2014b): 111–113 [diagnosis, description, illustr.].

Chelilimosina baloghi Papp, 2014b. Distr.: Afrotropical: Congo.

Chelilimosina baloghi Papp 2014b: 113 [both sexes, illustr.]. Type locality: Congo, Brazzaville, ORSTOM park. HT male (HNHM).

Genus Bregmosina Marshall, 2013

Bregmosina Marshall, 2013: 261 (??gender). Type species: Bregmosina bucki, original designation. – Marshall (2013): 261 [diagnosis, description, key, illustr.].

Bregmosina bucki Marshall 2013. Distr.: Neotropical: Costa Rica.

Bregmosina bucki Marshall 2013: 262 [both sexes, illustr.]. Type locality: Costa Rica, Volcan Tenorio, N Slope nr. Biajagua Biological Station.

HT male (INBC).

Bregmosina ephydriformia Marshall 2013. Distr.: Neotropical: Costa Rica, Venezuela.

Bregmosina ephydriformia Marshall 2013: 263 [both sexes, illustr.]. Type locality: Venezuela, Merida. HT male (MIZA).

Bregmosina obunca Marshall 2013. Distr.: Neotropical: Costa Rica, Guyana.

- 289 -

Bregmosina obunca Marshall 2013: 266 [both sexes, illustr.]. Type locality: Costa Rica, Alajuela. Volcan Tenorio, N slope nr. Bijagua Biological

Station. HT male (INBC).

Bregmosina howdeni Marshall 2013. Distr.: Neotropical: Ecuador.

Bregmosina howdeni Marshall 2013: 268 [both sexes, illustr.]. Type locality: Ecuador, Pichincha Province, 47 km S Sto Domingo, Rio Palenque

Biological Station. HT male (DEBU).

Bregmosina schizosterna Marshall 2013. Distr.: Neotropical: Ecuador.

Bregmosina schizosterna Marshall 2013: 268 [unique male, illustr.]. Type locality: Ecuador, Maquipucuna Biological Reserve.. HT male

(QCAZ).

Genus Ceroptera Macquart, 1835

Note: Papp 2014a gives new distributional records for several described Old World species and places several other into synonomy.

Ceroptera armata Papp, 2014a. Distr.: Afrotropical

Ceroptera armata Papp, 2014a: xx [both sexes, illustr.]. Type locality: ####. HT ### (###).

Ceroptera globosa Papp, 2014a. Distr.: Afrotropical

Ceroptera globosa Papp, 2014a: xx [both sexes, illustr.]. Type locality: ####. HT ### (###).

Ceroptera inermis Papp, 2014a. Distr.: Afrotropical

Ceroptera inermis Papp, 2014a: xx [both sexes, illustr.]. Type locality: ####. HT ### (###).

Ceroptera miniscula Papp, 2014a. Distr.: Afrotropical

Ceroptera miniscula Papp, 2014a: xx [both sexes, illustr.]. Type locality: ####. HT ### (###).

Ceroptera moroccana Papp, 2014a. Distr.: Palaearctic

Ceroptera moroccana Papp, 2014a: xx [both sexes, illustr.]. Type locality: ####. HT ### (###).

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Ceroptera nigra Papp, 2014a. Distr.: Afrotropical

Ceroptera nigra Papp, 2014a: xx [both sexes, illustr.]. Type locality: ####. HT ### (###).

Ceroptera setiscutellata Papp, 2014a. Distr.: Afrotropical

Ceroptera setiscutellata Papp, 2014a: xx [both sexes, illustr.]. Type locality: ####. HT ### (###).

Genus Coproica Rodani, 1861

Note: Bergeron et al. 2015 give new distributional records for several described Nearctic species in both main part and in appendix.

Coproica bifuracata Bergeron, Marshall & Swann, 2015. Distr.: Neotropical: Argentina, Bolivia, Brazil.

Coproica bifurcata Bergeron, Marshall & Swann, 2015: 19 [both sexes, illustr.]. Type locality: Argentina, Salta, Rosario de Lerma. HT male

(DEBU).

Coproica bispatha Bergeron, Marshall & Swann, 2015. Distr.: Nearctic: USA (AZ, CO, TX). Neotropical:

Barbados, Brazil, Costga Rica, Dominican Republic, Ecuador, Honduras, Jamaica, Mexico (CHI, ROO, TAB), St.

Kitts & Nevis. Venezuela.

Coproica bispatha Bergeron, Marshall & Swann, 2015: 20 [both sexes, illustr.]. Type locality: Ecuador, Galapagos, Floreana, Agriculture zone.

HT male (CNCI).

Coproica brachystyla Bergeron, Marshall & Swann, 2015. Distr.: Nearctic: USA (FL, SC, TX, UT [data suspect]); Neotropical: Argentina, Barbados, Belize, Bolivia, Brazil, Costa Rica, Dominican Republic,

Ecuador, Guatemala, Honduras, Mexico (CAM, CHI, COA, ROO, TAB), Panama, Paraguay, Puerto Rico, St.

Kitts, St. Martin, Venezuela.

Coproica brachystyla Bergeron, Marshall & Swann, 2015: 22 [both sexes, illustr.]. Type locality: Costa Rica, San Jose, San Carlos, Riosparaiso

Reserve, Pecari Stn., 16 km NNE Quepos. HT male (INBC).

- 291 -

Coproica diabolica Bergeron, Marshall & Swann, 2015. Distr.: Nearctic: Canada (ON), USA (AR, FL, GA, IL,

MS, NC, OH, OK, TN, TX); Neotropical: Brazil, Costa Rica, Dominica, Dominican Republic, Ecuador,

Guyana, Grenada, Jamaica, Mexico (CHI), St. Kitts, St. Lucia, St. Vincent, Trinidad, Venezuela..

Coproica diabolica Bergeron, Marshall & Swann, 2015: 24 [both sexes, illustr.]. Type locality: U.S.A, North Carolina, Bladen Co., Singletary

Lk. St. Pk. HT male (DEBU? Not noted).

Coproica emarginata Bergeron, Marshall & Swann, 2015. Distr.: Nearctic: USA (FL, MI [data suspect], MS,

TX); Neotropical: Belize, Brazil, Costa Rica, Dominican Republic, Grenada, Guatemala, Mexico (ROO),

Puerto Rico, St. Kitts, St. Vincent, Trinidad, Venzuela.

Coproica emarginata Bergeron, Marshall & Swann, 2015: 27 [both sexes, illustr.]. Type locality: Costa Rica, Alajuela, Volcan Tenorio, Bijagua

Biol. Stn.,. HT male (INBC).

Coproica galapagosensisBergeron, Marshall & Swann, 2015. Distr.: Neotropical: Ecuador (Galapogos Is.).

Coproica galapogosensis Bergeron, Marshall & Swann, 2015: 31 [both sexes, illustr.]. Type locality: Ecuador, Galapagos, Espanola Bahia

Manzanillo. HT male (QCAZ).

Coproica novacula Bergeron, Marshall & Swann, 2015. Distr.: Nearctic: USA (AZ, TX); Neotropical:

Argentina, Bolivia, Brazil, Costa Rica, Ecuador, Guatemala, Honduras, Mexico (JAL, MEX, MOR, OAX,

PUE).

Coproica novacula Bergeron, Marshall & Swann, 2015: 35 [both sexes, illustr.]. Type locality: Costa Rica, Cartago, Rio Grande de Orosi, near

Tapanti Nat. Pk.,. HT male (INBC).

Coproica testudinea Bergeron, Marshall & Swann, 2015. Distr.: Nearctic: USA (FL).

Coproica testudinea Bergeron, Marshall & Swann, 2015: 39 [both sexes, illustr.]. Type locality: U.S.A., Florida, Putnam Co., Hollister. HT male

(DEBU).

Genus Eulimosina Roháček, 1983

- 292 -

Eulimosina prominulata Su, 2013. Distr.: Oriental: China.

Eulimosina prominulata Su et al. 2013b: 199 [males, illustr.]. Type locality: China, Guangzi, Shangsi, Mt. Shiwanda. HT male (SACS).

Genus Herniosina Roháček, 1983

Herniosina erymantha Roháček, 2016. Distr.: Palaearctic: Greece.

Herniosina erymantha Roháček, 2016: 80 [unique male, illustr.]. Type locality: Greece, NW Peloponnese, Alepochori 0.5 km SE. HT male

(SMOC).

Herniosina hamata Roháček, 2016. Distr.: Palaearctic: Cyprus.

Herniosina hamata Roháček, 2016: 91 [both sexes, illustr.]. Type locality: Cyprus, Troodos Mts., Pedoulas env.. HT male (SMOC).

Genus Howickia Richards, 1951.

Howickia Richards, 1951a: 844 (feminine). Type species: Apterina trilineata Hutton, 1901, original designation. - Richards, 1965a: 457 [diagnosis in key]; Hackman, 1969a: 207 [list];

Richards, 1973: 389 [diagnosis]; Marshall, 1989b: 603 [Australasian/Oceanian catalog]; Marshall et al. 2014 [revision of New Zealand species, key to New Zealand species, synonymized Biroina and Apterobiroina].

Biroina Richards, 1973: 330 (feminine) [as subgenus of Leptocera Olivier, 1813; nom. n. for Biroella Duda, 1925]. Type species: Limosina myrmecophila Knab & Malloch, 1912, automatic. - Richards, 1973: 330–352 [redescription, revision of Australian species, key, illustr.];

Marshall, 1989b: 602 [as genus; Australasian/Oceanian catalog]; Papp, 1995a: 540–552 [as genus; diagnosis, revision of Oriental species, key, illustr.].

Biroella Duda, 1925: 74 (feminine) [as subgenus of Leptocera Olivier, 1813; a junior homonym of Biroella Bolívar, 1903 (Saltatoria: Eumastacidae)]. Type species: Limosina myrmecophila

Knab & Malloch, 1912, monotypy. - Duda, 1938: 23 [as subgenus of Limosina Macquart,

1835]; Hackman, 1969a: 207 [as genus; biogeography]; Richards, 1973: 330 [homonymy].

Biróella. - Duda, 1925: 76 [incorrect original spelling].

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Note: For any catalog update, all species previously in Biroina need to be transferred to Howickia. The species below are newly described species only.

Howickia bicolor Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia bicolor Marshall et al. 2014: 9 [both sexes, illustr.]. Type locality: New Zealand, North Island, Coppermine Island, Hen and Chicken

Islands. HT male (NZAC).

Howickia exasperata Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia exasperata Marshall et al. 2014: 10 [unique male, illustr.]. Type locality: New Zealand, Moehau, Okahutahi Stream. HT male (NZAC).

Howickia harrisoni Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia Marshall et al. 2014: 11 [both sexes, illustr.]. Type locality: New Zealand, North

Island, Coromandel Peninsula, 10 km E Thames. HT male (NZAC).

Howickia lepidostylus Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia lepidostylus Marshall et al. 2014: 13 [both sexes, illustr.]. Type locality: New Zealand, Moehau Mt., Coromandel Range, bush on upper Okahutahi Stream. HT male (NZAC).

Howickia mercurialis Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia mercurialis Marshall et al. 2014: 15 [both sexes, illustr.]. Type locality: New Zealand, Mercury Islands, Green Island. HT male

(NZAC).

Howickia nigrilegula Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia nigrilegula Marshall et al. 2014: 17 [both sexes, illustr.]. Type locality: New Zealand, Trounson Kauri Park, 11 km S of Waipoua SF.

HT male (NZAC).

Howickia nigriventer Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia nigriventer Marshall et al. 2014: 17 [both sexes, illustr.]. Type locality: New Zealand, North Island, Coromandel Peninsula, 10 km E

Thames. HT male (NZAC).

Howickia nudistylus Marshall 2014. Distr.: Australasian/Oceanian: New Zealand. - 294 -

Howickia nudistylus Marshall et al. 2014: 19 [unique male, illustr.]. Type locality: New Zealand, North Island, Te Paki. HT male (NZAC).

Howickia oliveri Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia oliveri Marshall et al. 2014: 20 [both sexes, male illustr.]. Type locality: New Zealand, Moehau, Okahutahi Stream. HT male (NZAC).

Howickia omamari Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia omamari Marshall et al. 2014: 20 [both sexes, illustr.]. Type locality: New Zealand, Northland, Omamari Beach Marsh. HT male

(NZAC).

Howickia regalis Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia regalis Marshall et al. 2014: 22 [both sexes, illustr.]. Type locality: New Zealand, Three Kings Island. HT male (NZAC).

Howickia cordata Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia cordata Marshall et al. 2014: 26 [both sexes, illustr.]. Type locality: New Zealand, North

Island, Tararua Range, Dundas Hut area. HT male (NZAC).

Howickia palmai Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia palmai Marshall et al. 2014: 28 [both sexes, illustr.]. Type locality: New Zealand, Mt. Kaukau, Wellington. HT male (NMNZ).

Howickia tangata Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia tangata Marshall et al. 2014: 28 [both sexes, illustr.]. Type locality: New Zealand, North Island, Putara Valley, 10 km W Eketahuna.

HT male (NZAC).

Howickia wahaika Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia wahaika Marshall et al. 2014: 35 [both sexes, illustr.]. Type locality: New Zealand, “Upper Maitai”. HT male (NZAC).

Howickia zonula Marshall 2014. Distr.: Australasian/Oceanian: New Zealand.

Howickia zonula Marshall et al. 2014: 35 [both sexes, illustr.]. Type locality: New Zealand, Big South Cape Island. HT male (NZAC).

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Genus Minilimosina

Subgenus Svarciella

NOTE SU ET AL PROVIDE KEYS AND NEW RECORDS FOR SEVERAL PREVIOUSLY DESCRIBED

SPECIES

Minilimosina gracilenta Su 2015. Distr.: Oriental: China.

Minilimosina gracilenta Lixin et al. 2015: 13 [unique male, illustr.]. Type locality: China, Jiangxi, Mt. Wuyi. HT male (SACS).

Minilimosina luteola Su, 2011. Distr.: Oriental: China.

Minilimosina luteola Su, 2011: 75 [males, illustr.]. Type locality: China, Yunnan, County Tengchong, Mt. Laifeng, HT male (SACS).

Minilimosina luteola.—Su et al. 2013c: 18–19 [redescription, illustr.].

Minilimosina obtusispina Su 2013. Distr.: Oriental: China.

Minilimosina obtusispina Su et al. 2013c: 19 [male, description, illustr.]. Type locality: China, Jiangxi, Guanshan, Donghe, HT male (SACS).

Minilimosina parafanta Su 2015. Distr.: Oriental: China.

Minilimosina parafanta Lixin et al. 2015:20 [both sexes, illustr.]. Type locality: China, Zhejiang, Mt. Tianmu, Grand Canyon, Qianmutian. HT male (SACS).

Minilimosina tapiehella Su 2015. Distr.: Oriental: China.

Minilimosina tapiehella Lixin et al. 2015: 22 [both sexes, illustr.]. Type locality: China, Hubei, Mt. Ta-pieh, County Luotian, Qingtaiguan. HT male (SACS).

Subgenus Allolimosina

Minilimosina cerciseta Su, 2011. Distr.: Oriental: China.

Minilimosina cerciseta Su 2011: 69 [males, illustr.]. Type locality: China, Hebei, Mt. Xiaowutai, Mt. Dongling, HT male (SACS).

Subgenus Minilimosina

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Minilimosina quadrispinosa Su, 2011. Distr.: Oriental: China.

Minilimosina quadrispinosa Su, 2011: 72 [males, illustr.]. Type locality: China, Liaoning, Mt. Qianshan, HT male (SACS).

Genus Mislocatus Papp 2014b

Mislocatus, Papp 2014b: 111 (masculine). Type species: Ceroptera ealensis Vanschuybroeck, original designation. – Papp (2014b): 111–116

[diagnosis, description, illustr.].

Genus Nearcticorpius

Nearcticorpus palaearctictum Su 2012. Distr.: Oriental: China.

Nearcticorpus palaearctictum Su et al. 2012: 342 [males, illustr.]. Type locality: China, Ningxia Hui Autonomous Region, Mt. Liupan, Longtan.

HT male (SACS).

Genus Oligochaetosella Papp 2014b

Oligochaetosella, Papp 2014b: 116 (feminine). Type species: Oligochaetosella inconspicua, original designation. – Papp (2014b): 116–118

[diagnosis, description, illustr.].

Oligochaetosella inconspicua Papp, 2014b. Distr.: Afrotropical: Ghana.

Oligochaetosella inconspicua Papp 2014b: 118 [both sexes, illustr.]. Type locality: Ghana, Tamale, 25 km on Damaongo road. HT male

(HNHM).

Genus Paralimosina Papp 1973

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Paralimosina australis Papp 2015. Distr.: Afrotropical: Burundi, Malawi, South Africa.

Paralimosina australis Papp 2015: 321 [both sexes, illustr.]. Type locality: South Africa, Eastern Cape, Hogsback, Wolf Ridge Road. HT male

(HNHM).

Paralimosina congoensis Papp 2015. Distr.: Afrotropical: Republic of Congo.

Paralimosina congoensis Papp 2015: 325 [unique female, illustr.]. Type locality: Republic of Congo, Sibiti. HT female (HNHM).

Paralimosina flavifacies Papp 2015. Distr.: Afrotropical: Tanzania.

Paralimosina flavifacies Papp 2015: 317 [both sexes, illustr.]. Type locality: Tanzania, Tanga, Amani. HT male (HNHM).

Paralimosina heteronerua Papp 2015. Distr.: Afrotropical: South Africa.

Paralimosina heteronerua Papp 2015: 327 [both sexes, illustr.]. Type locality: South Africa, KwaZulu-Natal, Southern Drakensberg, reedy meadow along Mlambonja River. HT male (HNHM).

Paralimosina paraustralis Papp 2015. Distr.: Afrotropical: South Africa.

Paralimosina paraustralis Papp 2015: pp [unique male, illustr.]. Type locality: South Africa, KwaZulu-Natal, Eshowe, Ngoye Forest Reserve.

HT male (NMSA).

Paralimosina sinelineata Papp 2015. Distr.: Afrotropical: South Africa.

Paralimosina sinelineata Papp 2015: 330 [both sexes, illustr.]. Type locality: South Africa, Eastern Cape, forest near R102 road. HT male

(HNHM).

Genus Paramosina Marshall & Yau, 2014

Paramosina Marshall & Yau, 2014: 394 (feminine). Type species: Paramosina hirsuta, original designation. – Marshall & Yau (2014): 394

[diagnosis, description, illustr.].

Paramosina hirsuta Marshall & Yau 2014. Distr.: Neotropical: Ecuador.

Paramosina hirsuta Marshall & Yau 2014: 394 [both sexes, illustr.]. Type locality: Ecuador, Pichincha: Cotopaxi Natl. Pk., Quebrada

Mishahuaicu. HT male (QCAZ). - 298 -

Genus Permixtolimosina Papp 2014b

Permixtolimosina, Papp 2014b: 119 (feminine). Type species: Permixtolimosina sexsetosa, original designation. – Papp (2014b): 119–122

[diagnosis, description, illustr.].

Permixtolimosina sexsetosa Papp, 2014b. Distr.: Afrotropical: South Africa.

Permixtolimosina sesetosa Papp 2014b: 122 [both sexes, illustr.]. Type locality: South Africa, KwalaZulu-Natal, Ndumu Game Reserve, main campe area. HT male (BMSA).

Genus Phthitia Enderlein, 1938

Phthitia basilata Su, 2011. Distr.: Oriental: China.

Phthitia basilata Su, 2011: 85 [both sexes, illustr.]. Type locality: China, Ningxia, Liupanshan, Heshangpu. HT male (SACS).

Phthitia globosa Su, 2013. Distr.: Oriental: China.

Phthitia globosa Su et al. 2013a: 159 [unique male, illustr.]. Type locality: China, Ningxia, Liupanshan, Longtan. HT male (SACS).

Phthitia longidigita Su, 2011. Distr.: Oriental: China.

Phthitia longidigita Su, 2011: 89 [unique male, illustr.]. Type locality: China, Liaoning, Jianchang, Bailangshan. HT male (SACS).

Phthitia longula Su, 2013. Distr.: Oriental: China.

Phthitia longula Su et al. 2013a: 163 [unique male, illustr.]. Type locality: China, Yunnan, Dali, Cangshan. HT male (SACS).

Phthitia pollex Su, 2011. Distr.: Oriental: China.

Phthitia pollex Su, 2011: 84 [males, illustr.]. Type locality: China, Liaoning, Jianchang, Bailangshan. HT male (SACS).

Phthitia sternipilis Su, 2013. Distr.: Oriental: China. - 299 -

Phthitia sternipilis Su et al. 2013a: 167 [both sexes, illustr.]. Type locality: China, Yunnan, Dali, Cangshan. HT male (SACS).

Genus Preepiphallus Papp 2014b

Preepiphallus, Papp 2014b: 123 (masculine). Type species: Preepiphallus nitidifacies, original designation. – Papp (2014b): 123 [diagnosis, description, illustr.].

Preepiphallus endrodyi Papp, 2014b. Distr.: Afrotropical: Ghana.

Biphallapodema oligodentata Papp 2014b: 123 [males, illustr.]. Type locality: Ghana, Ho, Abuadi-Kpeze, c. 50 km. HT male (HNHM).

Preepiphallus nitidifacies Papp, 2014b. Distr.: Afrotropical: Malawi, South Africa.

Preepiphallus nitidifacies Papp 2014b: 124 [both sexes, illustr.]. Type locality: Malawi, Ntchisi Forest Reserve. HT male (NMSA).

Genus Pseudocollinella Duda

Papp 2016 provides a key to the Oriental species

Subgenus Pseudocollinella Duda

Pseudocollinella marshalli Papp, 2016. Distr.: Oriental: Mongolia.

Pseudocollinella marshalli Papp, 2016: 5 [both sexes, illustr.]. Type locality: Mongolia, Uvs aimak, am Fluss Baruun-turuun gol neben Somon

Baruun-turuun. HT male (HNHM).

Pseudocollinella mongolica Papp, 2016. Distr.: Oriental: Mongolia.

Pseudocollinella mongolica Papp, 2016: 7 [both sexes, illustr.]. Type locality: Mongolia, Bajan Ölgij aimak, im Tal des Flusses Chavcalyn gol,

25 km O von Somon Cagannuur. HT male (HNHM).

Pseudocollinella pseudohumida Papp, 2016. Distr.: Oriental: China.

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Pseudocollinella pseudohumida Papp, 2016: 16 [both sexes, illustr.]. Type locality: China, Fragrant Hill [Xiangshan] Park, West Mountains Nat.

Reserves – 40 km NW Beijing. HT male (HNHM).

Subgenus Setiopacifrons Papp 2016

Setiopacifrons Papp, 2016: 19 (feminine) [as subgenus of Pseudocollinella]. Type species: Pseudocollinella dupliciseta Duda, 1925. Papp, 2016, original designation. – Papp (2016): 19–57 [diagnosis, key to Afrotropical & extra-Afrotropical species, illustr.].

Pseudocollinella communis Papp, 2016. Distr.: Oriental: Thailand.

Pseudocollinella communis Papp, 2016: 30 [males, illustr.]. Type locality: Thailand, Mae Fang N.P., over & along a forest brook. HT male

(HNHM).

Pseudocollinella congoana Papp, 2016. Distr.: Afrotropical: Burundi, Democratic Republic of Congo.

Pseudocollinella Papp, 2016: 20 [both sexes, illustr.]. Type locality: D. R. Congo: Oriental Prov., Likombo forest, 2 km SW Bomona. HT male

(IRSN).

Pseudocollinella formosensis Papp, 2016. Distr.: Oriental: Taiwan.

Pseudocollinella formosensis Papp, 2016: 36 [males, illustr.]. Type locality: Taiwan: Kaohsiung Hsien, Liukuei, Shan Ping LTER Site - creek valley, No. 13. HT male (HNHM).

Pseudocollinella japonica Papp, 2016. Distr.: Oriental: Japan.

Pseudocollinella japonica Papp, 2016: 36 [unique male, illustr.]. Type locality: Japan, Kyushu Is., Oike, Mt. Kurodake area.. HT male (HNHM).

Pseudocollinella koreana Papp, 2016. Distr.: Oriental: Korea.

Pseudocollinella koreana Papp, 2016: 40 [unique male, illustr.]. Type locality: Korea, Prov. Ryang-gang, Plateau Chann-pay, San-zi-yan. HT male (HNHM).

Pseudocollinella normalis Papp, 2016. Distr.: Afrotropical: Tanzania.

Pseudocollinella normalis Papp, 2016: 23 [males, illustr.]. Type locality: Tanzania, Muyuni, Morogoro reg.. HT male (HNHM).

Pseudocollinella paradupliciseta Papp, 2016. Distr.: Oriental: Taiwan.

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Pseudocollinella paradupliciseta Papp, 2016: 42 [both sexes, illustr.]. Type locality: Taiwan, Kaohsiung Hsien, Liukuei, Shan Ping LTER Site - over/along a creek. HT male (HNHM).

Pseudocollinella pilitibia Papp, 2016. Distr.: Oriental: Thailand.

Pseudocollinella pilitibia Papp, 2016: 44 [males, illustr.]. Type locality: Thailand, Nan Prov., Ban Na Lae nr Pua. HT male (HNHM).

Pseudocollinella prima Papp, 2016. Distr.: Afrotropical: Democratic Republic of Congo, Ghana, Namibia.

Pseudocollinella prima Papp, 2016: 23 [both sexes, illustr.]. Type locality: Ghana, Banda Nkwanta. HT male (HNHM).

Pseudocollinella setipuga Papp, 2016. Distr.: Oriental: Thailand.

Pseudocollinella setipuga Papp, 2016: 44 [males, illustr.]. Type locality: Thailand, Trang Prov., Ban Liphang. HT male (HNHM).

Pseudocollinella setisternalis Papp, 2016. Distr.: Afrotropical: South Africa.

Pseudocollinella setisternalis Papp, 2016: 25 [unique male, illustr.]. Type locality: Republic of South Africa, KwaZulu-Natal, Ndumo

Game R.. HT male (BMSA).

Pseudocollinella simplicisternum Papp, 2016. Distr.: Oriental: Taiwan.

Pseudocollinella simplicisternum Papp, 2016: 48 [unique male, illustr.]. Type locality: Taiwan: Kaohsiung Hsien, Liukuei, Shan Ping LTER Site- creek valley. HT male (HNHM).

Pseudocollinella tercia Papp, 2016. Distr.: Oriental: Thailand.

Pseudocollinella tercia Papp, 2016: 50 [males, illustr.]. Type locality: Thailand: Mae Fang N.P.. HT male (HNHM).

Pseudocollinella trifida Papp, 2016. Distr.: Oriental: Taiwan.

Pseudocollinella trifida Papp, 2016: 50 [unique male, illustr.]. Type locality: Taiwan: Taipei, Nanshih Chiao, Ha Lo-Da. HT male (HNHM).

Pseudocollinella vietnamensis Papp, 2016. Distr.: Oriental: Vietnam.

Pseudocollinella vietnamensis Papp, 2016: 52 [males, illustr.]. Type locality: Vietnam: O-qui-ho. HT male (HNHM).

Pseudocollinella vulnerata Papp, 2016. Distr.: Afrotropical: Burundi, Democratic Republic of Congo, Tanzania.

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Pseudocollinella vulnerata Papp, 2016: 27 [both sexes, illustr.]. Type locality: D.R. CONGO: Oriental Prov., Eyolo forest, ca. 2 km E Lieki. HT male (IRSNB).

Genus Rudolfina Roháček 1987

Rudolfina zhangi Su, 2017. Distr.: Oriental: China (JIL).

Rudolfina zhangi Su et al., 2017: 3 [males, illustr.]. Type locality: CHINA: Jilin Prov., Mountains Changbai. HT male (SACS).

Genus Subacuminiseta Papp 2014b

Subacuminiseta, Papp 2014b: 127 (feminine). Type species: Subacuminiseta minor, original designation. – Papp (2014b): 127–128 [diagnosis, description, illustr.].

Subacuminiseta minor Papp, 2014b. Distr.: Afrotropical: Ghana.

Subacuminiseta minor Papp 2014b: 129 [both sexes, illustr.]. Type locality: Ghana, Ho, Abuadi-Kpeze, c. 50 km. HT male (HNHM).

Genus Telomerina

Telomerina curvibasata Su, 2013. Distr.: Oriental: China.

Telomerina curvibasata Su et al. 2013d: 8 [males, illustr.]. Type locality: China, Jiangxi, Mt. Guan, River Dong. HT male (SACS).

Telomerina laterispinata Su, 2013. Distr.: Oriental: China.

Telomerina laterispinata Su et al. 2013d: 11 [males, illustr.]. Type locality: China, Jiangxi, Mt. Guan, River Dong. HT male (SACS).

Telomerina levicana Su, 2013. Distr.: Oriental: China.

Telomerina levicana Su et al. 2013d: 12 [males, illustr.]. Type locality: China, Jiangxi, Mt. Guan. HT male (SACS).

Telomerina tuberculata Su, 2013. Distr.: Oriental: China.

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Telomerina tuberculata Su et al. 2013d: 14 [males, illustr.]. Type locality: China, Jiangxi, Mt. Guan, River Dong. HT male (SACS).

Genus Volumosina Rohacek & Marshall 2017

Volumosina Rohacek & Marshall, 2017: ## (female). Type species: Volumosina voluminosa, original designation. – Rohacek & Marshall (2017):

444–460 [diagnosis, description, illustr.].

Volumosina voluminosa (Marshall, 1987). Distr.: Nearctic: Canada (ON), USA (NH).

Herniosina voluminosa Marshall 1987: 711 [both sexes, phylogenetic notes, illustr.]. Type locality: USA, New Hampsire, Coos Co., 3 mi NE

East Inlet Dam, Norton Pool. HT male (CNCI).

A1.2 DESCRIBED COPROMYZINAE SINCE LAST CATALOG UPDATE

Genus Dudaia Hedicke, 1923

Note – Papp and Norrbom 2015 give new distributional records of previously published species, including issues on the identification of some species.

Dudaia abdita Papp & Norrbom 2015. Distribution: Afrotropical: Kenya.

Dudaia abdita Papp & Norrbom, 2015: 38, [both sexes, illustr.]. Type locality: Kenya, Suam fishing hut, Mt. Elgon.. HT male (MNHN).

Dudaia aethiopica Papp & Norrbom 2015. Distribution: Afrotropical: Ethiopia.

Dudaia aethiopica Papp & Norrbom, 2015: 40, [unique male, illustr.]. Type locality: Ethiopia,. HT male (AMNH).

Dudaia albimana Papp & Norrbom 2015. Distribution: Afrotropical: Madagascar.

Dudaia albimana Papp & Norrbom, 2015: 43, [both sexes, illustr.]. Type locality: Madagascar, Prov. Fianarantsoa, 7 km W Ranomafana. HT male (USNM).

Dudaia brevis Papp & Norrbom 2015. Distribution: Afrotropical: Madagascar. - 304 -

Dudaia brevis Papp & Norrbom, 2015: 45, [unique male, illustr.]. Type locality: Madagascar, 30 Km West Fort Dauphin. HT male (USNM).

Dudaia communis Papp & Norrbom 2015. Distribution: Afrotropical: Democratic Republic of Congo, Republic of

Conto, Ghana, Nigeria, South Africa.

Dudaia communis Papp & Norrbom, 2015: 47, [both sexes, illustr.]. Type locality: Ghana, Kwadaso. HT male (HNHM).

Dudaia malagasiensis Papp & Norrbom 2015. Distribution: Afrotropical: Madagascar.

Dudaia malagasiensis Papp & Norrbom, 2015: 50, [both sexes, illustr.]. Type locality: Madagascar, Antsingy de Bekopaka Inst. Scient.

Madagascar. HT male (MNHN).

Dudaia microtuberculata Papp & Norrbom 2015. Distribution: Afrotropical: South Africa.

Dudaia microtuberculata Papp & Norrbom, 2015: 52, [both sexes, illustr.]. Type locality: South Africa, KwaZulu

Natal, Kosi Bay Nat. Res., picnic area. HT male (BMSA).

Dudaia pseudohumeralis Papp & Norrbom 2015. Distribution: Afrotropical: Democratic Republic of Congo.

Dudaia pseudohumeralis Papp & Norrbom, 2015: 58, [both sexes, illustr.]. Type locality: Democratic Republic of Congo, Kivu, Rutshuru (riv.

Fuku). HT male (IRSN).

Dudaia spangleri Papp & Norrbom 2015. Distribution: Afrotropical: Kenya.

Dudaia spangleri Papp & Norrbom, 2015: 60, [males, illustr.]. Type locality: Kenya, Ngong Forestry Station. HT male (USNM).

Dudaia steineri Papp & Norrbom 2015. Distribution: Afrotropical: Madagascar.

Dudaia steineri Papp & Norrbom, 2015: 61, [both sexes, illustr.]. Type locality: Madagascar, Prov. Fianarantsoa, 7 km W Ranomafana. HT male

(USNM).

A1.3 DESCRIBED SPHAEROCERINAE SINCE LAST CATALOG UPDATE

Genus Ischiolepta Lioy

Ischiolepta paradraskovitsae Su, Liu and Xu 2016. Distribution: Oriental: China.

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Ischiolepta paradraskovitsae Su, Liu & Xu, 2016: 2, [male, illustr.]. Type locality: China, Zhejiang, Tianmushan, Grand Canyon, Qianmutian.

HT male (SACS).

A1.4 DESCRIBED ARCHIBORBORINAE SINCE LAST CATALOG UPDATE

See Kits & Marshall 2011 and Kits & Marshall 2015. Antrops – 40 new species; Boreantrops – 31 new species,

Coloantrops – 1 new species, Maculantrops – 1 new species, Photoantrops – 1 new species, Poeciloantrops – 10 new species.

Antrops – 40 species

Antrops anovariegatus Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops anovariegatus Kits & Marshall, 2013: 20 [both sexes, illustr.]. Type locality: Ecuador, Napo, Lago Papllacta, nr. HT male (QCAZ).

Antrops aurantifemur Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops aurantifemur Kits & Marshall, 2013: 21 [both sexes, illustr.]. Type locality: Ecuador, Napo, Lago Papallacta, nr. HT male (QCAZ).

Antrops baeza Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops baeza Kits & Marshall, 2013: 22 [both sexes, illustr.]. Type locality: Ecuador, Napo, Baeza, 15 km NW. HT male (QCAZ).

Antrops bellavista Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops bellavista Kits & Marshall, 2013: 77 [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Bellavista Reserve. HT male (QCAZ).

Antrops biflavus Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops biflavus Kits & Marshall, 2013: 51 [unique male, illustr.]. Type locality: Ecuador, Loja, Cajanuma, Podocarpus Natl. Pk., trail Los

Miradores. HT male (UTPL).

Antrops bucki Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

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Antrops bucki Kits & Marshall, 2013: 24 [both sexes, illustr.]. Type locality: Ecuador, Napo/Pichincha, Papllacta Pass, 0°19'15"S 78°11'51"W.

HT male (QCAZ).

Antrops carpishensis Kits & Marshall, 2013. Distr.: Neotropical: Peru.

Antrops carpishensis Kits & Marshall, 2013: 53 [both sexes, illustr.]. Type locality: Peru, Huanuco, Paso Carpish, vic. Chinchao, 9°43'S 76°4'W.

HT male (FFMNH).

Antrops cochabamba Kits & Marshall, 2013. Distr.: Neotropical: Bolivia, Peru.

Antrops cochabamba Kits & Marshall, 2013: 62 [both sexes, illustr.]. Type locality: Bolivia, Cochabamba, Siberia, W. Comarapa. HT male

(CNCI).

Antrops cochinoca Kits & Marshall, 2013. Distr.: Neotropical: Argentina.

Antrops cochinoca Kits & Marshall, 2013: 68 [unique male, illustr.]. Type locality: Argentina, Jujuy, Cochinoca. HT male (DEBU).

Antrops coniobaptos Kits & Marshall, 2013. Distr.: Neotropical: Bolivia.

Antrops coniobaptos Kits & Marshall, 2013: 53 [males, illustr.]. Type locality: Bolivia, Cochabamba, along Hwy. 7, 17°14.28'S 65°53.37'W. HT male (FMNH).

Antrops coroico Kits & Marshall, 2013. Distr.: Neotropical: Bolivia.

Antrops coroico Kits & Marshall, 2013: 79 [both sexes, illustr.]. Type locality: Bolivia, La Paz, Coroico, Cero Uchumachi, 16°12'43"S

67°42'49"W. HT male (UASC).

Antrops cotopaxi Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops cotopaxi Kits & Marshall, 2013: 62 [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Cotopaxi Natl. Pk., Quebrada Mishahuaicu.

HT male (QCAZ).

Antrops didactylos Kits & Marshall, 2013. Distr.: Neotropical: Argentina, Chile.

Antrops didactylos Kits & Marshall, 2013: 27 [both sexes, illustr.]. Type locality: Chile, Maule, Altos del Lircay Natl. Res.. HT male (MNNC).

Antrops diversipennis Kits & Marshall, 2013. Distr.: Neotropical: Ecuador. - 307 -

Antrops diversipennis Kits & Marshall, 2013: 27 [both sexes, illustr.]. Type locality: Ecuador, Napo, Lago Papallacta, nr.. HT male (QCAZ).

Antrops fulginosus Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops fulginosus Kits & Marshall, 2013: 32 [both sexes, illustr.]. Type locality: Ecuador, Napo, Quito–Baeza road. HT male (QCAZ).

Antrops guandera Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops guandera Kits & Marshall, 2013: 72 [both sexes, illustr.]. Type locality: Ecuador, Carchi, Guandera For. Res., 15 km E San Gabriel. HT male (QCAZ).

Antrops guaramacalensis Kits & Marshall, 2013. Distr.: Neotropical: Venezuela.

Antrops guaramacalensis Kits & Marshall, 2013: 54 [both sexes, illustr.]. Type locality: Venezuela, Trujillo, Guaramacal Natl. Pk., 19.3 SE

Bocono, 9°14'11"N 70°11'7"W,. HT male (MIZA).

Antrops inca Kits & Marshall, 2013. Distr.: Neotropical: Bolivia, Peru.

Antrops inca Kits & Marshall, 2013: 81 [both sexes, illustr.]. Type locality: Peru, Cusco, Puente Pilco, ~5.3 km NNW Challabamba, along creek,

13°10'53"S 71°46'08"W,. HT male (MUSM).

Antrops juninensis Kits & Marshall, 2013. Distr.: Neotropical: Peru.

Antrops juninensis Kits & Marshall, 2013: 69 [unique male, illustr.]. Type locality: Peru, Junin, Ondores. HT male (MZLU).

Antrops manu Kits & Marshall, 2013. Distr.: Neotropical: Peru.

Antrops manu Kits & Marshall, 2013: 81 [unique male, illustr.]. Type locality: Peru, Cuzco, Paucartamba, Buenos Aires, km 132. HT male

(USNM).

Antrops mucarensis Kits & Marshall, 2013. Distr.: Neotropical: Chile.

Antrops mucarensis Kits & Marshall, 2013: 71 [both sexes, illustr.]. Type locality: Chile, Antofagasta, Mucar, on Argentina border. HT male

(CNCI).

Antrops niger Kits & Marshall, 2013. Distr.: Neotropical: Venezuela.

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Antrops niger Kits & Marshall, 2013: 39 [both sexes, illustr.]. Type locality: Venezuela, Merida, Apartaderos, Laguna Negra. HT male (FMNH).

Antrops papallacta Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops papallacta Kits & Marshall, 2013: 82 [both sexes, illustr.]. Type locality: Ecuador, Napo, Lago Papllacta, nr.. HT male (QCAZ).

Antrops pecki Kits & Marshall, 2013. Distr.: Neotropical: Bolivia, Colombia, Ecuador.

Antrops pecki Kits & Marshall, 2013: 65 [both sexes, illustr.]. Type locality: Ecuador, Napo, Quito–Baeza road, Papllacta. HT male (QCAZ).

Antrops podocarpus Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops podocarpus Kits & Marshall, 2013: 56 [both sexes, illustr.]. Type locality: Ecuador, Loja, Cajanuma, Podocarpus Natl. Pk., trail Los

Miradores. HT male (UTPL).

Antrops quadrilobus Kits & Marshall, 2013. Distr.: Neotropical: Colombia, Ecuador, Venezuela.

Antrops quadrilobus Kits & Marshall, 2013: 83 [both sexes, illustr.]. Type locality: Ecuador, Carchi, Bosque El Arrayan, 6 km E San Gabriel. HT male (QCAZ).

Antrops siberia Kits & Marshall, 2013. Distr.: Neotropical: Bolivia.

Antrops siberia Kits & Marshall, 2013: 57 [males, illustr.]. Type locality: Bolivia, Santa Cruz, Yungas de la Siberia, 26.4 km NW Comarapa,

17°49'37"S 64°39'11"W. HT male (UASC).

Antrops sierrazulensis Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops sierrazulensis Kits & Marshall, 2013: 83 [both sexes, illustr.]. Type locality: Ecuador, Napo, SierrAzul Res., 14 km W Cosanga. HT male (QCAZ).

Antrops tachira Kits & Marshall, 2013. Distr.: Neotropical: Venezuela.

Antrops tachira Kits & Marshall, 2013: 85 [both sexes, illustr.]. Type locality: Venezuela, Tachira, San Cristobal, 55 km NE. HT male (MIZA).

Antrops tequendama Kits & Marshall, 2013. Distr.: Neotropical: Colombia.

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Antrops tequendama Kits & Marshall, 2013: 57 [unique male, illustr.]. Type locality: Colombia, Cundimarca, C. Amara, Tequendama. HT male

(CNCI).

Antrops tetrastichus Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops tetrastichus Kits & Marshall, 2013: 73 [both sexes, illustr.]. Type locality: Ecuador, Napo, Quito–Baeza pass. HT male (QCAZ).

Antrops tumbrensis Kits & Marshall, 2013. Distr.: Neotropical: Chile.

Antrops tumbrensis Kits & Marshall, 2013: 72 [both sexes, illustr.]. Type locality: Chile, Antofagasta, Tubre. HT male (CNCI).

Antrops unduavi Kits & Marshall, 2013. Distr.: Neotropical: Bolivia.

Antrops unduavi Kits & Marshall, 2013: 75 [both sexes, illustr.]. Type locality: Bolivia, La Paz, Unduavi, 16°19'S 67°54'W,. HT male (UASC).

Antrops variegatus Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Antrops variegatus Kits & Marshall, 2013: 49 [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Tandapi, 35 km E. HT male (QCAZ).

Antrops versabilis Kits & Marshall, 2013. Distr.: Neotropical: Venezuela.

Antrops versabilis Kits & Marshall, 2013: 66 [both sexes, illustr.]. Type locality: Venezuela, Tachira, San Cristobal, 55 km NE. HT male

(MIZA).

Antrops vittatus Kits & Marshall, 2013. Distr.: Neotropical: Chile.

Antrops vittatus Kits & Marshall, 2013: 50 [both sexes, illustr.]. Type locality: Chile, Biobio, Laraquete, 37°10’S 73°10’W. HT male (MNNC).

Antrops yungas Kits & Marshall, 2013. Distr.: Neotropical: Bolivia, Peru.

Antrops yungas Kits & Marshall, 2013: 67 [both sexes, illustr.]. Type locality: Peru, Cusco, Wayqecha Biol. Stn., ~9 km NE Challabamba,

13°10'S 71°34'W. HT male (MUSM).

Antrops zongo Kits & Marshall, 2013. Distr.: Neotropical: Bolivia.

Antrops zongo Kits & Marshall, 2013: 85 [both sexes, illustr.]. Type locality: Bolivia, La Paz, Zongo Valley, 16°07'57"S 68°06'56"W. HT male

(UASC).

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Genus Boreantrops Kits & Marshall 2013

Boreantrops, Kits & Marshall 2013: 86 (masculine). Type species: Boreantrops mexicanus Steyskal, original designation. – Kits & Marshall

(2013): 127–128 [diagnosis, description, illustr.]. Kits & Marshall 2015 [redescription, key, revision, species, illus.]

Note: there are no distributional records for the previously described species included in either paper (2013,

2015) but they are newly transferred to the genus in Kits & Marshall 2013.

Boreantrops albipes Kits & Marshall, 2015. Distr.: Neotropical: Costa Rica.

Boreantrops albipes Kits & Marshall, 2015: 311 [both sexes, illustr.]. Type locality: Costa Rica, Alajuela, San Gerardo Biol. Stn. 10°52'51"N

85°23'20"W. HT male (INBC).

Boreantrops alytothrix Kits & Marshall, 2015. Distr.: Nearctic: Mexico.

Boreantrops alytothrix Kits & Marshall, 2015: 313[both sexes, illustr.]. Type locality: Mexico, Jalisco, Atenquique, 18 mi. W. HT male (DEBU).

Boreantrops apterus Kits & Marshall, 2015. Distr.: Neotropical: Costa Rica, Panama.

Boreantrops apterus Kits & Marshall, 2015: 313 [both sexes, illustr.]. Type locality: Panama, Chiriqui, Cerro Punta, 5 km ESE. HT male

(DEBU).

Boreantrops auranticeps Kits & Marshall, 2015. Distr.: Neotropical: Ecuador.

Boreantrops auranticeps Kits & Marshall, 2015: 334 [both sexes, illustr.]. Type locality: Ecuador, Napo, SierrAzul Res., 14 km W Cosanga,

0°40'55"S 77°56'69"W. HT male (QCAZ).

Boreantrops avignis Kits & Marshall, 2015. Distr.: Neotropical: Costa Rica, Venezuela.

Boreantrops avignis Kits & Marshall, 2015: 314 [both sexes, illustr.]. Type locality: Costa Rica, Alajuela, San Gerardo Biol. Stn, 10°52'51"N

85°23'20"W. . HT male (INBC).

Boreantrops boliviensis Kits & Marshall, 2015. Distr.: Neotropical: Bolivia.

- 311 -

Boreantrops boliviensis Kits & Marshall, 2015: 336 [both sexes, illustr.]. Type locality: Bolivia, La Paz, Cubre Alto Beni, 28 km E Caranavi,

15°40'31"S 67°29'21"W. HT male (UASC).

Boreantrops challabamba Kits & Marshall, 2015. Distr.: Neotropical: Peru.

Boreantrops challabamba Kits & Marshall, 2015: 337 [both sexes, illustr.]. Type locality: Peru, Cusco, Puente Pilco, 5.3 km NNW Challabamba.

HT male (MUSM).

Boreantrops costaricensis Kits & Marshall, 2015. Distr.: Neotropical: Costa Rica.

Boreantrops costaricensis Kits & Marshall, 2015: 316 [both sexes, illustr.]. Type locality: Costa Rica, Puntarenas, Monteverde Biol. Res.. HT male (INBC).

Boreantrops cryptopygium Kits & Marshall, 2015. Distr.: Neotropical: Bolivia.

Boreantrops cryptopygium Kits & Marshall, 2015: 354 [both sexes, illustr.]. Type locality: Bolivia, La Paz, Chulumani, Apa Apa Reserve,

16°21'15"S 67°30'21"W. HT male (UASC).

Boreantrops durango Kits & Marshall, 2015. Distr.: Nearctic: Mexico.

Boreantrops durango Kits & Marshall, 2015: 317 [both sexes, illustr.]. Type locality: Mexico, Durango, El Salto, 13 mi. W. HT male (CNCI).

Boreantrops emarginatus Kits & Marshall, 2015. Distr.: Neotropical: Guatemala, Mexico.

Boreantrops emarginatus Kits & Marshall, 2015: 338 [both sexes, illustr.]. Type locality: Guatemala, Guatemala, Santa Catarina Pinula. HT male

(DEBU).

Boreantrops friburguensis Kits & Marshall, 2015. Distr.: Neotropical: Brazil.

Boreantrops friburguensis Kits & Marshall, 2015: 318 [both sexes, illustr.]. Type locality: Brazil, Rio de Janeiro, Nova Friburgo, 10 km S Sitio

Edelweiss, Muri. HT male (MZSP).

Boreantrops guatemalensis Kits & Marshall, 2015. Distr.: Neotropical: Guatemala, Mexico.

Boreantrops guatemalensis Kits & Marshall, 2015: 320 [both sexes, illustr.]. Type locality: Guatemala, Zacapa, San Lorenzo, 7 km N. HT male

(DEBU).

Boreantrops hispidus Kits & Marshall, 2015. Distr.: Neotropical: Brazil. - 312 -

Boreantrops hispudus Kits & Marshall, 2015: 321 [both sexes, illustr.]. Type locality: Brazil, Minas Gerais, Barbacena, 20 km SW. HT male

(MZSP).

Boreantrops hondurensis Kits & Marshall, 2015. Distr.: Neotropical: Guatemala, Honduras.

Boreantrops hondurensis Kits & Marshall, 2015: 321 [both sexes, illustr.]. Type locality: Honduras, Francisco Morazan, Uyaca. HT male

(DEBU).

Boreantrops inbio Kits & Marshall, 2015. Distr.: Neotropical: Costa Rica, Guatemala, Honduras, Mexico,

Nicaragua, Panama.

Boreantrops inbio Kits & Marshall, 2015: 323 [both sexes, illustr.]. Type locality: Costa Rica, Cartago, Tapanti Natl. Pk., above Ranger Stn. HT male (INBC).

Boreantrops longiphallus Kits & Marshall, 2015. Distr.: Neotropical: El Salvador, Guatemala, Mexico.

Boreantrops longiphallus Kits & Marshall, 2015: 325 [both sexes, illustr.]. Type locality: Guatemala, Guatemala, Santa Catarina Pinula. HT male

(DEBU).

Boreantrops machinator Kits & Marshall, 2015. Distr.: Neotropical: Peru.

Boreantrops machinator Kits & Marshall, 2015: 339 [males, illustr.]. Type locality: Peru, Cusco, Wayqecha Biol. Stn., ~9 km NE Challabamba,

13°10'S 71°34'W. HT male (MUSM).

Boreantrops masneri Kits & Marshall, 2015. Distr.: Neotropical: Venezuela.

Boreantrops masneri Kits & Marshall, 2015: 326 [males, illustr.]. Type locality: Venezuela, Aragua, Henri Pittier Natl. Pk., Maracay-Choroni highway, km 18. HT male (MIZA).

Boreantrops oaxacensis Kits & Marshall, 2015. Distr.: Neoarctic: Mexico.

Boreantrops oaxacensis Kits & Marshall, 2015: 328 [both sexes, illustr.]. Type locality: Mexico, Oaxaca, 1.7 mi. W Jct Mex. 175-Yuvila Rd. HT male (DEBU).

Boreantrops peruvianus Kits & Marshall, 2015. Distr.: Neotropical: Peru.

Boreantrops peruvianus Kits & Marshall, 2015: 329 [both sexes, illustr.]. Type locality: Peru, Loreto, Teniente Lopez. HT male (DEBU).

- 313 -

Boreantrops pollex Kits & Marshall, 2015. Distr.: Neotropical: Ecuador.

Boreantrops pollex Kits & Marshall, 2015: 341 [both sexes, illustr.]. Type locality: Ecuador, Napo, Baeza. HT male (QCAZ).

Boreantrops punctipennis Kits & Marshall, 2015. Distr.: Neotropical: Bolivia.

Boreantrops punctipennis Kits & Marshall, 2015: 342 [both sexes, illustr.]. Type locality: Bolivia, La Paz, Coroica, Cerro Uchumachi,

16°12'43"S 67°42'49"W. HT male (UASC).

Boreantrops subemarginatus Kits & Marshall, 2015. Distr.: Neotropical: Ecuador, Venezuela.

Boreantrops subemarginatus Kits & Marshall, 2015: 343 [both sexes, illustr.]. Type locality: Venezuela, Merida, Tabay, La Mucuy, Truchicola trail. HT male (MIZA).

Boreantrops subfoveolatus Kits & Marshall, 2015. Distr.: Neotropical: Panama.

Boreantrops subfoveolatus Kits & Marshall, 2015: 330 [both sexes, illustr.]. Type locality: Panama, Chiriqui, Cerro Punta, 5 km ESE. HT male

(DEBU).

Boreantrops suchixtepecensis Kits & Marshall, 2015. Distr.: Nearctic: Mexico.

Boreantrops suchixtepecensis Kits & Marshall, 2015: 332 [both sexes, illustr.]. Type locality: Mexico, Oaxaca [San Miguel] Suchixtepec, 8 km

S. HT male (DEBU).

Boreantrops talamanca Kits & Marshall, 2015. Distr.: Neotropical: Costa Rica, Panama.

Boreantrops talamanca Kits & Marshall, 2015: 345 [both sexes, illustr.]. Type locality: Panama, Chiriqui, Cerro Punta, 2 km E. HT male

(DEBU).

Boreantrops wayqecha Kits & Marshall, 2015. Distr.: Neotropical: Peru.

Boreantrops wayqecha Kits & Marshall, 2015: 346 [both sexes, illustr.]. Type locality: Peru, Cusco, Wayqecha Biol. Stn., ~9km NE

Challabamba, 13°10'S 71°35'W. HT male (MUSM).

Boreantrops zacapa Kits & Marshall, 2015. Distr.: Neotropical: Guatemala.

Boreantrops zacapa Kits & Marshall, 2015: 333 [both sexes, illustr.]. Type locality: Guatemala, Zacapa, San Lorenzo, 7 km N. HT male

(DEBU).

- 314 -

Boreantrops zamora Kits & Marshall, 2015. Distr.: Neotropical: Ecuador.

Boreantrops zamora Kits & Marshall, 2015: 348 [both sexes, illustr.]. Type locality: Ecuador, Zamora Chinchipe, San Francisco, Res. Biol. San

Francisco, trail Canal, 3°58'30"S 79°4'25"W. HT male (UTPL).

Genus Coloantrops Kits & Marshall, 2013

Coloantrops Kits & Marshall, 2013: 87 (masculine). Type species: Coloantrops daedalus, original designation. – Kits & Marshall (2013): 87–89

[diagnosis, description, illustr.].

Coloantrops daedalus Kits & Marshall, 2011. Distr.: Neotropical: Chile.

Coloantrops daedalus Kits & Marshall, 2011: 87 [both sexes, illustr.]. Type locality: Chile, Los Lagos, Alerce Andino Natl. Pk., trail to Laguna

Fria, 41°30'S 72°37'W. HT male (MNNC).

Genus Maculantrops Kits & Marshall, 2013

Maculantrops Kits & Marshall, 2013: 89 (masculine). Type species: Borborus hirtipes, original designation. – Kits & Marshall (2013): 89–90

[diagnosis, description, key, illustr.].

Maculantrops altiplanus Kits & Marshall, 2013. Distr.: Neotropical: Bolivia.

Maculantrops altiplanus Kits & Marshall, 2013: 90 [both sexes, illustr.]. Type locality: Bolivia, La Paz, La Paz, 15 km NE, 16°24.6'S 68°02.9'W.

HT male (UASC).

Genus Photoantrops Kits & Marshall, 2013

Photoantrops Kits & Marshall, 2013: 92 (masculine). Type species: Photoantrops echinus, original designation. – Kits & Marshall (2013): 93–93

[diagnosis, description, illustr.].

- 315 -

Photoantrops echinus Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Photoantrops echinus Kits & Marshall, 2013: 93 [both sexes, illustr.]. Type locality: Ecuador, Napo, SierrAzul Lodge, 14 km W Cosanga. HT male (QCAZ).

Genus Poeciloantrops Kits & Marshall, 2013

Poeciloantrops Kits & Marshall, 2013: 95 (masculine). Type species: Poeciloantrops baorucensis, original designation. – Kits & Marshall

(2013): 95–96 [diagnosis, description, key, illustr.].

Poeciloantrops baorucensis Kits & Marshall, 2013. Distr.: Neotropical: Dominican Republic.

Poeciloantrops baorucensis Kits & Marshall, 2013: 96 [both sexes, illustr.]. Type locality: Dominican Republic, Pedernales, Las Abejas, 30 km

N of Caba Rojo. HT male (DEBU).

Poeciloantrops boraceiensis Kits & Marshall, 2013. Distr.: Neotropical: Brazil.

Poeciloantrops boraceiensis Kits & Marshall, 2013: 97 [both sexes, illustr.]. Type locality: Brazil, Sao Paulo, USP Biology Station. HT male

(MZSP).

Poeciloantrops crocidosternum Kits & Marshall, 2013. Distr.: Neotropical: Brazil.

Poeciloantrops crocidosternum Kits & Marshall, 2013: 98 [both sexes, illustr.]. Type locality: Brazil, Sao Paulo, Est. Biol. Boraceia. HT male

(MZSP).

Poeciloantrops dominicus Kits & Marshall, 2013. Distr.: Neotropical: Dominican Republic.

Poeciloantrops dominicus Kits & Marshall, 2013: 99 [both sexes, illustr.]. Type locality: Dominican Republic, Independencia, La Descubierta, 32 km NW, Sabana Real. HT male (DEBU).

Poeciloantrops flavifemur Kits & Marshall, 2013. Distr.: Neotropical: Bolivia.

Poeciloantrops flavifemur Kits & Marshall, 2013: 99 [both sexes, illustr.]. Type locality: Bolivia, La Paz, Coroico. HT male (UASC).

Poeciloantrops marensis Kits & Marshall, 2013. Distr.: Neotropical: Brazil.

Poeciloantrops marensis Kits & Marshall, 2013: 101 [both sexes, illustr.]. Type locality: Brazil, Parana, Curitiba, Sitio H. HT male (DZUP).

- 316 -

Poeciloantrops plaumanni Kits & Marshall, 2013. Distr.: Neotropical: Brazil.

Poeciloantrops plaumanni Kits & Marshall, 2013: 101 [both sexes, illustr.]. Type locality: Brazil, Santa Catarina, Nova Teutonia, 27°10'S

52°22'W. HT male (FMNH).

Poeciloantrops psilosternum Kits & Marshall, 2013. Distr.: Neotropical: Brazil.

Poeciloantrops psilosternum Kits & Marshall, 2013: 102 [both sexes, illustr.]. Type locality: Brazil, Sao Paulo, Barueri. HT male (DZUP).

Poeciloantrops stellans Kits & Marshall, 2013. Distr.: Neotropical: Ecuador.

Poeciloantrops stellans Kits & Marshall, 2013: 103 [both sexes, illustr.]. Type locality: Ecuador, Napo, Quito–Baeza pass. HT male (QCAZ).

Poeciloantrops vittifrons Kits & Marshall, 2013. Distr.: Neotropical: Brazil.

Poeciloantrops vitifrons Kits & Marshall, 2011: 104 [both sexes, illustr.]. Type locality: Brazil, Santa Catarina, Nova Teutonia, 27°10'S 52°22'W.

HT male (DEBU).

A1.5 PENDING SPECIES DESCRIPTIONS

The following Limosininae species are in preparation and are not complete citations, as they have yet to be published. Confirmation of the details is required after time of publication. Currently there are 28 Archiceroptera, 20

Bromeloecia, 1 Pectinosina, 9 Rudolfina, and 10 “Sabogramma” to be described based on the following entries.

Archiceroptera Papp

Note: there are new distributional records for both previously described species.

Archiceroptera adamas Paiero & Marshall, in prep. Distr.: Neotropical: French Guiana, Guyana.

Archiceroptera adamas Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: French Guiana, Maripasoula, Mitaraka, MIT-C-TOP,

2°13'59"N, 54°26'38"W. HT male (MHNM).

- 317 -

Archiceroptera addenda Paiero & Marshall, in prep. Distr.: Neotropical: Bolivia, Ecuador, French Guiana, Guyana,

Panama, Peru.

Archiceroptera addenda Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Ecuador, Esmeraldas, La Chiquita, 11 km SE San

Lorenzo. HT male (QCAZ).

Archiceroptera barberi Paiero & Marshall, in prep. Distr.: Neotropical: Belize, Colombia, Costa Rica, Ecuador,

Guatemala, Honduras, Mexico, Panama, Trinidad & Tobago, Venezuela.

Archiceroptera barberi Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Panama, Chiriquí, Hartmann's Finca, 15 km NW Hato de Volcán. HT male (DEBU).

Archiceroptera basilia Paiero & Marshall, in prep. Distr.: Neotropical: Ecuador, Peru.

Archiceroptera basilia Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, Cosanga, 2.5 km W, 0°35'24"S,

77°53'19"W. HT male (QCAZ).

Archiceroptera bilobata Paiero & Marshall, in prep. Distr.: Neotropical: Ecuador.

Archiceroptera bilobata Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Ecuador, Pinchincha, Alluriquin, 23km E, Chiriboyo

Ret.. HT male (QCAZ).

Archiceroptera bisetosus Paiero & Marshall, in prep. Distr.: Neotropical: Bolivia.

Archiceroptera bisetosus Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Bolivia, La Paz, Cumbre Alto Beni, 28 km E

Caranavi. HT male (UASC).

Archiceroptera braziliensis Paiero & Marshall, in prep. Distr.: Neotropical: Brazil.

Archiceroptera braziliensis Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Brazil, Paraná, Curitiba, 30 km SE, BR 277. HT male (MZSP).

Archiceroptera brevivilla Paiero & Marshall, in prep. Distr.: Neotropical: Guatemala, Mexico (CHI, HID, SLP,

VER).

Archiceroptera brevivilla Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Mexico, Hidalgo, Tlanchinol, 2.5mi N. HT male

(FMNH).

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Archiceroptera browni Paiero & Marshall, in prep. Distr.: Neotropical: Ecuador.

Archiceroptera browni Paiero & Marshall, in prep: pp [unique male, illustr.]. Type locality: Ecuador, Pichincha, Río Palenque Stn., 47 km S

Santo Domingo. HT male (QCAZ).

Archiceroptera caliga Paiero & Marshall, in prep. Distr.: Neotropical: Brazil, Colombia, Costa Rica, Ecuador,

Panama, Peru.

Archiceroptera caliga Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Rio Palenque. HT male (QCAZ).

Archiceroptera calligraphia Paiero & Marshall, in prep. Distr.: Neotropical: Belize, Costa Rica, Ecuador,

Guatemala, Honduras, Mexico.

Archiceroptera calligraphia Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Mexico, Chiapas, Lagunas de Montebello Parque

Nacional, Aqua Tinta. HT male (FMNH).

Archiceroptera cobolorum Paiero & Marshall, in prep. Distr.: Neotropical: Brazil, Colombia, Ecuador, Peru.

Archiceroptera cobolorum Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, Baeza, 15km NW. HT male

(QCAZ).

Archiceroptera crenulata Paiero & Marshall, in prep. Distr.: Neotropical: Colombia, French Guiana, Venezuela.

Archiceroptera crenulata Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Colombia, Leticia. HT male (DEBU).

Archiceroptera curvavilla Paiero & Marshall, in prep. Distr.: Neotropical: Costa Rica, Ecuador, Panama.

Archiceroptera curvavilla Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Rio Palenque. HT male

(QCAZ).

Archiceroptera dolabra Paiero & Marshall, in prep. Distr.: Neotropical: French Guiana.

Archiceroptera dolabra Paiero & Marshall, in prep: pp [males, illustr.]. Type locality: French Guiana, St. Laurent du Maroni, Maripasoula,

Mitaraka, MIT-DZ, 2°14'2"N,7 54°27'1"W. HT male (MHNM).

Archiceroptera llama Paiero & Marshall, in prep. Distr.: Neotropical: Guatemala, Mexico (CHI).

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Archiceroptera llama Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Guatemala, Sacatepéquez, Volcán Atitlán, Ref. Quetzal,

14°33'2"N, 91°11'32"W. HT male (UVGC).

Archiceroptera maniba Paiero & Marshall, in prep. Distr.: Neotropical: Guyana, Venezuela.

Archiceroptera maniba Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Guyana, Potaro-Siparuni, Mount Wokomung,

5°6'35"N, 59°49'15"W. HT male (ROME).

Archiceroptera masoni Paiero & Marshall, in prep. Distr.: Neotropical: Mexico (SIN).

Archiceroptera masoni Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Mexico, Sinaloa, Concordia, 20mi. E. HT male

(UNAM).

Archiceroptera megacercus Paiero & Marshall, in prep. Distr.: Neotropical: Bolivia, Brazil, Colombia, Costa Rica,

Ecuador, French Guiana, Guyana, Peru, Venezuela.

Archiceroptera megacercus Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, Tena, 12 km SW. HT male

(QCAZ).

Archiceroptera megavilla Paiero & Marshall, in prep. Distr.: Neotropical: Argentina, Costa Rica, Ecuador, Mexico

(CHI), Panama, Peru, Trinidad & Tobago, Venezuela.

Archiceroptera magavilla Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Panama, Chiriquí, Cerro Punta, 2 km W. HT male

(DEBU).

Archiceroptera mexicorona Paiero & Marshall, in prep. Distr.: Neotropical: Mexico (GUE).

Archiceroptera mexicorona Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Mexico, Guerro, Ixtapa, 45km NE. HT male

(UNAM).

Archiceroptera mitaraki Paiero & Marshall, in prep. Distr.: Neotropical: French Guiana.

Archiceroptera mitaraki Paiero & Marshall, in prep: pp [males, illustr.]. Type locality: French Guiana, St. Laurent du Maroni, Maripasoula,

Mitaraka, MIT-DZ-RBF1, 2°14'4"N, 54°27'2"W. HT male (MNHM).

Archiceroptera paracercus Paiero & Marshall, in prep. Distr.: Neotropical: Costa Rica, Ecuador.

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Archiceroptera paracercus Paiero & Marshall, in prep: pp [males, illustr.]. Type locality: Ecuador, Esmeraldas, La Chiquita, 11 km SE San

Lorenzo. HT male (QCAZ).

Archiceroptera pussula Paiero & Marshall, in prep. Distr.: Neotropical: Argentina, Bolivia, Ecuador.

Archiceroptera pussula Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Bolivia, Santa Cruz, Potrerillos de Guenda,

17°40'29"S, 63°27'22"W. HT male (UASC).

Archiceroptera ternum Paiero & Marshall, in prep. Distr.: Neotropical: Bolivia, Brazil, Costa Rica, Ecuador,

French Guiana, Panama, Peru, Venezuela.

Archiceroptera ternum Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Rio Palenque. HT male (QCAZ).

Archiceroptera triclavus Paiero & Marshall, in prep. Distr.: Neotropical: Brazil, Guyana.

Archiceroptera triclavus Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Brazil, Bahia, Porto Segure, 15 km NE,

Ecological Reserva “Pau-Brasil”. HT male (MZSP).

Archiceroptera uncinata Paiero & Marshall, in prep. Distr.: Neotropical: Bolivia, Colombia, Ecuador, French

Guiana, Guyana, Peru, Trinidad & Tobago, Venezuela.

Archiceroptera uncinata Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Venezuela, Bolivar, km40 Sta. Elena Icabaru Road.

HT male (MIZA).

Bromeloecia abundantia Yau & Marshall, 2018. Distr.: Nearctic: U.S.A. (AZ); Neotropical: Argentina, Brazil,

Belize, Bolivia, Colombia, Costa Rica, Ecuador, Guatemala, Guyana, Honduras, Mexico, Panama, Paraguay, Peru,

Venezuela.

Bromeloecia abundantia Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Rio Palenque. HT male (QCAZ).

Bromeloecia aculatus Yau & Marshall, 2018. Distr.: Neotropical: Belize, Bolivia, Brazil, Colombia, Costa Rica,

Cuba, Dominican Republic, Ecuador, Guatemala, Guyana, Mexico, Panama, Peru, Puerto Rico Trinidad & Tobago,

Venezuela. - 321 -

Bromeloecia aculatus Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Zamora-Chinchipe, Zamora, Podocarpus National

Park, Bombuscaro, El Mirador trail. HT male (UTPL).

Bromeloecia aurita Yau & Marshall, 2018. Distr.: Neotropical: Belize, Brazil, Costa Rica, Ecuador, Panama.

Bromeloecia aurita Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, 17 km NE Baeza. HT male (XXXX).

Bromeloecia balaena Yau & Marshall, 2018. Distr.: Neotropical: Bolivia, Ecuador.

Bromeloecia balaena Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Bolivia, La Paz, Coroico, Cerro Uchumachi, 16°12'43"S,

67°42'49"W. HT male (UASC).

Bromeloecia brachium Yau & Marshall, 2018. Distr.: Neotropical: Belize, Bolivia, Colombia, Costa Rica,

Ecuador, Guyana, Mexico, Panama, Peru, Trinidad & Tobago, Venezuela.

Bromeloecia brachium Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, Jatun Sacha Res., 6km E Misahuallí, 1°4'S,

77°37'W. HT male (QCAZ).

Bromeloecia cercarcuata Yau & Marshall, 2018. Distr.: Neotropical: Bolivia, Ecuador.

Bromeloecia cercarcuata Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, Tena, 12 km SW. HT male (QCAZ).

Bromeloecia coniclunis Yau & Marshall, 2018. Distr.: Neotropical: Bolivia, Ecuador, Peru, Venezuela.

Bromeloecia Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Bolivia, La Paz, Coroico, Cerro Uchumachi, 16°12'43"S,

67°42'49"W. HT male (UASC).

Bromeloecia diabolunguia Yau & Marshall, 2018. Distr.: Neotropical: Colombia, Venezuela.

Bromeloecia diabolunguia Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Colombia, Norte de Santander, 20 mi S Cucuta Qbda..

HT male (IAVH).

Bromeloecia ephippium Yau & Marshall, 2018. Distr.: Neotropical: Belize, Bolivia, Brazil, Costa Rica, Cuba,

Dominican Republic, Ecuador, Guatemala, Guyana, Mexico (CHI, OAX, VER), Panama, Peru, Trinidad & Tobago,

Venezuela.

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Bromeloecia ephippium Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Guyana, Mazaruni-Potaro, Tukeit Falls, Potaro River. HT male (DEBU).

Bromeloecia fractacincta Yau & Marshall, 2018. Distr.: Neotropical: Ecuador, Guyana, Venezuela.

Bromeloecia fractacincta Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Guyana, Potaro-Saparuni, Mount Wokomung, 5°6'35"N,

59°49'15"W. HT male (ROME).

Bromeloecia magna Yau & Marshall, 2018. Distr.: Neotropical: Costa Rica, Panama.

Bromeloecia magna Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Costa Rica, San Jose, Moravia Zurqui de Moravia, 10°2'58"N,

84°0'57"W. HT male (INBC).

Bromeloecia peloris Yau & Marshall, 2018. Distr.: Neotropical: Guyana, Venezuela.

Bromeloecia peloris Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Guyana, Potaro-Siparuni, Mount Wokomung, 5°6'35"N,

59°49'15"W. HT male (DEBU).

Bromeloecia pinna Yau & Marshall, 2018. Distr.: Neotropical: Ecuador.

Bromeloecia pinna Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, Baeza. HT male (QCAZ).

Bromeloecia ponsa Yau & Marshall, 2018. Distr.: Neotropical: Argentina, Bolivia, Ecuador, .

Bromeloecia ponsa Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, Lago Papallacta, 0°20'29"S, 78°10'23"W. HT male (QCAZ).

Bromeloecia ramus Yau & Marshall, 2018. Distr.: Neotropical: Bolivia, Colombia, Ecuador, Guyana, Peru,

Venezuela.

Bromeloecia ramus Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, Tiputini Biodiversity Stn., 0°36'50"S,

76°9'1"W. HT male (DEBU?).

Bromeloecia robustora Yau & Marshall, 2018. Distr.: Neotropical: Bolivia, Colombia, Ecuador, Peru, Trinidad and

Tobago, Venezuela.

Bromeloecia robustora Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Napo, Jatun Sacha Res., 6 km E Misahualli,

1°4'S, 77°37'W. HT male (QCAZ). - 323 -

Bromeloecia spathicercus Yau & Marshall, 2018. Distr.: Neotropical: Belize, Costa Rica, Guatemala, Mexico

(CHI, HID, OAX, VER).

Bromeloecia spathicercus Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Mexico, Veracruz, 4 mi N Huatusco. HT male (UNAM).

Bromeloecia triunguia Yau & Marshall, 2018. Distr.: Neotropical: Belize, Costa Rica, Ecuador, Guyana.

Bromeloecia triunguia Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Pichincha, Rio Palenque. HT male (QCAZ).

Bromeloecia undulata Yau & Marshall, 2018. Distr.: Nearctic: Mexico; Neotropical: Barbados, Brazil, Bolivia,

Colombia, Costa Rica, Ecuador, Grenada, Guatemala, Guyana, Honduras, Mexico, Panama, Paraguay, Peru,

Trinidad & Tobago, Venezuela.

Bromeloecia undulata Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Esmeraldas, 11 km SE San Lorenzo, La Chiquita.

HT male (QCAZ).

Bromeloecia wolverinei Yau & Marshall, 2018. Distr.: Neotropical: Ecuador, Panama, Puerto Rico, St. Kitts &

Nevis, Trinidad.

Bromeloecia wolverinei Yau & Marshall, 2018: pp [both sexes, illustr.]. Type locality: Ecuador, Galapagos, Santa Cruz 2 km N Bellavista. HT male (QCAZ).

Pectinosina carro Paiero & Marshall, in prep. Distr.: Neotropical:.

Pectinosina carro Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality:. HT male (XXXX).

Rudolfina Roháček

Note: there are new distributional records of previously described species included in the thesis.

Rudolfina bucki Paiero & Marshall, in prep. Distr.: Nearctic: Mexico (OAX).

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Rudolfina bucki Paiero & Marshall, in prep: pp [males, illustr.]. Type locality: Mexico, Oaxaca, Jct. Mex. 175-Yuvila Rd., 4.1 mi W. HT male

(FMNH).

Rudolfina exuberata Paiero & Marshall, in prep. Distr.: Nearctic: U.S.A. (AL, FL, LA, MO, MS, NC, NM, OK,

SC, TN, TX). Neotropical:Argentina, Belize, Bolivia, Brazil, Chile, Colombia, Costa Rica, Ecuador, Guatemala,

Guyana, Honduras, Mexico (CAM, CHI, GUE, OAX, PUE, TAB, TAM, VER, YUC), Paraguay, Peru, Trinidad &

Tobago, Venezuela.

Rudolfina exuberata Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: U.S.A., Florida, Marion Co., Ocala National Forest. HT male (DEBU).

Rudolfina howdeni Paiero & Marshall, in prep. Distr.: Nearctic: Mexico (HID, JAL, MEX, MOR, OAX, VER).

Rudolfina howdeni Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Mexico, Oaxaca, Ixtlan de Juarez, 6.6mi. N. HT male

(FMNH).

Rudolfina megepandria Paiero & Marshall, in prep. Distr.: Nearctic: Mexico (JAL, OAX).

Rudolfina megepandria Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Mexico, Jalisco, Atenquique, 18 mi. W. HT male

(FMNH).

Rudolfina newtoni Paiero & Marshall, in prep. Distr.: Nearctic: Mexico (OAX).

Rudolfina newtoni Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality:Mexico, Oaxaca, Jct. Mex. 175-Yuvila Rd., 4.1mi. W. HT male (FMNH).

Rudolfina pauca Paiero & Marshall, in prep. Distr.: Nearctic: Guatemala, Mexico (HID, MEX, MOR).

Rudolfina pauca Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Mexico, Mexico, Tenancingo, 1mi NE. HT male (FMNH).

Rudolfina pilosa Paiero & Marshall, in prep. Distr.: Nearctic: Mexico (OAX).

Rudolfina pilosa Paiero & Marshall, in prep: pp [males, illustr.]. Type locality: Mexico, Oaxaca, Jct. Mex. 175–Yuvila Rd., 2.0 mi W. HT male

(FMNH).

Rudolfina remiforma Paiero & Marshall, in prep. Distr.: Nearctic: Mexico (CHI).

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Rudolfina remiforma Paiero & Marshall, in prep: pp [both sexes, illustr.]. Type locality: Mexico, Chiapas, Bochil, 21mi. N. HT male (FMNH).

Rudolfina tumida Paiero & Marshall, in prep. Distr.: Nearctic: U.S.A. (WY).

Rudolfina tumida Paiero & Marshall, in prep: pp [unique male, illustr.]. Type locality: U.S.A., WY, Uinta Co., Evanston, 8mi. SE. HT male

(DEBU).

A1.6 References for Appendix 1 (citations for pending species descriptions are not currently included):

Bergeron, M., Marshall, S.A. & Swann, J.E. (2015) A review of the New World Coproica (Diptera: Sphaeroceridae)

with a description of 8 new species. Zootaxa, 3953, 1–157.

Kits, J.H. & Marshall, S.A. (2013) Generic classification of the Archiborborinae (Diptera: Sphaeroceridae), with a

revision of Antrops Enderlein, Coloantrops gen. nov., Maculantrops gen. nov., Photoantrops gen. nov., and

Poecilantrops gen. nov. Zootaxa, 3701, 1–113.

Kits, J.H. & Marshall, S.A. (2015) A revision of Boreantrops Kits & Marshall (Diptera: Sphaeroceridae:

Archiborborinae). Zootaxa, 3915 (3), 301–355.

Luk, S. & Marshall, S.A. (2014) A revision of the New World genus Aptilotella Duda (Sphaeroceridae:

Limosininae). Zootaxa, 3761, 1–156.

Marshall, S.A. (2013) Bregmosina, a new Neotropical genus of Limosininae (Diptera: Sphaeroceridae). Zootaxa,

3641(3), 260–270.

Marshall, S.A. (2014) Albostyla, a new genus of Neotropical Limosininae (Diptera: Sphaeroceridae). Zootaxa, 3793,

257–264.

Marshall, S.A. & Yau, T. (2014) Paramosina, a new genus of high Andean Limosininae (Diptera: Sphaeroceridae).

Zootaxa, 3872(4), 393–397.

Papp, L. (1991) Oriental Limosininae: new species and records (Diptera, Sphaeroceridae). Acta Zoologica

Academiae Scientiarum Hungaricae, 37(3–4), 225–251.

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Papp, L. (2014a) A review of the Old World species of Ceroptera Macquart, 1835 (Diptera: Sphaeroceridae). Acta

Zoologica Academiae Scientiarum Hungaricae, 60(2): 109–155.

Papp, L. (2014b) New genera of Afrotropical limosinine sphaerocerids (Diptera: Sphaeroceridae). Zootaxa, 3764(2),

101–130.

Papp, L. (2015) The first record of the genus Paralimosina L. Papp (Diptera: Sphaeroceridae) in the Afrotropical

region with descriptions of six new species. African Invertebrates, 54(2), 315–333.

Papp, L. (2016) An overview of the Old World species of Pseudocollinella Duda (Diptera: Sphaeroceridae) with

description of a new subgenus. Acta Zoologica Academiae Scientiarum Hungaricae, 62(1), 1–58.

Roháček, J. (2016) Herniosina Roháček: revised concept, two new species, new key and atalas of male and female

terminalia (Diptera, Sphaeroceridae), Zookeys, 609, 69–106.

Roháček, J. & Marshall, S.A. (2017) Volumosina, a new Nearctic genus for the rare old-growth forest fly Herniosina

voluminosa Marshall (Diptera: Sphaeroceridae). The Canadian Entomologist, 149(4), 444–460.

Su, L.-X. 2011. Lesser Dung Flies. Liaoning University Press, Shenyang, Liaoning, China.

Su, L.-X., Liu, C., Xu, J., & Wang, J. (2012) A new speices of the genus Nearcticorpus Roháček and Marshall 1982

from China (Diptera: Sphaeroceridae). The Pan-Pacific Entomologist, 88(3), 342–346

Su., L.-X., Liu, G.-C. & Xu, J. (2013a) Phthitia Enderlein (Diptera: Sphaeroceridae: Limosininae) in China, with

descriptions of three new species. Journal of the Kansas Entomological Society, 86(2), 155–170.

Su, L., Liu, G. & Xu, J. (2013b) A new sphaerocerid Eulimosina prominulata sp. nov. (Diptera) from China.

Oriental Insects, 47(4), 199–202.

Su, L.X., Liu, G.C., Xu, J. & Wang, J.F. (2013c) The genus Minilimosina (Svarciella) (Sphaeroceridae: Diptera)

from China with description of a new species. Oriental Insects, 47(1), 15–22.

Su, L., Liu, G. & Xu, J. (2013) The genus Telomerina Roháček (Diptera Sphaeroceridae) from China, with the

descriptions of four new species. The Pan-Pacific Entomologist, 89(1), 7–17.

Su, L., Liu, G. & Xu, J. (2015) A new species of Ischiolepta Lioy (Diptera: Sphaeroceridae) from China. Oriental

Insects, 49(1–2), 1–5.

Su, L., Xu, J. & Cong, G. (2017) A new species of the genus Rudolfina Roháček, 1987 (Diptera, Sphaeroceridae)

from north-east China, with a key to the known Holarctic species of Rudolfina. Oriental Insects, 51(4),

391–396. - 327 -

Yau, T. and S.A. Marshall. 2018 (submitted manuscript). A review of the genus Bromeloecia Spuler (Diptera:

Sphaeroceridae). Zootaxa.

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