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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
University of Guelph, 2017
Advisor:
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).
iii
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
iv
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 -
v
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 -
vi
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 -
vii
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 -
viii
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
ix
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 -
x
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 -
xi
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 -
xii
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;
xiii
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 -
xiv
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 -
xv
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 -
xvi
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 -
xvii
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 -
xviii
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 -
xix
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 -
- 1 -
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.
- 2 -
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
- 3 -
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.
- 4 -
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.
- 5 -
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.
- 6 -
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
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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.
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2.5 Figures
Figure 2.1. Head chaetotaxy: lateral, Archiceroptera venezolana (Richards) (debu01077469), and dorsolateral,
A. browni n. sp. (debu01077561).
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Figure 2.2. Thoracic chaetotaxy of Archiceroptera mahukani (debu00295088).
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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.
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Figure 2.4. Wing of two members of the Archiceroptera genus complex. A) Archiceroptera venezolana
(debu00378968); B) Rudolfina exuberata (debu00276674).
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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).
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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.
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Figure 2.7. Archiceroptera venezolana phallus. A) dorsal view; B) dorsolateral view; and C) lateral view. A-C)
from debu00373840.
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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).
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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
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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
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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.
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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.
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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.
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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
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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
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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
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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.
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1
3.4 Tables and Figures 2
Table 3.1. Morphological character states for the epandrial process group (EPG). 3
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
mahukani group 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
Archiceroptera
(ternum-group) 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
Archiceroptera
(Extension group) 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
Archiceroptera
brevivilla-group 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
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
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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.
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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.
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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).
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Figure 3.4. Strict Consensus Tree (A) and Majority Rules Tree (B) from the morphological
analysis of the epandrial process group.
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Figure 3.5. Morphological phylogeny of the epandrial process group, including species groups
of the Archiceroptera genus complex. One of 11 equal length trees.
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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
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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.
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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.
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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,
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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,
- 36 -
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
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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,
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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;
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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,
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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,
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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
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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-
- 43 -
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.
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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
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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.
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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).
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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
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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.
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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.
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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.
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Figure 4.6. Distribution of Pectinosina prominens and P. carro.
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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]
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
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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
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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
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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
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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.
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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.
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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
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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
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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.
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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.
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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
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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
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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
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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.
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- 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
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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.
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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:
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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,
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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
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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
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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
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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.
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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
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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,
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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-
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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,
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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
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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.
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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.
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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
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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).
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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..
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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.
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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).
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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
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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.
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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).
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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,
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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.
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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.
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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).
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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.
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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).
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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).
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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).
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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).
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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).
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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).
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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).
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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).
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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).
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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).
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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).
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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
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(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
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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
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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
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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.
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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.
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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.
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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.
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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
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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.
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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
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- 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
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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
- 135 -
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
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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.
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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.
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- 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
- 139 -
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
- 140 -
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.
- 141 -
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).
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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.
- 143 -
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
- 144 -
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.
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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
- 146 -
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;
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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.
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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.
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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
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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).
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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.
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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
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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
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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.
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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.
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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
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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
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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
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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
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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,
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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,
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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
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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.
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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.
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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:
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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
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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:
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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
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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
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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
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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
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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.
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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,
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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-
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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).
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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
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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.
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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.
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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,
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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.
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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–
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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
- 183 -
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
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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.
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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
- 186 -
(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
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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.
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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
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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
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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
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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,
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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",
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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
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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-
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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–
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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
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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.
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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
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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,
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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.
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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.
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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:
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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
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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
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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
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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.
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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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6.10 Archiceroptera Tables and Figures 1
Table 6.1. Morphological character states for phylogenetic study of Archiceroptera. 2 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
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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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Transition Zone revisited: the distributional congruence patterns of Passalidae (Coleoptera). Invertebrate
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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Marshall, S.A. 1982. A review of Nearctic Limosininae (Diptera: Sphaeroceridae) with revisions of selected genera
(PhD Thesis). University of Guelph, Guelph, Ontario. 320p.
Marshall, S.A. 1983a. A revision of the genus Aptilotus Mik in North America (Diptera, Sphaeroceridae). Canadian
Journal of Zoology, 61: 1910–1924.
Marshall, S.A. 1983b. The genus Bromeloecia Spuler in North America (Diptera: Sphaeroceridae: Limosininae).
Proceedings of the Entomological Society of Washington 85: 32–35.
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.
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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–
216.
Marshall, S.A. 2000a. A new genus of ant-like sphaerocerid from Mexico (Diptera, Sphaeroceridae). Studia
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.
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.
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(Diptera) from Colorado and Arizona. Proceedings of the Entomological Society of Washington, 99: 641–
644.
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Sphaeroceridae, Limosininae). Contributions in Science, 474: 1–27.
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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.
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Agriculture Canada Monograph.
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Nearcticorpus (Diptera: Sphaeroceridae). Annals of the Entomological Society of America, 75: 642–648.
Marshall, S.A. and J. Roháček. 1984. A revision of the genus Telomerina Roháček (Diptera, Sphaeroceridae).
Systematic Entomology, 9: 127–163.
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Sphaeroceridae: Limosininae). Studia Dipterologica, 7: 259–311.
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Canadian Journal of Zoology 71: 835–857.
Marshall , S.A. and S. Totton. 1995. Seven new species of Druciatus Marshall (Diptera: Sphaeroceridae;
Limosininae). Insecta Mundi 9: 291–299.
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Sphaeroceridae). European Journal of Entomology, 104: 573–599.
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world catalog update covering the years 2000–2010, with new generic synonymy, new combinations, and
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Scientiarum Hungaricae, 54 (Suppl. 2): 47–209.
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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
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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.].
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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).
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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).
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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.
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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.].
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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.
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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).
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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
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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.
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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).
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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.
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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.
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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.
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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.
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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.
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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).
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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).
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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.].
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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).
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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).
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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.
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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).
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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.
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Yau, T. and S.A. Marshall. 2018 (submitted manuscript). A review of the genus Bromeloecia Spuler (Diptera:
Sphaeroceridae). Zootaxa.