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HARVARD UNIVERSITY
ave tians
Ernst Mayr Library
of the Museum of
Comparative Zoology
MCZ LIBRARY
JUL 24 2012
HARVARD | UNIVERSITY
MCZ_
LIBRARY r EB 5 t Que HAN RSITY JOURNAL of the ENTOMOLOGICAL»y
Volume One Hundred and Thirty-Eight 2007 :
Published November 2007
JOURNAL of the ENTOMOLOGICAL SOCIETY
ONTARIO. | - Volume One Hundred and Thirty-Eight 7 =") 2007 : ur F Published November 2007 THE ENTOMOLOGICAL SOCIETY OF ONTARIO
OFFICERS AND GOVERNORS 2006-2007 President: Webmaster: B. HELSON D. B. LYONS
Natural Resources Canada, Canadian Forest Service 1219 Queen St E., Sault Ste. Marie, ON P6A 2E5 bhelson@nrcan.gce.ca
President-Elect:
R. HALLETT
Dept. of Environmental Biology,
University of Guelph, Guelph, ON NIG 2W1 rhallett@uoguelph.ca
Past President:
J. HUBER Natural Resources Canada, Canadian Forest Service c/o Eastern Cereal and Oilseed Research Centre 960 Carling Ave., Ottawa, ON K1A 0C6 huberjh@agr.ge.ca
Secretary:
D. HUNT
Agriculture and Agri-Food Canada G.P.C.R.C. 2585 County Road 20, Harrow, ON NOR 1G0 huntd@agr.gc.ca
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Natural Resources Canada, Canadian Forest Service 1219 Queen St E., Sault Ste. Marie, ON P6A 2E5 kbarber@nrcan.gc.ca
Librarian:
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Library, University of Guelph
Guelph, ON NIG 2W1
Directors:
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Royal Ontario Museum, Toronto, Ontario
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4890 Victoria Avenue North
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J. SKEVINGTON (2007-2009) Agriculture and Agri-Food Canada 960 Carling Ave., Ottawa, ON K1A 0C6
L. TIMMS (2006-2008) Faculty of Forestry, University of Toronto
Toronto, ON MS5S 3B3
S. VANLAERHOVEN (2006-2008) Dept. of Biology, University of Windsor
Windsor, ON NOB 3P4.
ISSN 1713-7845
of
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Ontario Ministry of Agriculture, Food and Rural Affairs 1 Stone Road West, Guelph, ON NIG 4Y2
EDITORIAL COMMITTEE
Scientific Editor:
M. H. RICHARDS*
Dept. of Biological Sciences, Brock University St. Catharines, ON L2S 3A1 miriam.richards@brocku.ca
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R. HARMSEN
Biology Department, Queen’s University Kingston, ON N7L 3N6
Y. MAUFFETTE Faculté des sciences, Département des sciences biologiques _ Université 4 Québec Montréal, Montréal, QC H3C 3P8 J. SKEVINGTON
Agriculture and Agri-Food Canada
Eastern Cereal and Oilseed Research Centre
960 Carling Ave., Ottawa, ON K1A 0C6
~
*Please submit manuscripts electronically to the Editor (miriam.richards@brocku.ca). |
JESO Volume 138, 2007
JOURNAL of the | ENTOMOLOGICAL SOCIETY OF ONTARIO
VOLUME 138 2007
You will notice that there are two Editors’ Forwards in this volume of the Journal of the Entomological Society of Ontario. On page 2 is the forward from Special Editor Steve Marshall that was originally printed in Volume 137, which I have been thinking of as ‘Pengelly I’, with the first five papers in this volume comprising ‘Pengelly II’. About two years ago, we began planning a special volume in memory of David Pengelly, an entomologist at the University of Guelph who inspired many Ontario and Canadian entomologists. It is a mark of the esteem in which his students and colleagues held him, that the memorial volume had to be split into two. I would like to express my warmest thanks and appreciate to Steve Marshall who acted as the Special Editor of the Pengelly volumes.
With the publication of Volume 138, I am delighted to note that JESO has resumed a timely publication schedule. Not only is the 2007 volume actually being published in 2007, but preparations are already underway for Volume 139, to be published in 2008. Another important change is electronic distribution of published manuscripts, with the added attraction that the electronic versions may include colour plates and maps, which are generally impossibly expensive to print. Electronic distribution will enable our authors’ work to reach the wider audience that it deserves, not least of all because it will become accessible to web search engines. These are important developments in bringing JESO to a wider audience, and everyone who has helped to get us to this point, should award herself or himself a pat or two on the back. I especially thank the JESO Editorial Board, the ESO Board, our patient Technical Editor, and the many reviewers whose hard work and enthusiasm for the Journal, have been the real impetus for these changes.
After that moment of self-congratulation, let me assure you that we are not resting on our laurels, but intend to continue building on the momentum we have established. First, we are committed to maintaining annual publication in the autumn of each year (dare we dream of bi-annual publication?), so please submit your work to JESO — we are already considering manuscripts for Volume 139 (2008), and the more, the merrier. Remember, all entomological research is welcome, whether its subjects are in Ontario or elsewhere. Second, we are planning to begin converting back volumes of JESO into electronic format for distribution from the website — eventually we would like to post all volumes of JESO and its predecessor, the Proceedings of the ESO, right back to volume 1. Third, the success of these Pengelly volumes has inspired suggestions for future special volumes devoted to particular entomological themes. All these plans will help to ensure a bright and buggy future for the Journal.
Happy reading!
Miriam H. Richards Editor
JESO Volume 138, 2007
JOURNAL of the ENTOMOLOGICAL SOCIETY OF ONTARIO
VOLUME 138 2007
It has been a pleasure for me to serve as Special Editor for the D. H. Pengelly tribute volumes, and to work with regular Editor Miriam Richards to line up a diverse assemblage of papers touching on the range of entomological disciplines influenced by Dave Pengelly (1922-2004). Dave was widely known, and is fondly remembered, as a fantastic teacher of entomology and as a huge contributor to the development of the University of Guelph Insect Collection; but he is also sorely missed as a seemingly infinite reservoir of entomological — trivia, anecdotes, insightful stories, jokes, support, and wise advice. Almost all of the authors in these volumes were strongly influenced by Professor Pengelly, most shared his enthusiasm and passion for collecting and identifying insects, and many worked with Dave to build up the University of Guelph Insect Collection. Although now widely appreciated as a treasure trove of information about faunal change and insect distribution while also serving as the foundation for a growing body of taxonomic research, the University of Guelph Insect Collection during Pengelly’s tenure was a beleaguered resource used mostly as a source of specimens for a myriad of entomology courses including the hands-on third year entomology labs that Dave taught every weekday. It is a testimonial to his foresight that he not only kept the collection from deterioration or destruction due to invasions of dermestid beetles and indiscriminate colleagues, he consistently built up the collection and increased its curatorial level annually. He did not do this alone, of course. Professor Pengelly had a contagious vision of what was needed to improve our understanding of Ontario’s insects, and his enthusiasm for the study of insect biology and diversity was such that he was always associated with dedicated students. Thousands of specimen labels in the insect collection bear the names of most of the authors of papers in these volumes, standing as permanent testimonials to a shared vision that started with students working with their mentor at University of Guelph, and which continues through the widely dispersed teaching, research, and publications by Dave’s students and colleagues today. I think he would have liked these sets of papers, and he would have been pleased to see them published in the journal that he supported so selflessly ... almost every copy of this journal sent out during the 16 year period “D.H.” was Secretary-Treasurer of the ESO was sent out by Dave personally. We all wish he were still here to send this one out!
Steve Marshall Special Editor
Review of Eustochus JESO Volume 138, 2007
REVIEW OF EUSTOCH US, A RARELY COLLECTED GENUS OF MYMARIDAE (HYMENOPTERA)
J.T. HUBER’? and E. BAQUERO? Canadian Forestry Service, Natural Resources Canada
Abstract J. ent. Soc. Ont. 138: 3-31
The four nominal species of Eustochus are reviewed. Four new species, E. confusus from Spain, E. pengellyi and E. yoshimotoi from North America, and E. nipponicus from Japan are described and a key to the eight species is given.
Résumé Nous revisons les quatre espéces nominales d’Eustochus. Nous décrivons quatre nouvelles espeéces, E. confusus d Espagne; E. pengellyi et E. yoshimotoi d’Ameérique du Nord, et E. nipponicus du Japon, et pourvoyons une clé d identification aux huit espéces.
Published November 2007
Introduction
The Holarctic genus Eustochus was established by Haliday (1833) for a species described by Curtis (1832) ina key as a species of Mymar—M. atripennis (which he attributed to Walker). Curtis’s laconic description, abstracted here from his key, reads: ‘abdomen petiolated, wings ciliated and perfect, ovipositor exserted, longer than the abdomen’. The collection information given next to the name atripennis states simply ‘June, amongst grass in a wood’. Graham (1982) suggested that perhaps only one specimen had been collected by Walker, probably at or near Southgate, Middlesex (England). Haliday (1833) redescribed E. atripennis (Curtis) in more detail, and included it as the only species within his new genus Eustochus. He mentioned that the species occurred ‘in autumn, among trees, but very rare’. Since then, the genus has been mentioned infrequently in the literature and only three more species have been described, based on very few specimens each. Here, we review the described species and describe four new ones from Spain, Canada, USA, and Japan.
' Author to whom all correspondence should be addressed.
* Correspondence address: Systematic Entomology, K.W. Neatby Building, 960 Carling Avenue, Ottawa, Ontario, Canada K1A 0C6, email: huberjh@agr.gc.ca
> Departamento de Zoologia y Ecologia, Faculdad de Ciencias, Universidad de Navarra, 31080, Pamplona, Spain
Huber and Baquero JESO Volume 138, 2007
Materials and Methods
This study is based on recent examination of about 115 specimens from the institutions listed below.
Eustochus species all appear to be very similar to one another so long descriptions for each species, as for E. besucheti Bakkendorf, would be repetitious; therefore, diagnoses only are given. The new species are described and illustrated based on females because males are known for only two species and are exceedingly rare (7 specimens known). Specimens from Japan were dissected and gold coated for the scanning electron micrographs (Figs. 1 —]5).
Morphological terms used follow Gibson (1997). Measurements, from slide- mounted specimens unless otherwise indicated, are given in micrometers (uum), and are as described in Huber (1987). The range is followed by the number of specimens measured, in parentheses. Primary type data is recorded as found on each label, with slashes indicating the beginning of each new line. Data from other specimens is given in a standardized format to simplify retrieval or comparison. Abbreviations used are: Fl —Fl,=funicle segment one to six, FWL or FWW=forewing length or forewing width, Fl, L (W)=flagellar segment length (width).
Acronyms of repositories: CNC—Canadian National Collection of Insects, Ottawa, Canada, J. Huber; FAFU-—Biological Control Research Institute, Fujian Agricultural and Forestry University, Fuzhou, China, N.-Q. Lin; MRSN—Museo Regionale di Scienze Naturali (Spinola Collection), Turin, Italy, G. Pagliano; MZNA—Museo de Zoologia, Universidad de Navarra, Pamplona, Spain, E. Baquero; MHNG—Museum d’ Histoire Naturelle, Geneva, Switzerland, C. Besuchet; UCRC—University of California, Riverside, CA, USA, S. Triapitsyn; USNM-—National Museum of Natural History, Washington, DC, USA, M. Gates.
Eustochus Haliday
Eustochus Haliday, 1833: 269 (key), 349 (description); Westwood, 1839: 78 (diagnosis); Walker, 1846: 50 (diagnosis); Foerster, 1847: 225 (diagnosis); Foerster, 1856: 117 (key); Blanchard, 1840: 293 (diagnosis); Dalla Torre, 1898: 428 (catalogue); Ashmead, 1904: 363 (key); Schmiedeknecht, 1909: 494 (key), 495 (diagnosis); Gahan and Fagan, 1923: 64 (type species designation); Schmiedeknecht, 1930: 450 (key); Kryger, 1950: 61 (description); Annecke and Doutt, 1961: 24 (generic comment); Debauche, 1948: 200 (description); Schauff, 1984: 50 (diagnosis, phylogeny); Yoshimoto, 1990: 57 (diagnosis); Xu and Lin, 2003: 65 (diagnosis).
Type species: Mymar atripennis Curtis, by monotypy.
Diagnosis. Body 670-1330 um in length (critical point or air dried specimens); brown to dark brown, with appendages somewhat lighter in colour. Head (except face) and mesosoma, scape, dorsal surface of gastral petiole, coxae and femora with distinct reticulate sculpture
(Figs. 1-5, 7-10, 13, 15, 35, 36, 38-43); gaster, underside of petiole, and remainder of legs
4
Review of Eustochus JESO Volume 138, 2007
FIGURES 1-6. Head, Eustochus sp. probably atripennis (Japan, Mt. Tsukuba). 1—dorsal; 2-lateral 3—anterior; 4—ventral; 5—posterior; 6—mouthparts.
Huber and Baquero JESO Volume 138, 2007
100 um
100 um
100 um
20m 20 um
FIGURES 7-15. Mesosoma and metasoma, Eustochus sp. probably atripennis (Japan, Mt. Tsukuba). 7—mesosoma, dorsal; 8-mesosoma lateral; 9-mesosoma ventral; 10—metasoma, dorsal; 1 1—metasoma, lateral; 12—metasoma, ventral; |3—petiole dorsal; 14—petiole, ventral; 15—petiole, lateral.
Review of Eustochus JESO Volume 138, 2007
pe
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FIGURES 16-20. Eustochus spp., female antenna, lateral. 16—atripennis (Germany, Bornheim); 17—besucheti (Switzerland, Laquintal); 18—nearcticus (Canada, MacKenzies Mt.); 19-triclavatus (paratype); 20—pengellyi (holotype).
Huber and Baquero JESO Volume 138, 2007
FIGURES 21-25. Eustochus spp., female antenna (except 25), lateral. 21—yoshimotoi (holotype); 22—?nearcticus (USA, Andrews Bald); 23—confusus (holotype); 24—nipponensis (holotype); 25—atripennis, male (Switzerland, Chancy).
Review of Eustochus JESO Volume 138, 2007 en VOR D0, 200
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FIGURES 26-27. Eustochus spp., wings. 26—atripennis (Germany, Bornheim); 27— besucheti (Switzerland, Laquintal).
Huber and Baquero JESO Volume 138, 2007
28
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FIGURES 28-29. Eustochus spp., wings. 28—nearcticus (NS, MacKenzies Mt.); 29— triclavatus (paratype).
Review of Eustochus JESO Volume 138, 2007 i I a 10, 20
FIGURES 30-31. Eustochus spp., wings. 30—pengellyi (holotype); 31—voshimotoi (holotype).
11
JESO Volume 138, 2007
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Review of Eustochus JESO Volume 138. 2007 i a OREO EU
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FIGURES 34-36. Eustochus spp., wings and bodies. 34—nipponicus, wings (holotype): 35—atripennis, body lateral (Germany, Bornheim); 36—besucheti, mesosoma dorsal (Switzerland, Laquintal).
13
Huber and Baquero JESO Volume 138, 2007
0.1mm
FIGURES 37-40. Eustochus spp., bodies. 37—besucheti, metasoma lateral (Switzerland, Laquintal), 38—nearcticus, lateral (NS, MacKenzies Mt.); 39-triclavatus, mesosoma + metasoma, lateral (paratype); 40—pengellyi, mesosoma + metasoma, dorsal (holotype).
14
Review of Eustochus JESO Volume 138, 2007 I i 0,
FIGURES 41-43. Eustochus spp., bodies. 41—yoshimotoi, lateral (holotype); 42- ?nearcticus, mesosoma + metasoma, lateral (USA, Andrews Bald); 43—confusus, lateral
(holotype).
15
Huber and Baquero JESO Volume 138, 2007
FIGURES 44-46. Eustochus spp. 44—nipponensis, body lateral (holotype); 45—atripennis, male head, anterior (Switzerland, Chancy); 46—atripennis, male genitalia, ventrolateral.
16
Review of Eustochus JESO Volume 138, 2007
0.4mm
FIGURES 47-50. Eustochus confusus (paratype). 47-antenna; 48—clava: 49—wings (forewing damaged); 50—forewing venation.
Huber and Baquero JESO Volume 138, 2007
smooth (Figs. 10-12, 14). Legs with 4 tarsal segments, the basal one not much longer than each of the others.
Female. Eyes of normal size but ocelli small, the posterior ocelli not much larger than the mesh diameter of surrounding scupture (Figs. 1-3). Mandibles either relatively short and not quite meeting medially or occasionally slightly overlapping medially, with two (Fig. 6) or, usually, three teeth. Antennal funicle 6-segmented and clava 2-segmented (Figs. 16-18, 20-24) or, rarely, 3-segmented (Fig. 19); flagellum with longitudinal sensilla on fl, (1 sensillum), fl, (2 sensilla), fl, (2 sensilla) and clava (1 sensillum on basal segment, 5 on apical segment) [distribution of longitudinal sensilla on clava of E. triclavatus not clear due to collapse and poor orientation]. Forewing oval, with a more or less distinct, comma- shaped, dark mark extending from basal half of marginal vein to posterior margin just beyond retinaculum (Figs. 26-34); venation 0.35-0.4 times length of forewing; marginal + stigmal vein longer than submarginal vein; hypochaeta directly in front of proximal macrochaeta or slightly basal to it and two, widely separated, distal macrochaetae present, the second distal macrochaeta near apex of venation near base of short stigmal vein. Petiole long and narrow (Figs. 13-15, 38, 40, 42, 44). Gaster smooth, deep, and somewhat compressed, with gt, the largest tergum (Figs. 10, 11, 13, 38-44); spiracle present on gt,; dorsal one or two cercal setae extremely long and curved (Figs. 11, 13), distinctly longer than ventral two setae. Ovipositor either very long and conspicuously exerted beyond gastral apex, or short and not extending beyond gastral apex.
Male. Forewing minute and hindwing absent, with very small eyes and ocelli absent, and large, tridentate mandibles that meet medially (Fig. 45); body with reticulate sculpture less pronounced than in female. Antenna with 9 flagellar segments, the apical two widely joined to form a loose clava (Fig. 25).
Discussion. Eustochus is most closely related to Caraphractus Walker, another strictly Holarctic genus (Schauff 1984). Members of both genera have strongly sclerotized bodies with distinct reticulation on the head, mesosoma, petiole, coxae, and scape, and they have similar mesosomal, metasomal, and wing structure. Sclerotization, surface sculpture, and wings may be convergent due to the habitats in which the species of both genera are found. A well sclerotized body and strengthened wings are needed for protection while moving around in forest litter (Ewstochus) or water (Caraphractus). The heavy surface sculpture may trap air around the body. A host shift from terrestrial to semiaquatic to aquatic host may have occurred in some ancestral Eustochus species, leading to species that are now classified as Caraphractus, which parasitize only Dytiscidae, as far as is known. Though hosts for Eustochus are unknown, a few specimens of Eustochus have been collected near water (see E. atripennis, below) and the change from hosts in forest litter or soil to hosts near and eventually in water may easily have occurred.
Biology. Hosts are unknown. Most specimens of Eustochus examined were collected from deciduous forests. A few were collected near water, and one specimen of £. nearcticus Yoshimoto from Alberta was collected from gravel in water. A few specimens appear to have been collected in more open habitats (heron rookery, edge of cornfield), though it
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Review of Eustochus JESO Volume 138, 2007
is not possible to know exactly what microhabitat they were in. Females and the males of E. atripennis and E. besucheti were collected near the base of an old trunk (Viggiani 1970), from soil under oak (specimen record, below), and from moss (Bakkendorf 1965), respectively. A few were also collected in pitfall or Malaise traps. Label data on many females from Switzerland indicated they were collected from dead leaves and the Belgian specimen (from Forét de Soignes, Rouge Cloitre) described by Debauche (1948) was collected by sifting dead leaves. Most likely, species of Eustochus parasitize hosts that lay their eggs in soil, surface litter, or mosses. The number of specimens collected near water suggests an aquatic host. A given species of Eustochus likely has more than one generation per year, judging from the range of dates when specimens were collected, e.g., March to September for E. atripennis.
Key to Eustochus species
Females
l Ovipositor exerted distinctly beyond apex of gaster, considerably longer than emenmens. SAT SOAN) LOC. eb! adouedatl ..lmortemaesls) Fano: 2
— Ovipositor not exerted beyond apex of gaster, shorter than metatibia (Figs. 38, 42, | ER kee SEUPEN Ng avd SE ee ee ee BE CE ges gee LY eee Mee eer eee | 4 nn 6
2(1) Fl, 3.5 times as long as wide, only slightly shorter than fl, (Figs. 16, 19) ............. 3
— Fl, at most 2.2 times as long as wide, distinctly shorter than fl, (Figs. 17, 20, 21)
3(2) Clava 2-segmented; fi, 2.1 times as long as wide (Fig. 16); forewing with longer venation (distance between second and third macrochaeta at least 1.8 times distance between first and second macrochaeta) .................00000eeeeeeeees E. atripennis (Curtis)
_ Clava 3-segmented; fi, 2.7 times as long as wide (Fig. 19); forewing with shorter venation (distance between second and third macrochaeta about 1.4 times distance between first and second macrochaeta) ................::0000ceee E. triclavatus Xu and Lin
4(2) Fl fi, each longer than wide (Figs. 20, 21); Nearctic ............cccceseseseseeeeseeeeseeees 5
— Fl.-fl, each as wide as long, quadrate (Fig. 17); European ............c:ccceeseeeseseeeeees empresa tie Nd Oat Cool ted sates La Acadaieds E. besucheti Bakkendorf
5(4) Forewing (Fig. 31) with distinct triangular asetose area behind venation (between level of hypochaeta and first distal macrochaeta) and in front of one and a partial second row of microtrichia that extend basally to level of submarginal vein; mandible with 3 teeth [Western North America] ..................... E. yoshimotoi sp. n.
= Forewing (Fig. 30) with narrow, linear asetose area behind venation (between level of hypochaeta and first distal macrochaeta) and in front of two rows of microtrichia that extend basally to level of submarginal vein; mandible with 2 teeth [Eastern
mmmmincds OS bile to) uo hid oul Lleida dina. E. pengellyi sp. n. 6(1) FWL/FWW less than 4.0; Nearctic ..................ccssssseeseeeees E. nearcticus Yoshimoto se nF OW aVoereater than:4.0; Palacarctic wi..2 :.6...aties. a 7
7(6) Mesosoma deep, about 1.5 times as long as high and dorsum distinctly curved in lateral view (Fig. 43); forewing (Fig. 33) with about 8 microtrichia on blade behind apex of submarginal vein + base of marginal vein, arranged in 1-2 indistinct
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rows; asetose area behind these (and in front of retinaculum) relatively wide [area concealed: behind -wirigsin Fig; 33} elia...cciw. soos. Ba E. confusus sp. n. - Mesosoma shallow (Fig. 44), about 1.8 times as long as high and dorsum much flatter in lateral view; forewing (Fig. 34) with about 15 microtrichia on blade behind apex of submarginal vein + base of marginal vein, arranged in 2-3 indistinct rows; asetose area behind these (and in front of retinaculum) relatively narrow wa ddd. Sots oi GMebereh eased, 2c ae es nies ates baat E. nipponicus sp. n.
Eustochus atripennis (Curtis) (Figs. 1-15 [but see comments below], 16, 25, 26, 35, 45, 46)
Mymar atripennis Curtis, 1832: folio 411 (description in key); Haliday, 1833: 350 (list).
Eustochus atripennis Haliday, 1833: 349 (description, transfer to Eustochus); Walker, 1846: 54 (list); Foerster, 1847: 226 (description), 233 (German record from Aachen); Blanchard, 1840: 293 (diagnosis); Dalla Torre, 1898: 428 (list); Ashmead, 1904: 363 (mention in key); Schmiedeknecht, 1909: 495 (list); Kryger, 1950: 63 (description); Debauche, 1948: 201 (description); Bakkendorf, 1965: 122 (key); Viggiani, 1970: 135 (male description); Trjapitzin, 1978: 967 (duplicate of Bakkendorf key); Graham, 1982: 221 (type material); Schauff, 1984: 51 (type material lost); Viggiani, 1989: 146 (male genitalia); Ulrich, 1999: 388 (collection record).
Diagnosis. Female. Body length 978 (820-1330, air and critical point dried specimens). Head width 201-251 (n=6). Mandibles each with 3 teeth (2 teeth in specimens from Japan that may be E. atripennis, see additional material, below), the middle tooth slightly the largest, the dorsal tooth blunt. Mesosoma length/height 1.29-1.44 (n=2). Ovipositor length 746-1007 (n=9), 1.55-1.78 times length of hind tibia; distinctly exserted beyond apex of metasoma, the exserted part slightly less than hind tibial length.
Antenna (Fig.16). Segment L (W) (n=10): scape 136-169 (32-41), pedicel 66-81 (30-36), fl, 65-95 (15-20), fl, 64-83 (17-20), fl, 56-77 (17-20), fi, 49-60 (22-32), fl, 42-51 (29-40), fl 38-45 (34-45), entire clava 130- 157 (55-79). Ratios of L/W: scape 3. 74- 4.79, pedicel 1.84-2.48, fl, 3.82-5.48, fl, 3.53-4.45, fl, 3.29-4.1, fl, 1.69-2.26, fi, 1.16-1.54, fi, 0.86-1.04, entire bias 1.84-2.46.
Wings (n=10). FWL 948-1282, FWW 323-454, FWL/FWW ratio 2.88-3.06. Distance between first and second distal macrochaetae 1.85-2.15 (n=6) times distance between proximal and first distal macrochaeta. HWL 841-1044, HWW 33-43.
Male. Body length 1075 (critical point dried specimen). Head brown, distinctly darker than yellowish brown body. Head (Fig. 45) large, with height 208, length on midline 149, width 307, and 1.55 width of mesosoma. Face in lateral view forming a distinct rounded protrusion just below eye level, with toruli facing obliquely upward and lower face and mouthparts strongly sunken in, in lateral view not visible due to protruding gena; face in anterior view with a curved row of 6 setae above mouth margin. Vertex small, without ocelli (cf. Viggiani 1970), and widely separated from back of head by occiput. Eye small.
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Malar space as long as eye and malar sulcus absent. Gena 3 times width of eye. Mouth about two-thirds width of head, with mandibles huge and capable of overlapping, each with three teeth, the dorsal one blunt and set back, not in line with the two ventral teeth. Labrum with one median seta. Mesosoma narrow and reduced. Pronotum visible in dorsal view, divided medially. Propleura visible in dorsal view as protruding ‘shoulders’ lateral to pronotum. Mesoscutum small, triangular, margined laterally by posterior of each pronotal lobe. Notauli absent. Scutellum apparently not divided into anterior and posterior portions and without placoid sensilla. Dorsellum narrow. Propodeum with small spiracle separated about 3 times its diameter from dorsellum; propodeal seta midway between anterior and posterior margins of propodeum, slightly more medial in position than spiracle. Pronotal lobe length (maximum) 46, mesoscutum length45, scutellum length 47, dorsellum length 14, propodeum length 122. Metasoma with petiole length 122. Gastral tergum1 overhanging entire petiole, approximate relative lengths of gt,-gt, (along dorsal margin, critical point dried specimens) 11, 15, 12, 9, 9, 9, 4. Spiracle apparently present. Cerci with two dorsal setae longer than ventral two and curved. Gaster length (critical point dried specimen) 614. Genitalia as in Fig. 46.
Forewing present (cf. Viggiani 1970) but minute and almost haltere-like. Hindwing absent.
Antenna (Fig. 25) with 9 flagellomeres, the apical two widely joined, clava-like. Relative proportions of segments L (W) [number of longitudinal sensilla, when present; if different between left and right antenna, both numbers are given]: scape 130 (35), pedicel 72 (38), fi, 42 (22), fl, 41 (22), fi, 39 (27/24) [1/0], fl, 41 (26/34) [1/2], fl, 36 (36) [2], fi, 36 (37) [2/3], fi, 43 (38) [2], fl, 42 (39) [3], fl, 44 (35) [3]. The widths of fl, and fi, differ between the left and right antennae due to the different numbers of longitudinal sensilla.
Discussion. This species is distinguished from the other species with long ovipositors by proportions of the funicle segments, microtrichial pattern behind the forewing venation, and relatively longer marginal vein. No other described species has such a long venation (distance between first and second distal macrochaeta at least 1.8 times distance between proximal and first distal macrochaeta).
The type material of E. atripennis is lost (Graham 1982). Haliday sent specimens to Spinola in Turin (MRSN) and Graham examined the one specimen of E. atripennis there. He suggested that it could be designated as neotype, if no undoubted Walker specimen were found. This specimen was examined by JH. It is still in good condition, exactly as Graham (1982) described it (complete, except right pair of wings missing) but is not designated as a neotype here because E. atripennis is not a problematic taxon and an objective definition of it is not necessary (ICNZ, 1999, Article 75). The specimen (MRSN) is labelled: 1. “Museo Zoologia/Torino - Italia”. 2. “Eustochus/ atripennis Haliday/ Lectotype °/ M de V. Graham, 1972”. This lectotype designation is incorrect, because the specimen was not from the original material seen by Curtis. If desired, the specimen certainly serves as an example of what Haliday meant when redescribing the species but there are many, more recently collected, specimens in several museums that can serve as well.
Material examined. Forty-seven females and | male on cards or points, 10 female and one male on slides (58 additional Swiss and one French specimen were examined many years
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prior to the present study; they are listed here). CZECH REPUBLIC, Bohemia: Praha- Stomovka, 18-19August 1999, L. Masner, riparian, yellow pan trap (2, CNC); Revnice environs, 20-21 August 1999, L. Masner, creek (2, CNC). ENGLAND, Avon: Bristol, Hallen Wood, 22 July 1925 (2, USNM); Berkeshire, Wytham, em., 8 September 1949, from soil under oak, G. C. Varley (22, BMNH); Cornwall, Scilly Is., Tresco, 21 September 1975, J. S. Noyes (2, BMNH); Devon, Birchette Wood, 13-21 August 1980 (2, BMNH); Dorset, Bournemouth, 13 July, 17 August, and September 1981, 28 June 1983, S.C. S. Brown (42, BMNH); Greater London, Richmond, 25 September 1907, C. Waterhouse (2, BMNH); Hampshire, New Forest, 24 June 1954, J. Murgetroyd (9, BMNH); Romsey, Awbridge, August, September 1981, June 1982. C. Vardy (92, BMNH). FRANCE, Haute- Savoie: Vongy, 11 June 1964, C. Besuchet, dead leaves (9, MHNG). GERMANY, North Rhine-Westphalia: Bornheim-Brenig, no date, M. Boness, on red currant (2, CNC). HUNGARY, Somogy: Mernye, 17 August-4 September 1985, N. D. Springate (32, CNC). SPAIN, Navarra: Artikutza, 29 May 1995, 600 m, 18 May 1997, 25 August—22 September, 6-20 October, and 20 October—17 November 1996, 590-610 m, L. Martinez de Murguia, Malaise trap (62, CNC, MZNA); Iratibizkar, 26 July 2000, E. Baquero, pitfall trap (29, CNC, MZNA). SLOVENIA: Bled, 5-12 August 1978, L. Huggert, luxuriant spruce forest, pan trap (2, CNC); Rateée, 31 July—7 August 1978, L. Huggert, edge of marshy area, pan trap, (2, CNC). SWITZERLAND, Geneva: Chancy, 8 May 1964, C. Besuchet, dead leaves (54, 232, CNC, MHNG); L’Allondon, 2 and 7 May 1959, C. Besuchet, sifting mosses (15°, MHNG); Ticino: Rancate, 8 August 1963, C. Besuchet, dead leaves (289, CNC, MHNG); Valais, Euseigne, 1000 m, 10 July 1970, C. Besuchet, det. Viggiani, 1976 (52, MHNG); Vouvry, 27 March 1967, at base of old stump, det Viggiani, 1970 (¢, 29, MHNG); Vaud, Cossonay, 4 August 1953, C. Besuchet, in old stump (2, MHNG).
Additional material (possibly E. atripennis). JAPAN, Honshu: Ibaraki, Mt. Tsukuba, 800 m, 18 September—2 October, 2-20 October 1989, M. J. Sharkey, pan trap (72, CNC). SOUTH KOREA, Kangwon-Chucheon: Nam-myeon, Hudong-li,°31 July—16 August 2003, Malaise trap in semi-shade, forest edge (2, CNC). The South Korean specimen is as large as the European (especially British) specimens, but fl, is longer so it is only tentatively identified as E. atripennis. The Japanese specimens are smaller than E. atripennis from Europe and because we are not sure if they are conspecific with it, we list them separately here. The scanning electron micrographs were taken from the Mt. Tsukuba specimens and may be E. atripennis. They have a long marginal vein but bidentate, instead of tridentate, mandibles.
Eustochus besucheti Bakkendorf (Figs. 17, 27, 36, 37) Eustochus besucheti Bakkendorf, 1965: 117 (description); Xu and Lin, 2003: 65 (list). Diagnosis. Female. Body length 712-772 (n=3, critical point dried specimens). Head width 192 (n=1). Mandibles each with 3 teeth. Mesosoma length/height about 1.35. Ovipositor length 585, distinctly exserted beyond apex of metasoma and 1.85 times length
of hind tibia.
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Antenna (Fig. 17). Segment L (W) (n=1): scape 135 (32), pedicel 64 (31), fl, 45 (18), fl, 45 (18), fl, 53 (20), fl, 33 (30), fi, 33 (36), fl, 33 (39), entire clava 123 (58). Ratios of L/W: scape 4.21, pedicel 2.05, fi, 2.48, fl, 2.56, fi, 2.68, fl, 1.1, fi, 0.92, fl, 0.84, entire clava 2.15.
Wings (Fig. 27). FWL 810, FWW 229, FWL/FWW ratio 3.54. Distance between first and second distal macrochaetae 1.33 times distance between proximal and first distal macrochaeta. HWL 689, HWW 34.
Male. Eustochus besucheti is the only other species besides E. atripennis whose male is known, based on a single specimen from Laquintal, the type locality. Apart from apparently narrower fi, and fi, in E. besucheti, the males of the two species appear to be identical.
Discussion. This species is distinguished from FE. atripennis, the only other European species with a long ovipositor, by its smaller size, shorter funicle segments, especially fi, (Fig. 17), shorter forewing venation, and less sclerotized posterior margin of the forewing just beyond the retinaculum.
Material examined. Three females on cards or point, one on slide. The holotype 2 and allotype ¢ (MHNG) were not examined. CZECH REPUBLIC, Bohemia: Studnice near Jablonec nad Nison, 860 m, 14 July 1964, V. Martinek (Q, CNC). SWITZERLAND, Turgau: Unterwasser, 1440 m, 4 August 1984, L. Masner, screen sweeping (2, CNC); Valais, Laquintal, 1400-1500 m, 1 July 1962, C. Besuchet, mosses (29, CNC). The Laquintal specimens examined were preserved in alcohol from the type locality, but not designated as paratypes by Bakkendorf (1965). They were critical point dried and point mounted by the senior author and distributed between MHNG and CNC, by permission of C. Besuchet.
Eustochus confusus Huber and Baquero, sp. nov. (Figs. 23, 33, 43, 47-50)
Diagnosis. Female. Body length 1100 (n=1, paratype on slide). Head width 230 (n=1). Mandibles tridentate, each with a small dorsal tooth and 2 larger teeth, the middle one thicker and longer than the ventral one. Mesosoma length/height about 1.5 (Fig. 43). Ovipositor length 338-356 (n=2), not exserted beyond apex of metasoma and less than (0.80-0.82 times) length of hind tibia.
Antenna (Figs. 23, 47, 48). Segment L (W) (n=2): scape 161-174 (38-40), pedicel 76-78 (32-33), fl, 64-66 (20), fl, 58-66 (21-24), fl, 51-52 (20-21), fl, 40-44 (28-31), fl, 38-41 (29-35), fl, 37-39 (29-41), entire clava 125-132 (46-48). Ratios of L/W: scape 4.21-4.39, pedicel 2.35-2.36, fi, 3.24-3.29, fl, 2.38-3.10, fl, 2.54-2.60, fl, 1.29-1.57, fi, 1.08-1.38, fi, 0.94-1.33, entire clava 2.58-2.87.
Wings (Figs. 33, 49). FWL 1083-1151, FWW 262, FWL/FWW ratio (n=1) 4.39. Distance between first and second distal macrochaetae 1.03-1.04 times distance between proximal and first distal macrochaeta (Fig. 50). HWL 857-916, HWW 33-41.
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Discussion. This species is distinguished from the only other species with a non-exserted ovipositor in the Palaearctic region, E. nipponicus, by the deeper mesosoma that is dorsally more rounded in lateral view (Fig. 43) compared to a shallower mesosoma that is dorsally flatter in lateral view (Fig. 44) and the wider asetose area between the retinaculum and rows of microtrichia behind the venation (Fig. 33). In Europe, E. confiusus is the only species with an ovipositor shorter than the hind tibial length. The other two, E. atripennis and E. besucheti have long, exserted ovipositors that are longer than the hind tibia. It differs from E. nearcticus by its narrower forewing (length/width ratio about 4.4).
Material examined. Two females on slides. HOLOTYPE ¥Y (MZNA) on slide labelled: 1. “MZNAXAO98a /Irati (Navarra, Spain), /18.1X.1982/ UTM:30TXN5460,/390 m, UNZYEC leg./ MUSEO DE ZOOLOGIA/UNIV. DE NAVARRA”. 2. “Eustochus confusus Huber and Baquero. Holotype Y”. The holotype is complete and was originally uncleared and mounted laterally in Hoyer’s medium under a single cover slip (Fig. 23, 33, 43). After being photographed it was cleared and remounted in Canada balsam. PARATYPE. Same data as holotype (9, MZNA).
Species name. The species name, confusus, is Latin for confusing. It refers to the initial confusion we had as to whether E. confusus was the same as E. nearcticus or not. If it had been, it would be the first Eutochus species occurring in both Europe and North America. While this is possible, because soil brought to North America as ballast in ships may have contained parasitized hosts, it is unlikely, given the occurrence of both species in localities that are not particularly near ports and that consist of native vegetation rather than human altered habitats.
Eustochus nearcticus Yoshimoto (Figs. 18, 28, 38, possibly also 22, 32, 42) Eustochus nearcticus Yoshimoto, 1990: 96 (description).
Diagnosis. Female. Body length 690-717 (n=2, critical point dried specimens). Head width 194 (n=1). Mandibles each with 3 teeth, the dorsal one smaller than the ventral two. Mesosoma length/height 1.55 (n=1) (Fig. 38). Ovipositor length 256-268 (n=2), 0.69-0.76 times length of hind tibia; not exserted beyond apex of metasoma.
Antenna (Fig. 18). Segment L (W) (n=2): scape 125-141 (30-32), pedicel 66-69 (29-30), fl, 55-66 (15), fl, 52 (19-20), fl, 44-46(17-18), fl, 41-47 (22-24), fl, 38-40 (29), fl, 36-37 (32-35), entire clava 124-129 (39-50). Ratios of L/W: scape 4.17-4.44, pedicel 2.30-2.33, fl, 3.66-4.42, fl, 2.62-2.76, fl, 2.47-2.68, fl, 1.73-2.09, fl, 1.32-1.41, fl, 1.05-1.12, entire clava 2.48-3.32.
Wings (Fig. 28). FWL 978-1048, FWW 278-283, FWL/FWW ratio 3.46-3.85. Distance between first and second distal macrochaetae 1|.03-1.09 times distance between proximal and first distal macrochaeta. HWL 778-863, HWW 30-31.
Discussion. Eustochus nearcticus is the only described species with a short ovipositor (Fig. 38) in the Nearctic region. It is distinguished from the two Palaearctic species with short
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ovipositors by proportions of the forewing and mesosoma. Eustochus nearcticus differs from E£. nipponicus by its deeper mesosoma (length/height ratio about 1.5 compared to about 1.8), with a more rounded dorsum in lateral view. It differs from E. confusus by its wider forewing (length/width ratio at most 4.08).
Several specimens from western North America (AB, BC, MT) and southeastern USA (NC) may belong to FE. nearcticus. We exclude them because we are not sure that they really are conspecific. They are listed separately below. Minor differences in their antennal proportions (funicle segments relatively short, somewhat as in E. confusus, in those specimens slide mounted and measured) size of bare area in front of fore-wing retinaculum, and height of mesosoma (e.g., Figs. 22, 32, and 42, all from a specimen from NC), may reflect intraspecific variation within.£. nearcticus but may also indicate that they are a separate species. A conservative approach is taken here and they are excluded from E. nearcticus until more material, especially from intervening areas, becomes available for detailed study. The antenna (Fig. 22) of the NC specimen resembles that figured by Yoshimoto (1990, fig 36) in that fl, is relatively short, compared to Fig. 18. Both of the latter figures are specimens from the type localities.
The specimens from British Columbia appear to have a flatter mesosoma, more resembling FE. nipponicus, than the point-mounted paratypes of FE. nearcticus. It is difficult to be sure how these specimens are distinguished from E. nipponicus. However, the western Nearctic specimens all appear to have fl, slightly longer than fl,, whereas E. nipponicus has fi, shorter than fl,. If they are indeed the same species as E. nipponicus then wider species limits would then have to be accepted. Similarly, FE. nearcticus specimens from eastern Canada may be the same as E. confusus, but, if so, wider species limits would again have to be accepted. Finally, the western specimens may represent a different species from either E. nipponicus, E. confusus, or E. nearcticus, and possibly could be defined geographically as well as morphologically. But since several specimens, possibly of E. nearcticus, are from Ontario it is difficult to define the two populations geographically. It is also possible that E. nipponicus and E. confusus are the extremes of only one species occurring across the Palaerctic region, but so far no Eustochus resembling these two species has been collected between Spain and Japan.
We prefer to treat the specimens with short-ovipositors as three species, with E£. nearcticus occurring in northeastern North America but not in the Palaearctic region, and EF. nipponicus distinct from E. confusus in the Palaeactic region. Much more material of these species is required, especially from intervening areas, to verify their status relative to one another and determine how many species really are involved.
The craterlike pits on the propodeum, visible on the scanning electron micrograph in figure 148 of Yoshimoto (1990), are an artifact, presumably due to remnants of a liquid deposit on the specimen. Cleared, slide mounts prepared of two specimens from the type locality do not show these pits; the sculpture is uniformly reticulate.
Material examined. Three females on points, two on slides. The holotype (CNC) is complete and in good condition on a point. Three paratypes from Nova Scotia, as listed in the original description, were also seen. The fourth paratype, from British Columbia, is a different species, assigned here to E. yoshimotoi, sp. n. (see below). The paratype from MacKenzies Mt. was slide mounted for detailed study. An additional specimen from
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MacKenzies ML, listed below, was also slide mounted; it was not included in the type series by Yoshimoto (1990), for unknown reasons,
CANADA, Nova Scotia: Cape Breton Highlands Nat. Park, MacKenzies Mt., 9 August 1983, J. E. H. & R. J. Martin (9, CNC),
Additional material, Thirteen other specimens that may be /, nearcticus were examined, 3 of them (MT and NC) are on slides. CANADA, Alberta: Waterton Lakes National Park, 2 August 1985, Cameron Creek on Akamina Parkway near mouth of Rowe Creek, gravel in riffle, 1. M. Smith (9, CNC). British Columbia: Kootenay National Park, Daer Pitts, 16- 30 July 2000, G. Gareau, MT, aspen (25°, CNC). Ontario: Brucedale Conservation Area
near Port Elgin, 19 April-16 June 1988, C. Dondale and J. Redner, pit fall trap at edge of
swamp (Y, CNC). UNIFPED STATES: Montana, Flathead Co., Glacier National Park, N. Fork Flathead area, S. Big Prairie, 3560°, T35N R21 W, sect. 16, 10-17 August 1993, M. A, Ivie, old growth light burn (24°, CNC). North Carolina: Jackson Co., Whiteside Mt., near Highlands, 1600 m, April-20 July 1987, CNC Hym. Team, oak forest (29, CNC); Swain Co., Andrews Bald, pitfall 51, N 35° 20°32” W 83° 39°29’, 10-24 May, 6-22 June, and 10- 25 September 2001, Parker, Stocks, Petersen (S°, CNC).
Eustochus nipponicus Huber and Baquero, sp. nov. (Figs. 24, 34, 44)
Diagnosis. Female. Body length 947-1100 (n=3, critical point dried specimens), Head width 188-198. Mandibles each apparently tridentate, with 2 large ventral teeth and a small, dorsal tooth. Mesosoma length/height 1.82 (Fig. 44). Ovipositor length 281 (holotype), not exserted beyond apex of metasoma and less than (0.91 times) length of hind tibia.
Antenna (Fig. 24). Segment L (W) (holotype): scape 138 (35), pedicel 69 (32), fl, 51 (19), fl, 51 (20), fi, 41 (20) fl, 44 (24), fl, 39 (29), fl, 37 (32), entire clava 110 (54). Ratios of L/W: scape 3.89, pedicel 2.19, fl, 2.71, fl, 2.62, fl, 2.01, fl, 1.10, fl, 1.36, fl, 1.15, entire clava 2.03.
Wings (Fig. 34). FWL 974, FWW 232, FWL/FWW ratio 4.09. Distance between first and second distal macrochaetae 1.04 times distance between proximal and first distal macrochaeta. HWL 823, HWW 34.
Discussion. This species is distinguished from /. confusus, the only other Palaearcic species with a short ovipositor, by the shallower mesosoma with a flatter dorsum in lateral view, and the narrower asetose area between the retinaculum and rows of microtrichia behind the venation (Fig. 34).
Material examined, Four females, one ona slide. HOLOTYPE Y (CNC) on slide labelled: |. “Japan: Honshu/, Iwate, lwaizumi/ Hitsutori, 770m/11-17.vili.1991/ A. Smetana [J47]”. 2. “Eustochus/ nipponicus/ Huber &/ Baquero/ Holotype Y”. The holotype is cleared and mounted laterally (Fig. 44) under a 6 mm cover slip, with the wings and one antenna under two additional cover slips. PARATYPES. Same locality data as holotype (29, CNC); Iwate, Kawai, Yoshibezawa, N 39° 37’ E 141° 31°, 500 m, 25 August 1996, L. Masner, screen sweeping (Y, CNC).
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Species name. The species is named from the Japanese name for the country of origin, Nippon.
Eustochus pengellyi Huber and Baquero, sp. nov. (Figs. 20, 30, 40)
Diagnosis. Female. Body length 666-896 [up to 973 when gaster somewhat inflated] (n=8, critical point dried specimens). Head width 184 (holotype). Mandibles each with 2 teeth. Mesosoma length/height 1.33-1.44 (n=3, critical point dried specimens). Ovipositor length 439, 1.40 times length of hind tibia and distinctly exserted beyond apex of metasoma.
Antenna. Segment L (W) (holotype): scape 121 (30), pedicel 62 (27), fl, 50 (14), fl, 46 (17), fl, 39 (20), fl, 38 (23), fl, 36 (27), fl, 35 (29), entire clava 105 (54). Ratios of L/W: scape 4.08, pedicel 2.29, fl, 3.63, fl, 2.62, fl, 1.97, fl, 1.66, fl, 1.33, fl, 1.22, entire clava 1.93.
Wings. FWL 799, FWW 228, FWL/FWW ratio 3.51. Distance between first and second distal macrochaetae 1.15 (1.30 on other wing) times distance between proximal and first distal macrochaeta. HWL 668, HWW 26.
Discussion. This species is distinguished from FE. yoshimotoi, the only other Nearctic species with a long, exserted ovipositor, by a narrower asetose area above the retinaculum that is separated from the venation by only one complete line of microtrichia (wider asetose area and at least a partial second line of microtrichia in E. pengellyi). It differs from E. atripennis by its shorter venation, from E. besucheti by antennal proportions, and from E. triclavatus by the number of claval segments. The specimens from Japan that we treat as possibly E. atripennis on the basis of a long venation, also have bidentate mandibles.
Material examined. Thirteen females, two on slides. HOLOTYPE 9 (CNC) on slide labelled: 1. “CANADA: ON/, Flint Hill near/ Kemptville/, 19-20.vii.1983/, L. Dumouchel”. 2. “Eustochus/ pengellyi/ Huber and Baquero °/ Holotype”. The holotype is cleared and mounted dorsally in Canada balsam under one 6 mm cover slip and the wings, and head, antennae and prothorax are under two other coverslips. PARATYPES. CANADA, Ontario: Frontenac Co., 5 km W. Chaffey’s Locks, Skycroft Campground, 9-14 July 1987, B. Hubley, MT (22, CNC); Thetford, 10-13 October 1982, A. Tomlin, FIT (2, CNC); Haliburton Forest and Wildlife Reserve, N 45° 15’ W 78° 35’, 12 July 2001, C. Vance, maple forest, MT (2, CNC); 7 km SE Westport, 134 m, N 44° 37°727’ W 76° 217545’, 1-31 August 2005, S. Peck, maple sugar bush, FIT (9, CNC). Quebéc, Lac Jean-Venne, N 45° 41’ W 76° 03’, 15-20 June, Osmunda marsh, YPT and 14-21 August 1995, L. Masner & J. Denis, Osmunda marsh creek, YPT (22°, CNC). UNITED STATES, Indiana: Laporte Co., Indiana Dunes National Lakeshore, Heron Rookery, 29 July 1997, R. Grundel, MT (22, CNC, USNM). Maryland: Prince George's Co., Beltsville-Agricultural Research Centre, 1-9 July1980, K. Thorpe, Malaise trap on corn field edge (2, USNM). Tennessee: Blount Co., Cades Cove, N 35° 2535’ W 83° 17”50’, 29 July1997, R. Grundel, MT (2°, CNC).
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Species name. The species is named in honour of Dr. David Pengelly, whose enthusiasm for teaching entomology encouraged several students to pursue insect taxonomy as a career.
Eustochus triclavatus Xu and Lin (Figs. 19, 29, 39) Eustochus triclavatus Xu and Lin, 2003: 66 (description).
Diagnosis. Female. Body length about 720 (n=1, ?paratype on slide). Head width 193. Mandibles each with 3? teeth (not clearly visible). Mesosoma length/height 1.35. Ovipositor length 415, distinctly exserted beyond apex of metasoma and 1.16 times length of hind tibia.
Antenna (Fig. 19). Segment L (W) (?paratype): scape—[not measurable] (ca. 23), pedicel 63 (24), fl, 47 (15), fl, 52 (15), fl, 47 (14), fi, 52 (20), fl, 45 (23), fi, 41 (23), entire clava 137 (40). tion of L/W: pedicel 2. 58, fi, 3. 14, fi, 3.46, fl, 3.25, fi, 2:57, fi, 1.96, fi, 1.77, entire clava ca. 3.42.
Wings (Fig. 29). FWL 857, FWW 264, FWL/FWW ratio 3.25. Distance between first and second distal macrochaetae 1.41 times distance between proximal and first distal macrochaeta. HWL 731, HWW 31.
Discussion. This species is distinguished from all others described so far by the three- segmented clava (two segmented in other species). It belongs to the group of species with distinctly exserted ovipositors.
Material examined. One female on slide. CHINA, Shanxi: Fengxian, 2°, 4 September 1999, N.Q. Lin, CNC. The locality (Baoji) given in the original description differs from the specimen label, quoted here; it is not certain whether this specimen is a paratype. The holotype 2 (FAFU) was not examined.
The paratype is uncleared, poorly oriented, and the clava of each antenna is slightly collapsed. It appears to have the division between claval segments 2 and 3 less distinct and perhaps incomplete, at least on one antenna.
Eustochus yoshimotoi Huber and Baquero, sp. nov. (Figs. 21, 31, 41)
Diagnosis. Eustochus yoshimotoi is one of two Nearctic species with a distinctly exserted Ovipositor. It is distinguished from the other, E. pengellyi, by the wider, more distinct asetose area behind the marginal vein (narrower in pengell/yi) and the tridentate mandible (bidentate in E. pengellyi).
Description. Female. Body length 845 (n=3, critical point dried specimens). Head width 211 (n=2). Mandibles each with 3 teeth. Mesosoma length/height 1.29-1.48. Ovipositor (Fig. 41) distinctly exserted beyond apex of metasoma, 485-525 (n=3) long and 1.24-1.32 times length of hind tibia.
Antenna (Fig. 21). Segment L (W) (n=2): scape 135-144 (32-33), pedicel 66-71 (32-33), fl, 55-60 (17), fl, 48-56 (17), fl, 44-54 (19-20), fi, 39-44 (27-29), fi, 39-41 (34),
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fi, 39-40 (38), entire clava 121-138 (57-59). Ratios of L/W: scape 4.06-4.28, pedicel 2.07- 2.10, fi, 3.27-3.51, fi, 2.80-3.19, fl, 2.25-2.87, fl, 1.43-1.55, fi, 1.14-1.20, fi, 1.05-1.06, entire clava 2.03-2.59. .
Wings (Fig. 31). FWL 950-994, FWW 281-298, FWL/FWW ratio 3.33-3.44. Distance between first and second distal macrochaetae 0.90-1.02 times distance between proximal and first distal macrochaeta. HWL 739-809, HWW 38-41.
Material examined. Seven females, four of them on slides. HOLOTYPE & (CNC) on slide labelled: 1. “USA, WA,/ Pierce Co./ Ashford/ 1-14.viti.1985/ L. Masner, Malaise trap”. 2.“Eustochus/ yoshimotoi °/ Huber and/ Baquero/ Holotype”. The holotype is cleared and mounted laterally under one 6 mm cover slip.in Canada balsam and the wings are under another coverslip. PARATYPES. Same data as holotype (42, CNC, USNM); CANADA, British Columbia, Vancouver Is., Mesache Lake, July 1984, MT, Sharkey—Johnson (°, CNC). UNITED STATES, California, El Dorado Co., Blodgett Forest, 27 August 1975, F. Andrews, M. Wasbauer, Pinus ponderosa log (2, UCRC). The paratype from British Columbia was designated originally as a paratype of E. nearcticus by Yoshimoto (1990) and bears the paratype label “Eustochus nearcticus”, as well as ours “Eustochus yoshimotoi’.
Species name. The species is named in honour of Carl Yoshimoto, the senior author's predecessor with the Canadian Forest Service, who was a taxonomist at the CNC and described the first North American species of Eustochus.
Acknowledgements
We thank K. Bolte for preparing the digital images and compiling the plates and the curators of the institutions listed above for loaning or donating material to us for study. B. Landry kindly checked the number of specimens of E. atripennis and E. besucheti in MHNG and provided their collection information. Through the kindness of G. Pagliano (MRSN), the senior author was able to borrow the Haliday specimen of E£. atripennis in the Spinola collection, and an additional specimen of one species was found and brought to my attention by S. Triapitsyn (UCRC). The manuscript was critically reviewed by G. Gibson and A. Bennett (CNC).
References
Annecke, D. P. and R. L. Doutt. 1961. The genera of the Mymaridae. Hymenoptera: Chalcidoidea. Entomology Memoirs. Department of Agricultural Technical Services, Republic of South Africa 5: 1—71.
Ashmead, W. H. 1904. Classification of the chalcid flies of the superfamily Chalcidoidea, with descriptions of new species in the Carnegie Museum, collected in South America by Herbert H. H. Smith. Memoirs of the Carnegie Museum 1(4): 225— 555.
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Bakkendorf, O. 1965. Description of a new subterranean species including a male and female of Eustochus Hal. Mitteilungen der Schweizerischen Entomologischen Gesellschaft 37: 117—122.
Blanchard, E. 1840. Histoire naturelle des insectes Orthopteres, Névroptéres, Hémiptéres, Hyménopteres, Lépidopteres et Dipteres. Paris: Duménil. 672 pp.
Curtis, J. 1832. British Entomology 9: folio 411. Plates 386—433 (with text).
Dalla Torre, C. G. de. 1898. Subfam. Mymarinae. pp. 422-431. Catalogus hymenopterorum hucusque descriptorum systematicus et synonymicus. Vol.5: Chalcididae et Proctotrupidae. Lipsiae [Leipzig]: Guilelmi Engelmann. 598 pp.
Debauche, H. R. 1948. Etude sur les Mymarommidae et les Mymaridae de la Belgique (Hymenoptera Chalcidoidea). Mémoires du Musée Royal d’Histoire Naturelle de Belgique 108: 1—248 + 24 plates.
Foerster, A. 1847. Ueber die Familie der Mymariden. Linnaea Entomologica 2: 195-233.
Foerster, A. 1856. Hymenopterologische Studien. II]. Heft. Chalcidiae und Proctotrupii. Aachen: Ernst ter Meer. 152 pp.
Gahan, A. B. and Fagan, M. M.1923. The type species of the genera of Chalcidoidea or chalcid-flies. Bulletin of the United States National Museum 124: 1-173.
Gibson, G. A. P. 1997. Morphology and terminology. pp. 16-44 /n Gibson, G. A. P., J. T. Huber, and J. B. Woolley (eds.), Annotated keys to the genera of Nearctic Chalcidoidea. Ottawa: NRC Research Press. 794 pp.
Graham, M. W. R. de V. 1982. The Haliday collection of Mymaridae (Insecta, Hymenoptera, Chalcidoidea) with taxonomic notes on some material in other collections. Proceedings of the Royal Irish Academy, B 82: 189-243.
Haliday, A. H. 1833. An essay on the classification of the parasitic Hymenoptera of Britain, which correspond with the Ichneumones minuti of Linnaeus. Entomological Magazine |: 259-276, 333-350.
Huber, J. T. 1987. Review of Schizophragma Ogloblin and the non-Australian species of Stethynium Enock (Hymenoptera: Mymaridae). The Canadian Entomologist 119: 823-855.
ICZN. 1999. International Code of Zoological Nomenclature, Fourth Edition. London: International Trust for Zoological Nomenclature c/o The Natural History Museum. 306 pp.
Kryger, J. P. 1950. The European Mymaridae comprising the genera known up to c. 1930. Entomologiske Meddelelser 26: 1—97.
Schauff, M. E. 1984. The Holarctic genera of Mymaridae (Hymenoptera: Chalcidoidea). Memoirs of the Entomological Society of Washington 12. 67 pp.
Schmiedeknecht, O. 1909. Hymenoptera fam. Chalcididae. Genera Insectorum 97: 1—550 + 8 pls.
Schmiedeknecht, O. 1930. Die Hymenopteren Nord- und Mitteleuropas. Jena: Gustav Fischer. 1062 pp.
Trjapitzin, V. A. 1978. Hymenoptera II. Chalcidoidea 18. Mymaridae. Opred Nasek. Evrop. Chasti SSSR. 516-538. [In Russian.] [English translation: Family Mymaridae (Mymarids). pp. 942—982 Jn Keys to the Insects of the European Part of the USSR. III. Part 2., Medvedev, G. S. (ed.), 1987. Oxonian Press, New Dehli. 1341 pp.]
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Ulrich, W. 1999. Phenology and life cycles of the parasitic Hymenoptera of a dry meadow on limestone. Polskie Pismo Entomologiczne 68: 383-405.
Viggiani, G. 1970. Description of the male of Eustochus atripennis Hal., 1833, and new terricolous spcies of Cleruchus Enock, with remarks on Anagrella Bkdf. (Hym., Mymaridae). Mitteilungen der Schweizerischen Entomologischen Gesellschaft 43: 135-142.
Viggiani, G. 1989 (1988). A preliminary classification of the Mymaridae (Hymenoptera: Chalcidoidea) based on the external male genitalic characters. Bollettino del Laboratorio di Entomologia Agraria “Filippo Silvestri” 45: 141-148.
Walker, F. 1846. Descriptions of the Mymaridae. Annals and Magazine of Natural History 18: 49-54.
Westwood, J. O. 1839. Synopsis of the genera of British insects: 78—79. [Issued with An Introduction to the Modern Classification of Insects. London: Longman, Orme, Brown, Green, and Longmans. |
Xu, M. and Lin, N.-Q. 2003. A new species of the genus Eustochus Haliday (Hymenoptera: Chalcidoidea: Mymaridae) in China. Entomologia Sinica 10: 65-68.
Yoshimoto, C. M. 1990. A review of the genera of New World Mymaridae (Hymenoptera: Chalcidoidea). Flora & Fauna Handbook no. 7. (Gainesville: Sandhill Crane Press). 166 pp.
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New genus and species of Perilampidae JESO Volume 138, 2007
A NEW GENUS AND SPECIES OF PERILAMPIDAE (HYMENOPTERA: CHALCIDOIDEA) WITH UNCERTAIN PLACEMENT IN THE FAMILY
J. M. HERATY! AND D. C. DARLING? Department of Entomology, University of California, Riverside, CA 92521, USA email: john. heraty@ucr.edu
Abstract J. ent. Soc. Ont. 138: 33-47
A new genus and species of perilampid wasp is described from Yemen and Israel. The features of the adult do not allow for accurate placement within any of the existing subfamilies of Perilampidae. The adults are similar to Chrysolampinae; however, the mandibles have a 3/2 formula and although the ventral surface of the male scape has distinct pores, these are not isolated within pits or depressions, which is characteristic of Chrysolampinae and Perilampinae. The labrum is similar to most Perilampinae, but the prepectus is associated with the mesepimeron and not the pronotum. The ovipositor is strongly expanded at the apex and scimitar-shaped, whereas the ovipositor in all other Perilampidae is needle-like. The morphological features that relate to the potential phylogenetic placement of this genus in the perilampid/ eucharitid complex are discussed.
Published November 2007
Introduction
The composition and higher level relationships of Perilampidae are uncertain. There has been debate over the inclusion of Akapalinae, Chrysolampinae, Echthrodapinae, and Philomidinae together with the more easily characterized Perilampinae within a single family (Ferriére and Kerrich 1958; Riek 1966; Graham 1969; Bouéek 1972, 1983, 1988; Burks 1979; Bouéek and Rasplus 1991; Darling 1986, 1995; Noyes 1990; Gibson et al. 1999). Echthrodapinae are now placed in Torymidae (Grissell 1995). Together or in part, these problematic subfamilies have also been proposed as a sister group or paraphyletic grade to Eucharitidae (Darling 1988, 1992; Gibson et al. 1999). Support for a close relationship between these groups is based almost entirely on morphology and behavior of the first-instar larva or planidium (Heraty and Darling 1984; Darling 1988, 1992; Heraty et al. 2004). The planidia of the eucharitid subfamilies, Gollumiellinae, Oraseminae, and Eucharitinae, are all very similar and several features support the monophyly of the Eucharitidae (Heraty and
' Author to whom all correspondence should be addressed.
? Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario, M5S 2C6, Canada and Department of Ecology and Evolutionary Biology, University of Toronto, Toronto, Ontario, Canada M5S 1A1; chrisd@rom.on.ca
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Darling 1984; Heraty et al, 2004), The life history and immature stages of Akapalinae, which has been placed in either Perilampidae or Eucharitidae, are unknown. In an analysis of 20 morphological features of immature stages, Chrysolampinae were sister to Philomidinae + (Perilampinae + Eucharitidae), with the latter three groups united by features of the planidium that include presence of a straplike tergite, absence of larval antennae, and an eversible postlabium (Darling 1992), Similar results were obtained from a combined larval and adult morphology dataset (Gibson et al. 1999), Initial results from molecular analyses with 28S-D2 ribosomal transcript supported monophyly of Eucharitidae (two genera), but not Perilampidae + Chrysolampinae, although with only four taxa sampled, the results are not reliable (Campbell et al. 2000).
Adults of Chrysolampinae and Perilampinae share two apomorphic features: one or more pores concentrated within distinct pits or depressions on the ventroapical surface of the male scape, and the gaster high and triangular in profile with the 2™ and 3" gastral tergites subequal in size and fused medially (Darling 1986, 1997; Gibson et al. 1999). Philomidinae have been tentatively placed within Perilampidae (Noyes 1990, 2002), but other than possessing a similar compressed antennal flagellum and a similar gaster shape, there are no compelling adult characters to support this grouping. Similarly, the only adult feature of consequence grouping Perilampidae and Eucharitidae is the flap-like labrum with marginal setae (Darling 1988a); the labrum of Philomidinae is very different and more similar to Chaleididae (Darling 1988a; Gibson et al, 1999),
Chrysolampinae are easily recognized but difficult to define with synapomorphies. Apart from what defines Perilampidae as a whole, Chrysolampinae have only a single prominent synapomorphy — the mandibles each with two sharp apically positioned teeth (Bouéek 1972, 1988; Darling 1986). Darling (1986, 1988) proposed that the flap-like structure of the labrum with marginal setae (and no digits) could be another synapomorphy, although some species that are referable to Chrysolampus have since been discovered to have a digitate labral margin, as is typical of Eucharitidae and some Perilampinae (Darling, unpublished),
Perilampinae are easily recognized and have anumber of diagnostic and apomorphic features (Bouéek 1978, 1988). Synapomorphies of Perilampinae include fusion or at least a very close association between the prepectus and pronotum, and pronotum with a dorsal collar (Bouéek 1978). The labrum of Perilampinae is flap-like, digitate, and medially incised (Darling 1988). Most species also have a pair of aboral digits or sessile setae, and a pair of translucent areas that are interpreted as vestigial sockets (Darling 1988). The groundplan structure in Perilampidae is considered to be a fan-like labrum with marginal digits as in Eucharitidae, although aboral digits are possibly a groundplan feature (Darling 1988). Perilampinae all have 3/2 dentate mandibles, but this is likely plesiomorphic for Chalcidoidea (Bouéek 1978).
Philomidinae do not help to resolve the relationships of Chrysolampinae and Perilampinae. The subfamily is highly autapomorphic in morphology. Adults share with Perilampidae a compressed, densely setose antennal flagellum, and the basal tergite (Gt, ) with distinct dorsal and lateral panels, but the labrum is sclerotized, flap-like, and broadly attached ventrally to the epipharynx with scattered surface setae (as in Chalcididae), the prepectus is swollen and shoulder-like, and the pronotum is reduced and obscured in dorsal view (Darling 1988; Heraty and Darling, unpublished), Like Chrysolampinae, the mandibles
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are bidentate with sharp apical teeth. If Philomidinae are part of this same lineage, as suggested by the morphology of the first-instar larva, then the sails am mandibles are potentially plesiomorphic for Chrysolampinae.
Recently, a species representing a new genus of chalcidoid wasp was collected in Israel and Yemen. It has features shared with both Chrysolampinae and Perilampinae that suggests that it may be either a plesiomorphic perilampid or a potential sister group to one or both of these subfamilies. Our purpose here is not to provide a final and formal placement of this genus, but to establish its name and provide a detailed discussion of important features as a prologue to more in-depth morphological and molecular studies that are in progress on the higher relationships of the perilampid-eucharitid complex.
Methods and Materials
Terms follow Darling (1988; for mouthparts), Heraty (2002), Gibson (1997), and Heraty and Quicke (2003; for ovipositor). Our terms differ in one major aspect involving the apex of the antennal flagellum. In many Chalcidoidea with an apparent 13-segmented antenna, the apical segment of the clava often has a small button-like apical sensillar area beyond flagellomere 11 (f1,,; antennomere 13) that is differentiated to the same degree as between fi'° and fl'', but with no multiporous plate sensilla (Fig. 2, f1,,). The archaic family Rotoitidae has a 14-segmented antenna (Boucek and Noyes 1988), and we consider this button-like structure in these and other Chalcidoidea as homologous to the 14" flagellomere of Rotoitidae. Therefore, a 14-segmented antenna is a more general condition across Chalcidoidea. Colour versions of the figure plates are accessible from JMH or http:// hymenoptera.ucr.edu.
Jambiya vanharteni n. gen. and n. sp. (Figs. 1-23)
Etymology. Jambiya is treated as an arbitrary combination of letters; the gender is considered to be female. The name is based on the name of the small curved Yemeni dagger of the same name which is similar in form to the female ovipositor. The species is named after Tony van Harten, the collector of the specimens from Yemen.
Current placment. J/ncertae sedis within Perilampidae (see discussion below).
Diagnosis. In Jambiya, the mandibles are 3/2 toothed (Fig. 6), the epipharynx has a single pair of epipharyngeal setae (Fig. 4), the labrum is flap-like and digitate with paired aboral digits and translucent areas (Fig. 5), the petiole is membranous ventrally (Fig. 15), the ventral surface of the male scape has scattered minute pores without prominent pits or depressions (Figs. 7, 8), the mucro is long and acuminate (Fig. 20), and the ovipositor is apically expanded and dentate (Fig. 20). This genus is similar to Chrysomalla (Chrysolampinae), which also has the distinct carina demarking a pronotal collar and a ventrally membranous petiole, but differs in several key features considered as diagnostic of the subfamily. Jambiva differs from all known genera of Perilampinae by having the labrum not medially incised, ventral
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surface of the male scape without any distinct pits (although minute pores are present), pronotum and prepectus + mesepisternum loosely articulated, ovipositor with the ventral valves expanded and flattened, and in several aspects of the forewing venation (parastigmal break, linear stigmal sensilla, short postmarginal vein), Without any congeneric species for comparison, all of the generic diagnostic features are also relevant for species recognition.
Female. Length |.1-1.3 mm. Body color black with iridescent bluish reflections; tarsi pale brown; ovipositor valves light brown. Wings hyaline, forewing venation brown.
/ F
a “4
60 sm
FIGURES 1-8, Jambiya vanharteni. \-female head in frontal view; 2—female antenna; 3—male antenna, 4—epipharynx; 5—labrum; 6—mandibles; 7—male scape, ventral view; 8— closeup of inset box from Fig. 7, Abbreviations; anl-anellus; f1,,flagellomere 12; pdl— pedicel,
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FIGURES 9-15. Jambiya vanharteni. 9-female habitus; 10—female head and mesosoma, dorsal view; 11—13, male mesosoma: 11-—subdorsal view, 12-—sublateral view, 13- posterodorsal view; 14—petiole in dorsal view, closeup of Fig. 13; 15, petiole and first sternite in subventral view. Abbreviations: axg—axillular groove; clc—collar; crs—crenulate sulcus; fmd—femoral depression; fre—frenum; Gs,—gastral sternite 1; Gt,—gastral tergite 1; occ—occiput; pre—prepectus; vmpt—ventral margin of petiole.
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FIGURES 16-22. Jambiya vanharteni. 16—forewing; 17—hind wing; 18—stigmal vein; 19- forewing vein; 20—apex of gaster, lateral view; 21—male genitalia, ventral view; 22—ovariole. Abbreviations: aed—aedeagus; dgs—digitus; dv—dorsal valve; muc—mucro; par—paramere; phb—phallobase; pmv—postmarginal vein; stg—stigmal vein; unc—uncus; vv—ventral valve; 2vf-second valvifer; 3v—third valvula.
Head. Rounded in frontal view (Fig. 1); eyes bare; median ocellus anterior to lateral ocelli. Frons and lower face smooth and shining with scattered semi-erect setae; ocellar- ocular depression absent; scrobal depression finely reticulate, shallow with rounded lateral margins, and extending 2/3 distance to median ocellus; vertex lateral to ocelli transversely strigate; occiput circularly colliculate (Fig. 6), dorsal occipital margin with a sharp carina (Fig. 10, occ), carina extending to oral fossa although partially obscured ventrally by fine postgenal striae. Clypeus smooth with few scattered short setae, margins demarked dorsally and laterally by weakly impressed smooth sulci that are rounded at dorsolateral margins of clypeus (clypeus nearly semicircular), clypeal margin slightly rounded with a transverse shelf (Fig. 1); supraclypeal area indistinct. Malar sulcus demarked by a weak carina (Fig.
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transverse ridge
Dees Foe Pi een oye" i iy rw. ee ee Tiwi oS
FIGURE 23. Jambiya vanharteni, ovipositor: A—dorsolateral view of apex; B—ventrolateral view. Abbreviations: dv—dorsal valve; go-gonangulum; Gt —gastral tergites 7; lw—line of weakness between third valvula and second valvifer; trs—triangular sclerite; v—valvula; vf— valvifer; vv—ventral valve.
6); hypostomal lobes broadly separated. Mandibles opposing, 3/2 dentate with teeth long and subequal (Fig. 6), basal articulation with gena membranous medially. Oral fossa broad. Epipharynx with single pair of stout epipharyngeal setae (Fig. 4). Labrum with 9 marginal acuminate digits, 2 aboral digits and 2 medial aboral translucent areas/vestigial sockets (Fig. 5); palpi 4/3 segmented. Antenna 14-segmented; scape narrow and elongate with no indication of ventral pits or pores in slide mounted antenna; anellus transverse (Fig. 2, anl); length of flagellum less than height of eye; funicle 7-segmented, all segments broader than long and slightly increasing in width apically; clava with four distinct, partially fused segments, apical segment (f1,,) small and button-like; multiporous plate sensilla large and numerous on all flagellomeres except flagellomeres 1—2 and flagellomere 12 (Figs. 2, 3); numerous socketed setae on pedicel and flagellum.
Mesosoma. Pronotum with distinct transverse collar demarked by a transverse carina (Figs. 9-12, clc); collar swollen and projecting laterally anterior to spiracle (Fig. 13), anterior face imbricate and mostly bare, mostly smooth posterior to carina and with scattered semi- erect setae; pronotum overlapping mesoscutum (Fig. 11). Mesoscutum mostly smooth with scattered setae; notauli deeply impressed and converging but not meeting at midline (Figs.
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10, 11). Scutoscutellar sulci diagonal, meeting anteriorly at transscutal articulation; lateral axillar lobe rounded and smooth (Figs. 10, 11); scutellar disc smooth medially and weakly imbricate laterally, with scattered setae; axillular groove (axg) present but broad and only slightly raised above surface (Fig. 11). Frenal line present and complete dorsally; frenum (fre) broadly rounded, smooth medially and weakly striate laterally, extended only slightly over metanotum (Figs. 11-13). Metanotum with broad rounded flange laterally, overlapping propodeum but not overlapping propodeal spiracle (Fig. 13). Propodeum reticulate medially with a smooth nucha; spiracle separated from anterior margin by its diameter, margin of spiracle thick and raised (Figs. 11, 13); callus weakly sculptured with several long setae (Figs. 11, 12); metepimeral groove present and foveate. Femoral depression (fmd) broadly and shallowly impressed and weakly reticulate (Fig. 12); transepimeral sulcus present; transepisternal sulcus present and associated with a weak carina (Fig. 12); posterior margin of upper mesepimeron even and without any indication of the metathoracic spiracle. Prepectus (pre) associated with mesepimeron, not fused with pronotum, and in a different plane from pronotum (Fig. 12); dorsal half of prepectus ovoid with a broad foveate medial depression (Fig. 12), overlapping base of tegula (Fig. 12), ventral half strongly narrowed and associated with anterior inflected margin of mesepisternum; mesothoracic spiracle exposed and prominent (Fig. 12). Propleura broadly separated ventrally, meeting only near anterior margin (Fig. 12). Coxae smooth to weakly imbricate with scattered long setae; midcoxa less than half size of hind coxa, hind coax semiglobose and with several prominent long hairs dorsally; femora expanded medially; tibiae densely setose; fore tibia obliquely truncate apically and with one large, curved, and bifid spur; mid tibia with a single long spur almost as long as basitarsus; hind tibia with two spurs, longest half as long as basitarsus; all tarsi 5-segmented; claws simple. Forewing venation distinct (Figs. 16, 18, 19); submarginal vein with 7 long setae dorsally; parastigma with hyaline break and 2 campaniform sensilla along posterior margin (Fig. 19); stigmal vein broadened apically and with distinct uncus and 4 campaniform sensilla, stigmal vein roughly perpendicular to forewing margin; postmarginal vein extending beyond stigmal vein, but equal in length to vertical height of stigmal vein (Fig. 18); basal area bare except for track of setae along cubital vein, speculum irregular (mostly setose), costal cell with dense short setae; disc with dense fine setae and distinct marginal fringe, setal tracts on wing disc absent (Fig. 16). Hind wing venation complete, no trace of basal vein (submarginal vein parallel and continuous); 3 hamuli; fringe present.
Metasoma. Petiole short and transverse, visible only as narrow smooth band dorsally (Fig. 14), ventrolateral margins of petiolar tergite (vmpt) not fused and petiole membranous ventrally (Fig. 15); membrane attaching to anterior margin of gastral sternite; not overlapping. Gastral terga smooth to very lightly coriaceous; basal tergites (Gt,.,) tightly appressed, following tergites distinctly overlapping and articulating; Gt, depressed medially, anteriorly with medial crenulate depression, laterally with irregular patch of setae (Fig. 13); following tergites with single row of prominent setae (Fig. 9). Cerci circular with 5 setae. Sclerotized epiproct absent. Basal sternite (Gs,) with transverse crenulate sulcus (crs) and raised flat anterior region (Fig. 15); hypopygium with long, bare mucro (Fig. 20, muc). Ovipositor sheaths long and parallel (Fig. 20, 2vf and 3v), second valvifer fused dorsally with a strong internal ridge across dorsal apical margin (Fig. 23), third valvula (3v) separated from
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New genus and species of Perilampidae JESO Volume 138, 2007
second valvifer by oblique suture (Fig. 23, posterior ventral margin of 2vf indicated by arrow), and apically with tuft of setae; ventral valve abruptly expanded apically, laterally flattened and scimitar shaped with 2 apical prongs (Fig. 20, vv); dorsal valve apically with 2 pronounced basally-directed, keel-like spines (Fig. 20, dv, one spine showing) and a minute spine subapical to these.
Male. Length, 1.02 mm. Dark brown to black. Antennal scape smooth ventrally (Fig. 7), minute pores present on ventral surface of scape, but visible only at high magnification, and scape lacking pits or depressions surrounding pores (Fig. 8). Gt, subtriangular and setose. Genitalia elongate (Fig. 21); aedeagus articulating and with prominent apodemes; parameres long and thin with a single apical seta; digitus broad, rounded, and with 2 marginal spines.
Ovarian egg. (Fig. 22). Length 0.15-0.16 mm. Cylindrical without apical stalk or surface sculpture.
Holotype. Female: YEMEN, 12 km NW Manakhah, 05 May—17 June 2002, A. van Harten; deposited in CNCI. Paratypes. YEMEN: same locality and collector data, but 3 July—21 August 2001 (2), 6 July—21 August 2002 (2, 24; 2 with slide mount of antenna and one set of wings, base of metasoma on card, ovipositor in vial on pin; 6 with head on card and slide mount of wings, antenna and genitalia; body used for SEM); ISRAEL: Arava Valley, 0.2 km N Hazeva Field, N 30°46’56” E 35°14’39”, 26-27 April 1996, 450 ft, school in small wadi, M. E. Irwin (@). All specimens deposited in CNCI.
Discussion
We are confident that Jambiya belongs to the family Perilampidae. The problem remains as to which subfamily it should be associated with, or if it deserves a new subfamily status. Several features are of interest in the placement of Jambiya and also affect our current treatment of the existing subfamilies of Perilampidae.
Male scape. In all Perilampinae and Chrysolampinae, the ventral subapical region of the male scape has distinct pits that are usually visible using simple light microscopy (Darling 1986, 1983, 1988b). Each of these pits is associated with | or more pores that are likely associated with pheromone glands (Darling 1986); pores do not appear in the surrounding interstices. The male scape of Philomidinae is reticulate without any visible pores, even in slide mounts (Heraty and Darling, unpublished). Males of some Eucharitidae (some Gollumiella, Psilocharis, and Neolosbanus) have scattered ventral pores visible only in slide preparations or under high magnification using SEM (Heraty 1994, 2004), however these are never associated with pits. Males of Jambiya have minute pores scattered over the ventroapical surface that are visible only with SEM, and no pits (Figs. 7, 8); pores are absent in the female scape. The presence of these scattered pores is likely a plesiomorphic condition among the pteromaloid Chalcidoidea, and possibly all Chalcidoidea (Heraty, unpublished). The association of pores with distinct pits is the derived feature of Chrysolampinae and Perilampinae which is not present in Jambiya.
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Heraty and Darling ~ JESO Volume 138, 2007
Antennal flagellum. The antennal flagellum is similar in Chrysolampinae and Perilampinae (cf. Figs 2, 3). The groundplan segmentation of the flagellum includes a distinct transverse anellus (f1,), followed by seven articulated funicular segments (f1, ,) that are usually broader than long and slightly increasing in width apically, and finally a four segmented clava (f1, ,,) with the flagellomeres fused but clearly demarked. The apical flagellomere (fl,,) is small and button-like. Multiporous plate sensilla (MPS) and numerous stout socketed sensilla are present on at least flagellomeres 4-11 and the MPS are always absent on the anellus and flagellomere 12. Although there are exceptions within Chrysolampinae (Brachyelatus, Austrotoxeuma), the flagellum is compressed and much shorter than the head height, and often barely extending to the clypeal margin. Philomidinae differ only in that some males have ventrally ramose antennae, whereas they are always simple in other Perilampidae. In Eucharitidae, the antennal flagellum is much more elongate, and f1,, is never present in the same form (may be a complete additional segment in some more derived taxa, cf. Heraty 2002). Except for the absence of MPS on the second flagellomere, the antenna of Jambiya is similar to that of most Perilampidae; however many of these features are found in other Chalcidoidea and it is difficult to ascertain which attributes are synapomorphic for Perilampidae.
Epipharynx. Chrysolampinae and Philomidinae have two patches of epipharyngeal seta, whereas Perilampinae, all Eucharitidae, and Jambiya have a single pair of stout setae. The polarity of this character is uncertain (Darling 1988).
Labrum. The labrum of Jambiya is similar in some regards to both Chrysolampinae and Perilampinae, but not Philomidinae. A weakly sclerotized, flap-like labrum with evenly placed digits along the apical margin is known for only some Chrysolampus (Darling, unpublished). In most Chrysolampinae, digits are absent and the setae are arranged evenly along the apical margin (Darling 1988a). Within Perilampidae, aboral digits and paired transluscent areas are known only within Perilampinae (Darling 1988a). Furthermore, the labrum of Perilampinae is excised medially and arranged into two distinct lobes, with the translucent areas located along the inner margin of the lobes (Darling 1988a). Darling (1988a) proposed that aboral digits in Perilampinae were likely derived. Their presence in Jambiya, in combination with other characters, would suggest that aboral digits might be plesiomorphic for Perilampinae. The labral digits of Jambiya are finely tapered apically; they may be either tapered or spatulate in Chrysolampinae and Perilampidae; tapered digits are considered to be plesiomorphic (Darling 1988).
Pronotal-prepectal association. Jambiya and Chrysolampinae have the prepectus closely associated with the mesepimeron and broadly separated from the pronotum by a wedge- shaped gap along most of its anterior length (Fig. 12). The size of the gap is related to the orientation of the pronotum, and can be more correctly correlated with a close association of the prepectus with the mesepipleuron and not the pronotum. This form of the prepectus is typical for Pteromalinae, Torymidae, and other chalcidoids, and is presumed to be plesiomorphic. Philomidinae have a prepectus unique in Chalcidoidea. The articulation between the pronotum and prepectus is rigid and closely appressed, but the prepectus remains separate and uniquely overlaps the posteroventral margin of the pronotum. In
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New genus and species of Perilampidae JESO Volume 138, 2007
Perilampinae, Akapalinae, and Eucharitidae, the prepectus is more closely associated with the pronotum, and is fused to the pronotum in all Perilampinae (except Steffanolampus) and within Eucharitidae (Gollumiellinae and all Eucharitini) (Heraty 2002). Close association with eventual fusion of the pronotum and prepectus is considered the derived state (Heraty 2002).
Forewing venation. The wing venation of Jambiya is almost identical to that found in Chrysolampus or Chrysomalla (Chrysolampinae), including the presence of a hyaline break in the parastigma, shape of the stigmal vein and stigma, and length of the postmarginal vein (Figs. 16-19). The only difference is the linear arrangement of the campaniform sensilla along the uncus, which are clustered in Chrysolampinae (cf. Darling 1986). Perilampinae have no hyaline break, a distinctly different stigma, and a much longer postmarginal vein. The venation of Philomidinae is similar to Chrysolampinae, but without a hyaline break and they have numerous campaniform sensilla in the stigma that are arranged in a dense cluster (Heraty, unpublished).
Petiole. The petiole of Jambiya is short and smooth, with the ventral margins of the petiolar sclerite broadly separated by a membranous area, and the apex of the petiolar scperite abuts with the anterior margin of the first gastral sternite (neither overlapping or inserted). A ventrally membranous petiole is found in Chrysomalla (Chrysolampinae) and females of Indosema and Timoderus (Eucharitidae: Oraseminae). A ventrally membranous petiole is known in some Pteromalidae (cf. Heydon 1989) and whether it is a derived or plesiomorphic state is uncertain. In taxa with a fused petiole (i.e. Orasema in Oraseminae), the sternal antecostal muscles attach between the crenulate sulcus of the first gastral sternite and the ventral midline of the petiole (Heraty, unpublished). However, in Jambiya, these same antecostal muscles divide anteriorly and attach to the ventrolateral margins of the sclerotized portion of the petiole, suggesting that the remnant of the ventral (first) sternite of the petiole has been split. If so, this could be an apomorphic state within Chalcidoidea, although it is potentially plesiomorphic for Perilampidae or Perilampidae + Eucharitidae.
Gastral tergites. Perilampinae and Chrysolampinae have the gaster usually high and triangular in profile, with the first and second tergites usually similar in size, fused dorsally, and Gt, has a large lateral panel (Darling 1986, 1997). In these two subfamilies, the margins of the basal two terga are abutting and non-overlapping dorsally. Philomidinae have the basal tergite longer than the second and also overlapping the second tergite; the basal tergite (Gt,) has a lateral panel, but this may be associated more with the odd shape of the gaster, which is strongly compressed dorsoventrally. The basal gastral tergites of Jambiya are subequal in length and with Gt, abutting Gt, (Fig. 9). These terga are not fused, and there is no distinct lateral panel on Gt,. Although the basal tergite is broadly depressed medially, the gaster is rounded and not triangular (Fig. 9). Other than having closely associated and abutting basal tergites, the gaster of Jambiya is not similar to other Perilampidae.
Ovipositor. The ovipositor of Jambiya is unique within Chalcidoidea. The ovipositor valves
in Chrysolampinae and Perilampinae are simple and needle-like. An expanded ovipositor with prominent ridges or spines is associated with oviposition into cavities formed in leaf
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tissue occurs in the three subfamilies of Eucharitidae (Gollumiellinae, Oraseminae, and some Eucharitinae) and Akapalinae, although oviposition habits are unknown in this latter subfamily (Heraty 1994, 2002; Heraty et al. 2004). It is difficult to postulate a use for the peculiar ovipositor in Jambiya, however its form may suggest a habit of oviposition into leaf surfaces as is known for some Eucharitidae (Gollumiellinae, Oraseminae, and some Eucharitinae; Heraty 2002; Heraty and Quicke 1998). This raises the possibility that expanded ovipositor and leaf ovipositions are plesiomorphic for the perilampid/eucharitid complex.
Egg. Jambiya has a fusiform smooth egg (as inferred from ovarian eggs). Perilampinae have a fusiform egg with a sculptured surface (Heraty and Darling 1984; Darling and Roberts 1999) whereas Chrysolampinae have a fusiform, although slightly dumbbell-shaped egg, with a smooth surface (Darling and Miller 1991). The eggs of some Eucharitidae are fusiform and smooth, but most species have a stalked egg, both in the ovariole and after the egg is deposited (Heraty 1994, 2002). A fusiform egg is likely plesiomorphic.
None of the morphological features discussed above accurately places Jambiya within any of the perilampid subfamilies, and at best, the features of Jambiya confuse the putative synapomorphies of Chrysolampinae and Perilampinae. Although closest to Chrysolampinae, and especially the genus Chrysomalla, Jambiya lacks any of the defining characteristics of the subfamily. The antennae are similar in general form (antennal formula, compact flagellum, and dense setation) to all three subfamilies of Perilampidae, but the lack of pits on the male scape would seem to exclude this genus from either Chrysolampinae or Perilampinae. The single pair of epipharyngeal setae is shared with Perilampinae and Eucharitidae, but not Chrysolampinae or Philomidinae. The 3/2 mandibular formula excludes Jambiya from Chrysolampinae and Philomidinae. The labrum has a composite of features shared with Chrysolampinae and Perilampinae, but not Philomidinae. The pronotal- prepectal complex is likely plesiomorphic and uninformative. The wing venation is almost identical with some Chrysolampinae, but with a different arrangement of campaniform sensilla. Polarity is difficult to assess for the forewing vein features, but Jambiya is very different from either Philomidinae or Perilampinae. The petiole is membranous ventrally and similar to some Chrysolampinae and Eucharitidae, but the polarity of this feature is uncertain. The gastral tergites are closely associated and abutting, but otherwise it is not similar to other Perilampidae. The ovipositor is unique within Chalcidoidea, but has some similarities with Eucharitidae and Akapalinae that may be suggestive of oviposition into leaf tissue. Lastly, the egg is fusiform which is characteristic of Perilampinae and some Eucharitidae, but this is likely a plesiomorphic feature. In summary, Jambiya might well deserve subfamily status within Perilampidae. A combined morphological and molecular analysis is currently in progress which should resolve not only the placement of this enigmatic wasp, but also the phylogeny of the perilampid/eucharitid complex.
New genus and species of Perilampidae JESO Volume 138, 2007
Acknowledgements
This paper is dedicated to Dr. ‘DP’ David Pengelley. He was an extraordinary mentor that inspired undergraduate students to achieve goals that were far and above what they would have ever expected of themselves. JMH in particular owes his entire career to the passion for insects that was instilled by studying under DP as both an undergraduate and as one of his last graduate students. We would like to thank Gary Gibson (CNCI) and Roger Burks (UCR) for bringing these specimens to our attention. Johan Liljeblad provided comments on an earlier draft of this manuscript. We acknowledge the support of NSF grants DEB-0108245 and EF-0341149 to JMH and an NSERC Discovery Grant to DCD.
References
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Darling, D.C. 1986. Revision of the New World Chrysolampinae (Hymenoptera: Chalcidoidea). Canadian Entomologist 118: 913-940.
Darling, D. C. 1988a. Comparative morphology of the labrum in Hymenoptera: the digitate labrum of the Perilampidae and Eucharitidae (Chalcidoidea). Canadian Journal of Zoology 66: 2811-2835.
Darling, D. C. 1988b. A review of the genus Krombeinius (Hymenoptera: Perilampidae) with a reexamination of generic limits and phylogenetic relationships and the
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descriptions of two new species. Journal of the New York Entomological Society 96: 63-81.
Darling, D. C. 1992. The life history and larval morphology of Aperilampus (Hymenoptera: Chalcidoidea: Philomidinae), with.a discussion of the phylogenetic affinities of the Philomidinae. Systematic Entomology 17: 331-339.
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Gibson, G. A. P., J. M. Heraty, and J. B. Woolley. 1999. Phylogenetics and classification of Chalcidoidea and Mymarommatoidea — A review of current concepts (Hymenoptera, Apocrita). Zoologica Scripta 28: 87—124.
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Heydon, S. L. 1989. Review of Nearctic Rhicnocoelia and Callimerismus with a discussion of their phylogenetic relationships (Hymenoptera: RieeRD naa) Journal of the New York Entomological Society 97: 347-357.
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Gut bacteria from Neodiprion abietis JESO Volume 138, 2007
PRELIMINARY EXAMINATION OF GUT BACTERIA FROM NEODIPRION ABIETIS (HYMENOPTERA: DIPRIONIDAE) LARVAE |
B. WHITTOME', R. I. GRAHAM?, AND D. B. LEVIN? Department of Biosystems Engineering, University of Manitoba, E2-376 EITC, Winnipeg, Manitoba, Canada R3T 5V6 email: levindb@cc.umanitoba.ca
Abstract - J. ent. Soc. Ont. 138: 49-63
The gut microbiotas of insects are important for many processes, including digestion, nitrogen fixation, and nutrient recycling. Bacterial 16S ribosomal DNA (rDNA) extracted from excised Neodiprion abietis larval guts was amplified using PCR. Two combinations of primers produced six fragments that were separated using Denaturing Gradient Gel Electrophoresis (DGGE). The DNA fragments were sequenced directly. BLAST-n analysis and comparison-rank searches, using the Ribosomal Database Project II, revealed four predicted bacterial species, one that had similarity to Alphaproteobacteria and three that aligned with Gammaproteobacteria. Phylogenetic analysis by maximum parsimony and neighbour joining confirmed these findings and suggest that Rahnella, Yersinia, Enterobacter, and a Caulobacter-like species inhabit the N. abietis larval gut.
Published November 2007
Introduction
The balsam fir sawfly, Neodiprion abietis (Hymenoptera: Symphyta: Diprionidae), is an indigenous phytophagous insect in North America. The larvae feed predominantly on balsam fir (Abies balsamea Mill), but will also consume white spruce (Picea glauca Moench) and black spruce (Picea mariana Mill.) (Wallace and Cunningham 1995). Outbreak populations typically occur every 5-15 years, lasting 4-5 years in duration (Piene et al. 2001; Moreau et al. 2005). Larvae emerge in early summer after overwintering as eggs sheltered in the needles of the host plant. Male larvae pupate after their fifth instar, whereas female larvae may go through an additional instar before pupation. Adults emerge in late summer and, after mating, females lay eggs in current year foliage, using a saw-
‘Department of Biology, University of Victoria, Victoria, British Columbia, Canada V8W 2Y2
? Population Ecology Group, Faculty of Forestry and Environmental Management, University of New Brunswick, Fredericton, New Brunswick, Canada E3B 6C2
> Author to whom all correspondence should be addressed.
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like ovipositor. The majority of current knowledge regarding diprionid sawflies is based on ecological (Wallace and Cunningham 1995; Li et al. 2005; Moreau et al. 2005) and anatomical studies (Bordas 1895; Maxwell 1955). Current knowledge of the structure and organization of the sawfly digestive tract is limited, and almost nothing is known about the microbiota of sawfly guts.
The insect gut is a complex and highly structured organ. Gut morphology and function are dependent on several factors: the insect taxon, its stage of development, feeding behaviour of each developmental stage, food source, environment that the insect inhabits, and the inhabiting microorganisms (Wigglesworth 1972; Chapman 1985; Nation 2002; Dillon and Dillon 2003). The first comparative review of hymenopteran guts was made, by Bordas (1895), describing the macro-morphology of guts from selected insects of every family in the order Hymenoptera. Sixty years later, Maxwell (1955) compared the internal anatomy of larvae from 132 species, in eleven families of North American and European sawflies, within the suborder Symphyta. Neither of these reviews on sawfly gut morphology mentioned gut microbes.
The natural microbiota of the gut represent microbial-host interactions that range from pathogenic to obligate mutualism. Studies that define the composition of microbial communities in the digestive system have primarily been performed using termites, tsetse flies, aphids, and cockroaches (Dillon and Dillon 2003). Recent interest in insect endosymbionts, such as bacteria in the genera Wolbachia, Buchnera, or Wigglesworthia bacteria, has increased our knowledge about relationships of microbes with their insect hosts. Termite microbiota are best characterized, primarily because the functional roles of gut microbes in other insects have not been investigated (Brune 1998; Bignell 2000; Breznak 2000). In termites, microbes are mainly located within specialized regions and structures of the gut. The majority of the termite microbiota are found in pouches of the hindgut, where bacterial densities can reach 10'' cells per ml of gastric fluid (Breznak and Pankratz 1977). In the midgut, microbial communities are typically sparse and tend to localize between the microvilli of the epithelial cells (Breznak and Pankratz 1977). Microorganisms may colonize the gut wall, attach to surfaces such as spines, or course freely in the lumen (Bignell 2000). Depending on the termite species and its food source, the functional role of the microbes may range from fermentation and hydrogen production to nitrogen recycling and carbon elimination (Breznak and Pankratz 1977; Brune 1998; Bauer et al. 2000; Bignell 2000; Brauman et al. 2001).
Recent studies of gut microbiota in the ant genera Camponotus, Solenopsis, and Tetraponera (Hymenoptera: Formicidae), have shown that bacteria localize to bacteriocytes within the midgut and the pouch of the hindgut (Shannon et al. 2001; Sauer et al. 2002; van Borm et al. 2002a; Li et al. 2005). These symbiotic microbiota are members of Alpha, Beta, and Gamma divisions of Proteobacteria, as well as Flavobacteria (van Borm et al. 2002a), and including a novel candidate genus, Blochmannia (Sauer et al. 2000; Sauer et al. 2002). Classification of these microbes was accomplished by culture-independent methods as these bacteria often cannot be cultured outside of their hosts (Schroder et al. 1996). In addition, media for culturing has typically been developed for medical studies and the growth conditions for fastidious microorganisms are often lacking, leading to misrepresentative sampling of the gut microbiota (Dillon and Dillon 2003). To surmount these difficulties, sequence analysis of 16S ribosomal DNA (rDNA) has become widely accepted as a tool
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for investigating unculturable microbes in these often complex communities (Hongoh et al. 2003a, b).
This manuscript represents the first preliminarily analysis of the gut microbiota of a diprionid species. PCR was used to amplify bacterial 16S ribosomal DNA (rDNA), extracted from excised larval guts, and along with Denaturing Gradient Gel Electrophoresis (DGGE) revealed 4 distinct DNA products. BLAST-n analysis and comparison rank searches of the Ribosomal Database Project II (RDP II) database showed similarity to Alphaproteobacteria and Gammaproteobacteria. Maximum parsimony and neighbour joining analyses confirmed these observations.
Materials and Methods
Larval collection
Balsam fir branches, containing Neodiprion abietis larvae, were collected from forest stands near Old Man’s Pond (near Corner Brook), Newfoundland, Canada (N 49° 05°59’ W 57° 56°05’). Larvae were maintained on balsam fir in paper bags at 4°C. Head capsule widths of healthy larvae were measured using a dissecting microscope with a calibrated objective. Larvae with head-capsule widths between 0.68-1.4 mm, corresponding to 2™ to 4" instar larvae, were harvested for histological preparation and extraction of total DNA from the excised gut.
PCR amplification, DGGE, and sequencing of bacterial /6S gene
Larvae harvested for molecular characterization of sawfly-gut bacteria were surface sterilized with a 60 second wash in 5% bleach, followed by a 60 second rinse in DEPC-treated water (0.1% diethyl pyrocarbonate). Larvae were submerged in sterile phosphate buffered saline (PBS, pH 7.4), and anterior and posterior segments were excised just posterior of the head capsule and immediately anterior to the eighth proleg, respectively. The cuticle was secured and the gut was pulled from the body cavity. The excised gut was transferred to fresh PBS and the peritrophic membrane, containing the food bolus, was pulled from the gut lumen using forceps. The gut tissue was immediately placed into RNAlater (Ambion Inc., Austin, Texas) and stored at -20°C.
DNA was purified using TRIzol (Invitrogen Co., Burlington, Ontario), following the manufacturer’s protocol. Two primer sets were used to ensure amplification of the targeted bacterial 16S rDNA. Primers p984f-GC (5’-CGCCCGGGGCGCGCCCCGGGCGGGGCGG GGGCACGGGGGGAACGCGCCGAACCTTAC-3’) and pl40Ir (5’-GCGTGTGT ACAAGACCC-3’) were used to amplify the V6 to V8 regions of 16S ribosomal DNA (Nobel et al. 1996; Frederick and Caesar 2000). Primers p515f-GC (5’-CGCCC GGGGCGCGCCCCGGGCGGGGCGGGGGCACGGGGGGCCAGCAGCCGCGGTAA -3’) and p806r (5’°-GGACTACCAGGGTATCTAAT-3’) were used to amplify the variable V4 region of 16S rDNA (Relman 1993). PCR mixtures of 50 ul volume contained reaction buffer (10 mM Tris-HCI pH 8.3 at 25°C, 50 mM KCl, 1.5 mM MgCl, 0.001% gelatin), 10 uM each of dATP, dTTP, dCTP, and dGTP, 0.1 uM of each primer, | unit Zag polymerase (Qiagen, Mississauga, Ontario) and approximately 10 ng insect genomic DNA template. PCR was conducted using a Mastercycler EP thermal cycler (Eppendorf, Mississauga,
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Ontario), with the following settings: (1) 94°C for 5 min, | cycle; (11) 94°C for 30 sec, 52°C for 30 sec, 72°C for 45 sec, 40 cycles; (iii) 72°C for 5 min, 1 cycle. On completion of thermal cycling, 10% of the reaction was loaded on a 1% agarose gel and electrophoresed in 1X TBE buffer (90 mM Tris Borate, pH 8.3, 2 mM EDTA) for 2 hrs at 60 V. The gel was stained with ethidium bromide and visualised using UV illumination.
Subsequently, the PCR products (50% of the reaction) were separated by DGGE using the DCode system (BioRad) according to the: manufacturer’s instructions. Gels consisted of 1 mm thick 6% polyacrylamide with a denaturing gradient of 30-70 % (100% denaturant corresponds to 7 M urea and 40 % vol/vol deionized formamide) and 1X TAE buffer (90 mM Tris Acetate, pH 8.3, 2 mM EDTA) for 16 hours. Electrophoresis was performed at 60°C and 80 V in 1X TAE running buffer for 16 hours. Gels were stained with SYBR Gold nucleic acid stain (Invitrogen) for 30 minutes and images captured upon UV illumination. DNA bands were excised with a sterile razor blade and placed in 100 ul of sterile distilled H,O. The samples were placed at 94°C for 5 minutes to elute the DNA from the polyacrylamide and were stored at 4°C overnight. Five uL of the supernatant were used as template to reamplify the individual DNA bands. The PCR conditions were the same as above, but with only 30 cycles of amplification. The PCR products were gel purified using the QIAquick Gel Extraction Kit (Qiagen), and samples stored at -20°C until ready for sequencing. Sequencing was performed by Ontario Genomics Innovation Centre, using an ABI 3730 DNA Analyzer (BigDye version 3.1).
Sequence data was analysed using BLAST-n (http://www.ncbi.nlm.nih.gov/ BLAST) and the Similarity Rank program of the RDP II (http://rdp.cme.msu.edu/seqmatch/ seqmatch intro.jsp) (Maidak et al. 1999), to determine similarity with known bacterial species (in the database). Closely related species, as well as gut microbiota listed in recent publications (Boursaux-Eude and Gross 2000; Sauer et al. 2000; Shannon et al. 2001; van Borm et al. 2002b; Hongoh et al. 2005), were used to construct phylogenetic trees using neighbour joining and maximum parsimony algorithms, with 1000 bootstrap replicates. Phylogenetic and molecular evolutionary analyses were conducted using MEGA version 3.1 (Kumar et al. 2004).
Results
DGGE separated four 16S rDNA fragments when using the p984f-p1401r primer set (Table 1, #1 to 4), and two fragments after amplification from the p515f-p806r primer set (Table 1, #5 and 6). BLAST-n analysis and similarity rank comparisons to the RDP II sequence database predicted four bacteria matches: Rahnella sp. (sequence #1, GenBank Accession No. EF140875), Yersinia sp. (sequences #2-4, GenBank Accession No. EF 140876- EF 140878 ), an Enterobacteriaceae (sequence #5, GenBank Accession No. EF140880), and an Alphaproteobacteria (sequence #6, GenBank Accession No. EF140879).
Phylogenetic analysis confirmed the predicted identities of the first four bacteria and showed their close relationship to other known insect-gut microbes in the Enterobacteriaceae family of Gammaproteobacteria. Maximum parsimony and neighbor joining analyses suggest that sequence #1 was most closely related to Rahnella aquatilis. Both analyses weakly supported the clustering of sequences #2-4 with Yersinia, with the degree of their
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Gut bacteria from Neodiprion abietis
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Uncultured N. abietis bacteria # 2 Uncultured N. abietis bacteria# 3 | —-———— Uncultured N. abietis bacteria # 4 | Yersinia aleksiciae Wigglesworthia ssp. of Glossina brevipalpis Yersinia pestis Brenneria quercina 79 Symbiont of Formica fusca Serratia maracens 52 Rhanella spp.
54 Yersinia rhodei Plesiomanas shigelloides 64 Candidatus Blochmannia silvicola Candidatus Blochmannia rufipes Candidatus Blochmannia socius Candidatus Blochmannia herculeanus Buchnera aphidicola Uncultured sheep mite bacteria Unclassified Pseudomonadaceae of Tetraponera binghami 51 Candidatus Clostridium massiliensis Candidatus Clostridium timonensis Clostridium thermocellum Chloroflexus aggregans Bacillus megaterium 61 87 Bartonella henselae 69 Wolbachia melophagi Unclassified Rhizobiaceae of Tetraponera binghami Wolbachia maritima Unclassified Methylobacteriaceae of Acromyrmex octospinosus sd 79 Caulobacter leidyia 99 Unidentified eubacterium AM084885 Unidentified eubacterium of Myrmeleon mobilis DQ163946 Unidentified eubacterium AJ459874 Burkholderia spp. of Tetraponera binghami Wolbachia inokumae Wolbachieae incompatibility symbiont of Nasonia vitripennis
96
77
53
FIGURE 1. Phylogenetic analyses of bacterial 16S ribosomal DNA gene sequences amplified from insect guts. Maximum parsimony (A and C) and neighbour joining trees (B and D) were inferred using the Mega 3.1 program with 1000 bootstrap repetitions. Support values >50% are listed at nodes. Sequences of 16S ribosomal DNA from the microbiota of N. abietis larval guts are indicated with arrows. Bacteria identified by sequences #1-— 4 (from primer set p984f-GC/p140Ir) are represented in Trees A and B, while bacteria identified by sequences #5 and #6 (primer set p515f-GC /p806r) are represented in Trees C and D. Boxes indicate groups referred to in Table 1.
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Uncultured WN. abietis bacteria # 2 B 83 Yersinia aleksiciae 52
Yersinia pestis 62 Brenneria quercina 85 Symbiont of Formica fusca Serratia maracens 64 Rhanella spp. 87 Uncultured N. abietis bacteria # 1 Rhanella aquatilis 69 rersinia rhodei Plesiomanas shigelloides 87 Candidatus Blochmannia silvicola Candidatus Blochmannia rufipes Candidatus Blochmannia socius Candidatus Blochmannia herculeanus Buchnera aphidicola Burkholderia spp. of Tetraponera binghami 99 Uncultured sheep mite bacteria Unclassified Pseudomonadaceae of Tetraponera binghami Flavobacterium of Tetraponera binghami 99 — Wolbachia inokumae Wolbachieae incompatibility symbiont of Nasonia vitripennis 97—=_ Bartonella henselae 91 Wolbachia melophagi Unclassified Rhizobiaceae of Tetraponera binghami Wolbachia maritima Unclassified Methylobacteriaceae of Acromyrmex octospinosus 91 Caulobacter leidyia 99 Unidentified eubacterium AJ459874 Unidentified eubacterium of Myrmeleon mobilis DQ163946 Unidentified eubacterium AM084885 Bacillus megaterium Chloroflexus aggregans 80 Candidatus Clostridium massiliensis 56—_§$.. Candidatus Clostridium timonensis Clostridium thermocellum
FIGURE 1. Continued
5D
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¢ Yersinia pestis 2 Unidentified eubacterium A Y537571
Plesiomanas shigelloides Uncultured NV. abietis bacteria # 5 Yersinia aleksiciae
Yersinia rhodei Rhanella aquatilis Brenneria quercina Rhanella spp. DQ217650 Serratia maracens
andidatus blochmannia socius Candidatus Blochmannia herculeanus Candidatus Blochmannia rufipes Candidatus Blochmannia silvicola Wigglesworthia ssp. of Glossina brevipalpis
Buchnera aphidicola
Burkholderia spp. of Tetraponera binghami 92 Uncultured sheep mite bacteria
Unclassified Pseudomonadaceae of Tetraponera binghami 97 Candidatus Clostridium massiliensis
Candidatus Clostridium timonensis 96 Wolbachia inokumae Wolbachieae incompatibility symbiont of Nasonia vitripennis Wolbachia maritima Clostridium thermocellum Chloroflexus aggregans Lactococcus lactis Bacillus subtilis 52 Bacillus megaterium 53 Lactobacillus maltaromicus Carnobacterium piscicola
67
Caulobacter leidyia Bartonella henselae
67 Wolbachia melophagi Unclassified Rhizobiaceae of Tetraponera binghami Unclassified Methylobacteriaceae of Acromyrmex octospinosus Unclassified Alphaproteobacteria DQ516567
80
Unidentified eubacterium of Myrmeleon mobilis DQ163946 Unidentified eubacterium AM084885
Unidentified eubacterium AJ459874
Unidentified eubacterium AJ874181
Unclassified Caulobacteria AY807064
Uncultured N. abietis bacteria # 6
FIGURE 1. Continued
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60 98 99 73 88 58 55 95 58
FIGURE 1. Continued
JESO Volume 138, 2007
Unidentified eubacterium AJ459874
Unidentified eubacterium AM084885
Unidentified eubacterium of Myrmeleon mobilis DQ163946 Uncultured N. abietis bacteria # 6 Unclassified Caulobacteria AY807064 Unidentified eubacterium AJ874181
—
Unclassified Alphaproteobacteria DQ516567
Bartonella henselae
Wolbachia melophagi
Unclassified Rhizobiaceae of Tetraponera binghami Unclassified Methylobacteriaceae of Acromyrmex octospinosus
Flavobacterium of Tetraponera binghami
Candidatus Clostridium massiliensis
Candidatus Clostridium timonensis
Clostridium thermocellum
Lactococcus lactis
Bacillus subtilis
Bacillus megaterium
Lactobacillus maltaromicus
Carnobacterium piscicola
Wolbachia maritima
Wolbachia inokumae
Wolbachieae incompatibility symbiont of Nasonia vitripennis Candidatus Blochmannia rufipes
Buchnera aphidicola
Candidatus Blochmannia silvicola
Wigglesworthia ssp. of Glossina brevipalpis
Burkholderia spp. of Tetraponera binghami
Uncultured sheep mite bacteria
Unclassified Pseudomonadaceae of Tetraponera binghami Candidatus Blochmannia socius
Candidatus Blochmannia herculeanus
erratia maracens Yersinia aleksiciae
Yersinia rhodei
Uncultured N. abietis bacteria #5 Brenneria quercina
Plesiomanas shigelloides
Yersinia pestis
Unidentified eubacterium AY537571 Rhanella aquatilis Rhanella spp. DQ217650 oroflexus aggregans
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relatedness to Yersinia aleksiciae varying (Table | and Figure 1). The three 16S rDNA sequences showed 98.7% identity to each other and were approximately 320 bp in length.
The identity of the bacterium from which sequence #5 was derived was not determined beyond Enterobacteriaceae because results from maximum parsimony and neighbor joining analyses were inconsistent. The difficulty in confirming the identity of this bacterium may be a result of the small amplicon size (180 bp). However, sequence #6 (184 bp), which was amplified with the same primer set, clearly clustered with the Alphaproteobacteriaceae. Maximum parsimony analysis suggested that bacteria, from which sequence #6 was derived, belonged to the genus Caulobacter.
Discussion
The microbiota identified by 16S rDNA sequences from N. abietis gut tissues include those that have been found ubiquitously in the environment and likely originated from the host’s diet (Selenska-Pobell et al. 1995; Dillon and Charnley 2002; Sprague and Neubauer 2005). Similarly, other free-living microbial species have been isolated from other sawfly gut tissues, including Pristiphora geniculata, Acantholyda erythrocephala, and Pikonema alaskensis (R. Graham, unpublished data). Fragments #1-5 (based on 16S rDNA sequences) represent bacteria that belong to the Gammaproteobacteria, specifically those in the Enterobacteriaceae family of Gram-negative, anaerobic microbes.
Neither Rahnella aquatilis nor Yersinia aleksiciae have been published as insect gut microbes, although R. aquatilis has been isolated from both chicken ticks (Montasser 2005) and the intestinal contents of snails (Brenner et al. 1998). An uncultured Rahnella sp. was reported in GenBank (Accession # U84730) from an isolate of the microbial gut flora from the coleopteran genera Phaleria and Latreille (Tenebrionidae). Rahnella spp. have been isolated from foliage (Hashidoko et al. 2002; Izumi et al. 2006) and ferment several polysaccharides (Brenner et al. 1998). Additionally, Rahnella spp. have been recognized as strong nitrogen fixers (Brenner et al. 1998; Izumi et al. 2006). This characteristic would be important for nitrogen recycling in nutrient-poor diets and possibly promote its retention as a symbiont within the gut, perhaps originally acquired through the sawfly’s diet.
Species of Yersinia have been isolated from other insect guts (Ulrich et al. 1981); therefore it is not surprising that we found related bacteria in the gut of N. abietis. No beneficial characteristics have been attributed to Yersinia. Their ubiquitous presence in soils and detritus suggest that this bacterium is more likely to be a transient microbe ingested with food matter, rather than part of the permanent flora of the sawfly gut.
16S-sequence analysis of fragment #5 and subsequent phylogenetic comparisons to other Gammaproteobacteria was inconsistent and poorly resolved. Maximum parsimony indicated that the closest relative to the N. abietis bacteria was Plesiomanas shigelloides, while neighbour joining analysis suggested that Y. rhodei was more closely related. BLAST-n searches of the 16S ribosomal sequence commonly aligned Serratia spp. with high degrees of identity (97%). Yersinia, Rahnella, and Serratia spp. have been shown to cluster closely together in a Group B of the enterobacterial genera, with the main signature nucleotides located between positions 590-649 (Sproer et al. 1999). The p515f-p806r primer set amplifies the variable V4 region of 16S rDNA between base pairs 627 and 807. This
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region only overlaps the signature nucleotides by 22 bp, making a positive identification difficult. Therefore bacterial sequence #5 can only be classified as an Enterobacteriaceae until further data is collected. .
Finally, 16S-sequence analyses indicated that bacterium #6 was a member of the Alphaproteobacteria, showing high similarity with uncultured bacteria of insect larvae and soil (GenBank AJ459874, DQ163946, and AM084885; Table 1; Figure 1 C, D). An uncultured Caulobacter (GenBank AY 807064) aligned within the unidentified Alphaproteobacteria, supported by a 98% bootstrap value, suggesting that the N. abietis bacterium #6 may be Caulobacter-like. Although Caulobacteria have typically been isolated from aquatic environments, a few isolates have been reported from the intestinal contents of a millipede (Abraham et al. 1999) and the mite Jetranychus urticae (Hoy and Jeyaprakash 2005). If N. abietis bacterium #6 is a Caulobacter, this microbe may play a key role in nutrient acquisition since Caulobacteria have been shown to uptake phosphorus from nutrient-poor environments (Gonin et al. 2000). Chemical analyses of current year foliar nutrients have reported phosphorous levels at 900-4000 ppm along the eastern US coastline and in the Laurentide-Onatcheway region of Québec, Canada (Bauce et al. 1994; Richardson 2004). Although foliar chemical data for balsam fir growing in Newfoundland could not be found, it is known that phosphorous levels decline rapidly in trees growing in harsh conditions (Richardson 2004). Due to the often severe climate of Newfoundland, one would predict phosphorous levels at the lower end of the range reported.
The diversity of the gut microbiota of N. abietis, using a PCR prospecting approach, is relatively low compared to the variety of microbes observed in termite and cockroach guts (Cruden and Markovetz 1984; Hongoh et al. 2003a). Approximately 270 phylotypes have been detected in the gut of Reticulitermes speratus and the bacteria were classified into 9 of the 20 phyla of eubacteria (Hongoh et al. 2003a). In contrast, only 6 phylotypes were detected in N. abietis and were classified within a single eubacterial phylum (Proteobacteria).
Although low levels of bacterial diversity within insect guts are not uncommon, the microbiota are generally composed of multiple phyla. The gut of the gypsy moth, Lymantria dispar (Order Lepidoptera) has a microbial diversity that ranges from 7 to 15 phylotypes, depending on its diet source (Broderick et al. 2004). A total of 13 genera were identified from larvae feeding on all diet sources and were classified within the Actinobacteria, the Bacteroidetes/Chlorobi group, Firmicutes, and Proteobacteria. Similar results were obtained from cultured isolates and 16S sequence analysis of microbes detected within the midgut of Culex quinquefasciatus (Order Diptera), where bacteria from 13 genera were identified (Pidiyar et al. 2004). The majority of mosquito bacteria belonged to the Gammaproteobacteria class (60% of cultured and 46% of culture-independent), while Actinobacteria and Firmicutes constituted the remainder of the bacterial types.
While diet influences the acquisition of bacterial flora observed in insect guts, morphology is often a significant factor affecting the diversity of the gut microbiota. Many termites and cockroaches have evolved complex and convoluted guts (Wigglesworth 1972; Brune and Friedrich 2000) that allow the retention of bacteria in specialized fermentation structures. Insects possessing simple and straight alimentary canals, such as the Diprionidae, Lepidoptera, and many Diptera, generally have a lower diversity of gut microbes (Dillon and Dillon 2003). Due to the selective diet of N. abietis and the simple morphology of its gut, the low level of bacterial diversity is not unexpected.
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Nonetheless, 16S rDNA sequencing and phylogenetic analyses identified six phylotypes in the larval gut of the balsam fir sawfly; four of the bacteria were clustered with Rahnella sp. and Yersinia sp., while the other two bacteria were determined to belong to the Enterobacteriaceae and Caulobacteriaceae. Whether or not they are present as obligate endobionts, they may variously play significant roles as associated microflora in sawfly larvae.
Acknowledgements
This work was supported by grants from the BioControl Network of Canada and the National Sciences and Engineering Research Council of Canada (NSERC). The authors would like to thank Chris Lucarotti for his valuable comments and editing.
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Dillon, R. J. and V. M. Dillon. 2003. The gut bacteria of insects: nonpathogenic interactions. Annual Review of Entomology 49: 71—92.
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Hashidoko, Y., E. Itoh, K. Yokota, T. Yoshida, and S. Tahara. 2002. Characterization of five phyllosphere bacteria isolated from Rosa rugosa leaves, and their phenotypic and metabolic properties. Bioscience Biotechnology and Biochemistry 66: 2474— 2478.
Hongoh, Y., P. Deevong, T. Inoue, S. Moriya, S. Trakulnaleamsai, M. Ohkuma, C. Vongkaluang, N. Noparatnaraporn, and T. Kudo. 2005. Intra- and interspecific comparisons of bacterial diversity and community structure support coevolution of gut microbiota and termite host. Applied and Environmental Microbiology 71: 6590-6599.
Hongoh, Y., M. Ohkuma, and T. Kudo. 2003a. Molecular analysis of bacterial microbiota in the gut of the termite Reticulitermes speratus (Isoptera; Rhinotermitidae). FEMS Microbiology Ecology 44: 231-242.
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Izumi, H., I. C. Anderson, I. J. Alexander, K. Killham, and E. R. Moore. 2006. Endobacteria in some ectomycorrhiza of Scots pine (Pinus sylvestris). FEMS Microbiology Ecology 56: 34-43.
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Revision of North American Sobarocephala JESO Volume 138, 2007
REVISION OF THE NORTH AMERICAN SOBAROCEPHALA (DIPTERA: CLUSIIDAE, SOBAROCEPHALINAE)
OWEN LONSDALE'AND STEPHEN A. MARSHALL? Entomology Department, Smithsonian Institution, National Museum of Natural History, Rm. CE-607, 10" & Constitution Ave. NW, Washington, D.C., 20560-0168 email: Neoxabea@hotmail.com
Abstract J. ent. Soc. Ont. 138: 65-106
The 17 species of Sobarocephala Czerny, 1903 occurring in the Neartic north of Mexico are reviewed and keyed. Sobarocephala wirthi spec. nov. and S. pengellyi spec. nov. are described from the eastern United States. Descriptions, illustrations, and distribution maps are provided for all Nearctic species.
_ Published November 2007
Introduction
Sobarocephala Czerny, 1903, as redefined by Lonsdale and Marshall (2006), is a large, predominantly neotropical genus with 17 Nearctic species and at least a dozen described and undescribed species from the Afrotropical, Australian, and Oriental Regions. The Nearctic species do not form a monophyletic group, but instead belong to a number of independent lineages, several of which are tropical in origin. The North American fauna, including two new eastern species allied to S. /atifrons (Loew), is here revised.
North American Sobarocephala species are small (2.4-5.6 mm), thin, pale yellow flies, often with distinctive brown patterning. Two male genitalic characters define the genus (a basal shield on the distiphallus and a “thumb” on the lateral lobe of the distiphallus (Lonsdale and Marshall 2006)), but Sobarocephala is most easily separated from other Nearctic Clusiidae by the absence of interfrontal bristles (found in Clusiodes Coquillett, 1904 and Clusia Haliday, 1838), the possession of inclinate anterior fronto-orbital bristles (reclinate in Clusiodes and Craspedochaeta Czerny, 1903), and the presence of a well- developed dorsal preapical bristle on the mid tibia (absent in Clusia and Heteromeringia Czerny, 1903). A key to the North American genera of Clusiidae was provided in Sods (1987); we follow Woodley (1984) in treating the single Nearctic Chaetoclusia (C. affinis Johnson) identified in that key as Sobarocephala.
Although the Nearctic Sobarocephala were recently treated by Sabrosky and Steyskal (1974), considerable new material has become available since then, allowing us
' Author to whom all correspondence should be addressed. Insect Systematics Lab, Department of Environmental Biology, University of Guelph,
Guelph, Ontario, Canada NIG 2W1
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to reconsider the Nearctic fauna on the basis of over 1200 specimens. We here add two species to the genus, provide an updated key, list new distributional data, and draw attention to previously unrecognized sister-group relationships.
Materials and Methods
Material from the following institutions was examined for this revision: California Academy of Sciences, San Francisco (CASC); Carnegie Museum of Natural History, Pittsburgh (CMNH); Canadian National Collection, Ottawa (CNCI); University of Guelph Insect Collection, Guelph (DEBU); Entomological Museum of Utah State, Logan (EMUS); Instituto Nacional de Biodiversidad, Santo Domingo de Heredia (INBC); Museum of Comparative Zoology, Cambridge (MCZC); North Carolina State University, Raleigh (NCSU); Texas A&M University, College Station (TAMU); Tel Aviv University, Tel Aviv (TAUI); United States National Museum, Washington, D.C. (USNM).
Specimen preparation and terminology follows that in Lonsdale and Marshall (2006). The M.,,, ratio is defined as the length of the ultimate section of vein M divided by the length of the penultimate section. Size ranges include both sexes. Label data for non-type material is not listed for the relatively abundant S. flaviseta (Johnson), S. latifacies Sabrosky & Steyskal, and S. /atifrons (Loew). Maps only include Nearctic records.
Generic Diagnosis
Pedicel with pronounced outer and inner angulate extensions. Cell bm open or closed. Interfrontal bristle absent. Anterior fronto-orbital bristle inclinate. Vein R, bare dorsally. Thorax usually with two postsutural dorsocentral bristles (rarely one or three in some tropical species). Presutural intra-alar bristle absent or weak in Nearctic species. Mid tibia with preapical dorsal bristle. Male 6" spiracle moved into membranous region anterior to annulus. Ventrolateral lobe of hypandrium setose and relatively large. Distiphallus almost always with basal shield (present in all Nearctic species); lateral lobe of distiphallus often with “thumb” (Fig. 20).
Key to the species of Sobarocephala north of Mexico
l. Arista densely plumose (Figs. 1, 2,6, 7). Bristles usually yellow. Gena eee shiny. First flagellomere with dorsal stripe or spot around base of arista .............. AY 3 — Arista sparsely plumose or pubescent (Figs. 3—5). Bristles light lotto to erry Gena pilose to silvery tomentose. First flagellomere entirely yellow dorsally (S. texensis and some S. setipes with light infuscation around base of arista and S. flava females with anterior margin lightly pigmented) ..................eeeeeeeseeeeeeseeeeseeeeeseeees > Z: Ocellar bristles minute to absent. Scutum with two pairs of spots; scutellum brown, at least in part. Sides of frons converging posteriorly. Female tergite 8 yellow. Surstylus with tubercle—like bristles apical. Southern United States ................8. 3
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- Ocellar bristles well-developed. Scutum and scutellum yellow. Sides of frons parallel. Female with basal half of tergite 8 brown. Surstylus with tubercle-like bristles along length of distal and posterior margins. Eastern United States ........... 4
3 Scutellum, laterotergites, and epandrium entirely brown (Fig. 1). Gena shiny. Cerci flush with distal margin of epandrium (Fig. 28). Surstylus subrectangular with long palisade-like bristles along inner-distal margin (Figs. 27, 28). Distiphallus large and spoon-shaped; lateral lobe without thumb (Fig. 29). Southeastern United States, Central America, Bahamas, Colombia ...........................9. quadrimaculata So6s
~ Scutellum brown laterally (Fig. 2), laterotergites brown lateral to scutellum, and epandrium yellow. Gena silvery tomentose. Cerci projecting (Fig. 25). Surstylus triangular with apical tubercle-like bristles (Figs. 24, 25). Distiphallus relatively small and thin; lateral lobe with thumb (Fig. 26). Florida ...........0.ccecccecesssceeesseeeeeees mre tae OR UW, SLU IRGR LQ Raii tcc ea. UE S. cruciger Sabrosky & Steyskal
4. Bristles and epandrium yellow (Fig. 6). Wing clear. Hind tibia and tarsi yellow. Frons yellow. Occiput occasionally pilose. Eastern United States 42.29... id:
RR SO ah NL a le S. affinis (Johnson)
- Bristles ind sereondnitdin brown (Fig. 7). Wing very lightly clouded on distal 1/3 along R,,,. Hind tibia and tarsi sometimes brownish. Frons sometimes with anterior margin white. Occiput usually pilose. Eastern and southeastern United States SEENON A RIAN Re OMNI: Sos cniva daaansadsase aces S. muesebecki Sabrosky & Steyskal
3; Sides of frons converging posteriorly. Notopleuron and/or scutellum sometimes with een aET RS esate Milan lisa Gaile tel) Je. hosluck! AL RAL tic 6
- Sides of frons parallel. Neither notopleuron nor scutellum with white spots ..........8
6. Scutellum white. Scutum with brown supra-alar spot and basal quadrate stripe; notopleuron, postpronotum, and posterolateral scutal spots white (Fig. 5). Arista pubescent. Florida, Dominican Republic ............8. atrifacies Sabrosky & Steyskal
— Scutellum yellow. Scutum predominantly yellow, sometimes with brown markings; white markings, if present, restricted to notopleuron and postpronotum (Fig. 3). Arista short-plumose .. Uae MMe oon
h Bristles brown. Cell - edosedl) iNest saliote sia potipleunon anid postpronotum white, at least in part (Fig. 3). Fore tarsi yellow. Female abdomen yellow with thin central stripe on tergite 3 and with wide light brown stripe (narrowing anteriorly) on tergites 4 and 5. Tubercle-like bristles on surstylus confined to apex (Fig. 19). Widespread mv NortlvAmeriva ! oti i0.0. ek ed ee S. flaviseta (Johnson)
— Bristles black. Cell bm open. Notum yellow, sometimes with lateral margin brown (Fig. 4). Fore tarsi light brown, at least apically. Female abdominal tergites 2-5 with small brown anteromedial spot; tergites 6 and 7 dark brown. Tubercle-like bristles on surstylus extending onto agg margin (Fig. 22). Northeastern North America and Utah .. 6. vissesessssseeeeeesedd» latifacies Sabrosky & Steyskal
8. Scutellum betes hie (Hig. 14). Frons sometimes with orange tint (darkest posteriorly). Scutum with one pair of wide postsutural stripes (connected posteriorly). Basicon North Atmerica y).) 00100. 8200. Wa. S. dreisbachi Sabrosky & Steyskal Scutellum yellow, at least in part. Frons yellow. Pattern on scutum variable ......... 9
9. Scutellum usually with central brown stripe (sometimes only brown medially or apically), but if yellow with distal infuscation on ventromedial surface, then surstylus
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14.
hook-shaped (some S. /atifrons; te 48) or subtriangular (some S. wirthi; Fig. 45)
Scutellum entirely yellow or brown lateiuliys never yellow with ventromedial infuscation. Surstylus — rounded or bifid, but never hook-like or triangular d 14
Stina on » scmplinp peer at i anlipeiat (Bigs fbi 17). Austad Saidtiis ahecth
Female tergite 2 with wide central stripe. Male abdomen with wide central stripe on tergites 2-5. Male sternite 5 with posteromedial comb of bristles. Surstylus small, lobate, and without tubercle-like bristles (Fig. 60) ............ccccceceecceeeeeesseceeeeeeessseeees 1]
Stripe on scutellum evenly brown or darkest at apex (Fig. 10). Acrostichal bristle present. Female tergite 2 brown along posterior margin or with thin central stripe. Male abdomen entirely yellow or with variable light brown pattern. Male sternite 5 without comb. Surstylus at least 2/3 length of ap with tubercle-like bristles along distal 2/3 of posterior margin (Fig. 43) .. ihe sake 2 Scutum yellow with one pair of brown spots on latonanan ie Gemunieon® one pair of postsutural stripes (Fig. 15). First flagellomere yellow. Bristles light brown. Fore tibia yellow. Occiput pilose. Small bristle in front of anterior dorsocentral. Female tergite 6 with wide central stripe. Laterotergites sometimes brown laterally. Distiphallus short, and lateral lobe small and ovate (Fig. 61). Eastern North America amin Ab Mttawaleciss caval Aik 6..420ir es ARKS S. setipes Melander & Argo (in part) Scutum yellow, sometimes with light brown prescutellar spot extending onto scutellum (Fig. 17). First flagellomere with light infuscation around base of arista. Bristles brown. Fore tibia light brown. Occiput shiny. No small bristle in front of anterior dorsocentral. Female tergite 6 yellow. Laterotergites yellow. Distiphallus longer than half length of phallapodeme, and lateral lobe well-developed (Fig. 58). Tekasiiaale wea wie... obdneedh eau S. texensis Sabrosky & Steyskal Bristles light brown. Scutum with light brown lateral markings (Fig. 12). Male face light brown medially and female face sometimes orange medially. Epandrium dark brown. Surstylus small, rounded, and 2/3 length of epandrium (Fig. 42). Female tergite 6 brown, and tergites 2-4 yellow except for thin central brown stripe. Alabama, Georpiew aivieiaio discs. ds cvallev. Wicket og nde S. pengellyi spec. nov. Bristles dark brown to black. Scutum with variable lateral markings. Face yellow in both sexes. Epandrium predominantly or entirely yellow. Surstylus almost as long as epandrium. Female tergite 6 brown laterally and (sometimes) basally, and tergites 2-4 dark laterally (sometimes also dark medially). Eastern United States ............. 13 Surstylus hook-shaped (Fig. 48). Female abdomen sometimes with central brown stripe in addition to lateral markings (Fig. 10). Epandrium sometimes with small
basalspot sss Anh. el 4a ee ee has. S. latifrons (Loew) Surstylus broad and rounded apically (Fig. 45). Female abdomen yellow medially (Fig. 11): . Epandrium entirely yellow .:¢1.22i2240 aetowleuleling S. wirthi spec. nov.
Female abdomen mostly or entirely yellow on tergites 1-5 and dark brown on tergite 6. Male abdomen entirely yellow and gena thin (height less than half that of first flagellomere). Surstylus large, lobate, and widest distally. Male cerci rounded and slightly projecting wwiJak. os eeqewe een. beeen dliw.. see. Jogleeiee
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16.
Female abdomen with wide central stripe on tergites (2) 3-6 (interrupted on tergite 5 in some S. lachnosternum and on tergite 6 in S. interrupta). Male abdomen with wide central stripe, but if entirely yellow, gena as high as first flagellomere. Surstylus less than half length of epandrium, small, and rounded or bifid. Male cerci flush with distal margin of epandrium or sunken .. te satcoheg ..16 First flagellomere yellow in both sexes, \ eit anterior margin n lightly oa SS in females (Fig. 8). Wing lightly (sometimes indistinctly) clouded along R,,, and costa (darkest distally). Female Se aes Eastern United States, Texas, Utah
i ...9. flava Melander & Argo First Racclidmere fark ae to i blacks on jereal half (Fig. 9). Wing clear. Female laterotergites brown, at least in part. Northeastern North America, Kansas ............... EN eS Se ee: ane. eon S. atricornis Sabrosky & Steyskal Postsutural stripe serrate on anterior margin. Scutellum brown laterally (rarely yellow) (Fig. 13). Female tergite 6 yellow, sometimes with anteromedial spot. No small bristle in front of anterior dorsocentral. Anterior lateral scutellar bristle minute or absent. Surstylus bifid (Fig. 51). Male cerci flush with distal margin of epandrium (Fig. 52). Posterior margin of male sternite 5 without posteromedial comb. Southeastern United States, Illinois ......... S. interrupta Sabrosky & Steyskal Postsutural stripe entire and rounded on anterior margin, if present. Scutellum usually yellow, but if lateral corner brown (some S. setipes), small bristle present in front of anterior dorsocentral. Female tergite 6 brown, at least medially. Anterior lateral scutellar bristle not much smaller than posterior bristle, if at all. Surstylus rounded. Male cerci sunken below distal margin of epandrium. Male sternite 5 with SE IE IS. SEE ee Se eS ers 17 Notopleuron brown posteriorly (females sometimes with most of notopleuron and postpronotum brown) and scutum with one pair of wide brown postsutural stripes (Fig. 15). Acrostichal bristle absent. Length 2.4-2.9 mm. Female abdomen with wide brown stripe on tergites 2-6. Eastern North America .............csceeesseeeeseeeeeeeeeeees en en eo S. setipes Melander & Argo (in part) Scutum yellow with dark brown quadrate spot on notopleuron (sometimes faded in males and teneral specimens) (Fig. 16). Acrostichal bristle present. Length 3.9- 4.9 mm. Female abdominal pattern as follows: tergite 2 sometimes brown along posterior margin, tergites 3 and 4 with wide central stripe, tergite 5 with narrow central stripe, and tergite 6 dark brown. Eastern and central North America ............. ee ta sa radacarescesaonseccesccansncs S. lachnosternum Melander & Argo
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FIGURES 1-9. 1—Sobarocephala quadrimaculata Soos; male (left) and female abdomen (right). 2—S. cruciger Sabrosky & Steyskal; male (left) and female abdomen (right). 3—S. flaviseta (Johnson); female. 4—S. latifacies Sabrosky & Steyskal; male (left) and female abdomen (right). 5—S. atrifacies Sabrosky & Steyskal; female. 6—S. affinis (Johnson); male (left) and female abdomen (right). 7—S. muesebecki Sabrosky & Steyskal; male (left) and female abdomen (right). 8—S. flava Melander & Argo; male (left) and female (right; arista not illustrated). 9-—S. atricornis Sabrosky & Steyskal; male (left) and female (right; head not illustrated).
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FIGURES 10-17. 10—Sobarocephala latifrons (Loew); male (left) and female (right; head not illustrated). 11—S. wirthi spec. nov.; male (left) and female abdomen (right). 12—S. pengellyi spec. nov.; male (left) and female abdomen (right). 13—S. interrupta Sabrosky & Steyskal; male (left) and female abdomen (right). 14—S. dreisbachi Sabrosky & Steyskal; male (left) and female abdomen (right). 15—S. setipes Melander & Argo; male (left) and female (right; head not illustrated). 16—S. Jachnosternum Melander & Argo; male (left) and female abdomen (right). 17—S. texensis Sabrosky & Steyskal; male.
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Species Descriptions Sobarocephala affinis (Johnson, 1913) (Figs. 6, 39-41; Map 1)
Chaetoclusia affinis Johnson, 1913: 101. Melander & Argo, 1924: 9. Sobarocephala testacea Soos, 1964: 449. Sabrosky & Steyskal, 1974: 378. Sobarocephala affinis, Woodley, 1984: 120.
Redescription (Fig. 6)
Male. Body length 2.8-4.4 mm. Bristles yellow. Two dorsocentral bristles with one minute bristle in front of anterior dorsocentral. Acrostichal bristle absent. Two pairs of well-developed lateral scutellar bristles. Arista densely plumose. Sides of frons parallel. Body yellow except as follows: fore tibia usually light brown to brown; fore tarsi dark brown; coxae, base of femora, posterior half of katepisternum, parafacial and gena white; first flagellomere with dark spot around base of arista; occiput occasionally pilose. Male from Alabama with one pair of small spots behind suture, and lateral margin of postpronotum (and part of notopleuron) brown. M__,, ratio 3.9-4.6. Wing clear. Cell bm open. Face flat.
Female. Externally similar to male except as follows: abdomen with median stripe on tergites (2) 3-4; tergites 5-7 and anterior half of tergite 8 brown.
Male terminalia. (Figs. 39-41) Similar to S. muesebecki, except pregonite ovate and hypandrium with four minute distal bristles.
Distribution: United States: AL, FL,GA, MA, NC, NJ, NY, VA (Map 1).
Holotype. UNITED STATES, VA: Cape Henry, 26 June 1939, A. L. Melander, 3, USNM.
Additional material examined. UNITED STATES, AL: Kushla, 22 July 1914, A. H. Sturtevant, 4, USNM; Baldwin Co., Bon Secour Nat. Wildlife Refuge, 5-7 May 1994, S. A. Marshall, 4, DEBU; Bon Secour (site 4), Malaise, 19 October 2004, 30.30N, 88.74W, E. Benton, 2, DEBU. FL: Alachua Co., Gainesville, Austin Cary Forest, insect flight trap, CO, baited, G. B. Fairchild, 6 August 1976, 23, 22, USNM; 9 August 1976, 3, 9, USNM;; 13 August 1976, 29, USNM; 20 August 1976, 4, 29, USNM; 17 September 1976, ©, USNM; 30 July 1976, ¢, USNM; 26 July 1976, 2, USNM; 16 August 1976, 9, USNM; Gainesville, Pine Hill Estates, H. V. Weems Jr., Malaise trap, 26 September 1973, 2, USNM; 3 October 1973, 2, USNM; Gainesville, Pierce’s homestead, 8 April 1976, W. H. Pierce, flight intercept trap, 2, USNM; Mobile Co., Camp Scoutshire (site 24), Malaise trap, 14 December 2004, 31.05N, 88.18W, E. Benton, 9, DEBU. GA: Liberty Co., St. Catherine’s Isl., 18-21 September 1972, F. C. & B. J. Thompson, 3, AMNH; 24-28 April 1972, Thomas & Picchi, 9, AMNH. MA: Concord, 19 July 1961, W. W. Wirth, marsh, 3, USNM. NC: Wake Co., 7 air mi. SW of Raleigh off rd., Malaise trap, C. S. Parron, 21 September 1985, 3, NCSU; 14 August 1985, 3, NCSU; 2 July 1985, 9, NCSU; 30 August 1985, 9, NCSU. NY: Kalfleisch, 15 August 1962, R. S. Huntington, 2, USNM.
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MAP 1. Distribution of Sobarocephala affinis (Johnson) (circle) and S. muesebecki Sabrosky & Steyskal (dot).
Comments. While the sister-species Sobarocephala muesebecki and S. affinis are both largely restricted to the eastern and southeastern coastal United States, the range of S. muesebecki extends further west into Texas and Pennsylvania. Males of S. muesebecki and S. affinis are easy to separate using the colour of the epandrium, but females are much more difficult to distinguish: the bristles of S. muesebecki are darker and the anterior margin of the frons is light yellow to white, but these characters are often difficult to see in poorly preserved specimens.
Sobarocephala atricornis Sabrosky & Steyskal, 1974 (Figs. 9, 30-32; Map 2) Sobarocephala atricornis Sabrosky & Steyskal, 1974: 382.
Redescription (Fig. 9).
Male. Body length 3.0-3.3 mm. Bristles dark brown. Two dorsocentral bristles. Acrostichal bristle present. Two pairs of well-developed lateral scutellar bristles. Arista sparsely plumose. Sides of frons parallel.. Notum yellow, usually with light to dark spot on and (usually) behind notopleuron (pigment sometimes indistinct). Pleuron and legs light yellow to white. Head largely yellow, with ocellar tubercle brown and ventral half of first
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flagellomere black; parafacial and gena white and silvery tomentose. Abdomen yellow. M,,, ratio 4.4. Wing clear or with cloud around apex of R,,, (sometimes filling first radial cell). Cell bm open. Face flat.
Female. As described for male except as follows: bristles black; scutum yellow with lateral margin dark brown (sometimes faded) (one female from Ontario (DEBU) with scutum yellow excluding brownish lateral margin on notopleuron); scutellum sometimes with thin brown border along lateral margin; laterotergites brown, sometimes yellow below scutellum; face white; fore tarsi sometimes light brown; tergites 2-4 sometimes with small spot on posterior margin; tergite 6 and basal margin of tergite 7 brown; lateral stripe present along length of abdomen (not visible dorsally).
Male terminalia. (Figs. 30-32) Sternite 5 evenly setose. Sclerites of annulus well-developed. Width of epandrium 4/5 height and length 3/5 height. Surstylus 2/3 height of epandrium, slightly wider on distal half; tubercle-like bristles along length of posterior margin, but more concentrated apically. Cerci projecting and rounded; bristles short with several slightly longer central bristles. Ventral lobe of hypandrium long with three minute distal bristles; arm short and truncate. Phallapodeme well-developed. Pregonite ovate, membranous, and possibly fused to hypandrium. Postgonite and basiphallus small. Epiphallus long, projecting, and perpendicular to basiphallus. Distiphallus half length of phallapodeme, and lateral lobe thin, hooked, spinulose distally, and without thumb.
243
Distribution. Canada: ON, PQ. United States: KS, MA, MI, NY (Map 2). Holotype. CANADA, ON: Maynooth, 8 July 1965, J. F. McAlpine, 3, CNCI.
Allotype. UNITED STATES, MI: Claire Co., 23-28 July 1959, R. R. Dreisbach, 9, USNM.
Paratypes examined. CANADA, ON: Ottawa, J. R. Vockeroth, 12 July 1959, 29, CNCI; 9 July 1962, 2, CNCI. PQ: Abbotsford, 23 June 1937, G. Shewell, 2, CNCI. UNITED STATES, MI: Monroe Co., J. Truchan, ex. Malaise trap, 1 August 1965, ¢, USNM; 1 July 1965, 2, USNM.
Additional material examined. CANADA, ON: Innisville, 12 July 1963, W. R. M. Mason, 2, CNCI; White R., July 1980, S. A. Marshall, 2, DEBU; Five Points, 25-30 July 1983, Malaise trap, J. Thompson, 9, DEBU; Wellington Co., University of Guelph Arboretum, ex. dung, O. Lonsdale, 9 August 2004, 2, DEBU [in alcohol]; 19 July 2005, 6, DEBU [in alcohol]; University of Guelph Arboretum, B. Brown, unbaited pitfalls, 22 May—8 July 1983, 29, DEBU; mushroom baited pitfalls, 19 July 1983, 22, DEBU; Guelph, 11 July 1981, J. Ernst, 2, DEBU; Fergus, Malaise trap, S. A. Marshall, 16 July 1990, 22, DEBU; 9 July 1990, 2, DEBU; 10 July 1990, 2, DEBU; Rondeau, 25 July 1981, S. A. Marshall, dung trap, 2, DEBU; Lambton Co., Port Franks, Watson Property, pans, J. Skevington, 15-18 July 1996, 52, DEBU; 31 July—6 August 1996, 2, DEBU; 27 June—2 July 1996, °, DEBU; 18-22 July 1996, 2, DEBU; Sault Ste. Marie, Ft. Creek Cons. Area, 8 July 1998, K. N. Barber, sweeps, Jmpatiens, sedge, fern, 46°32.5’N, 84°20.8’W, 3, DEBU; Sault Ste. Marie, Birchwood Park, 27 July 1986, K. N. Barber, mixed forest, 2, DEBU; Essex Co.,
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Point Pelee N. P., Visitor Centre, Malaise trap and pans, 6-11 August 2000, O. Lonsdale, 2°, DEBU; Ottawa, J. R. Vockeroth, damp, second-growth Acer-Betula wood, 16 July 1989, 3, CNCI; 4 July 1991, 9, CNCI; 7 mi E Griffith, 11 July 1990, B. E. Cooper, 4, CNCI:; McDonald Island, St. Lawrence Isl. N. P., A. Carter, 15 July 1976, 2, CNCI; Ottawa, 13 August 1974, J. R. Vockeroth, 2, CNCI. UNITED STATES, KS: Riley Co., Manhattan, 8 July 1966, Malaise trap, G. F. Hevel, 2, USNM. NY: Corlear Bay, L. Champlain, June 1939, R. C. Shannon, 3, USNM; Ulster Co., Cherrytown, 4 mi NNW Kerhonkson, 15-30 July 1971, P. & B. Wygodzinsky, 9, AMNH.
Comments. Sobarocephala atricornis can be readily diagnosed by a ventrally black first flagellomere (hence its name). If the antennae are missing, females can be identified by their characteristic notal and abdominal patterns. Male S. atricornis are more difficult to identify because, like many other North American Sobarocephala, they are weakly pigmented and have dark bristles, however they can be distinguished by their clear wings and brown supra- alar spots.
MAP 2. Distribution of Sobarocephala atricornis Sabrosky & Steyskal.
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Sobarocephala atrifacies Sabrosky & Steyskal, 1974 (Fig. 5; Map 3) Sobarocephala atrifacies Sabrosky & Steyskal, 1974: 379.
Redescription (Fig. 5) |
Female. Body length 3.8-4.4 mm. Bristles light brown. Two dorsocentral bristles. Acrostichal bristle present. Two pairs of well-developed lateral scutellar bristles. Arista pubescent. Sides of frons converging posteriorly. Thorax mostly yellow; notopleuron, postpronotum, scutellum, laterotergites beside scutellum and posterior notal spots white; one pair of dark brown supra-alar spots and one large posteromedial spot; if face yellow, anterior margin of scutum brown, including inner half of postpronotum. Coxae white. Legs mostly yellow, with tarsi dirty white to light brown, and fore tarsomeres 2-5 and distal 1/3 of tarsomere | brown. Head predominantly yellow, with parafacial and gena white, face black (yellow in some Dominican Republic specimens), and ocellar tubercle brown. Abdomen yellow with tergites 3-6 dark brown (tergites 5 and 3 yellow on anterior corner). Cerci brown. M_,, ratio 3.7, Wing clear. Cell bm closed. Face flat.
Male. Unknown.
Distribution. Florida (Map 3), Dominican Republic.
MAP 3. North American distribution of Sobarocephala atrifacies Sabrosky & Steyskal (diamond) and S. guadrimaculata Sos (circle).
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Holotype. UNITED STATES, FL: Pinellas Co., High Point, 17 November 1971, K. Hickman, 2, USNM.
Paratypes. UNITED STATES, FL: Dade Co., Miami, 3 November 1971, G. R. Searis, McPhail trap in Loquat tree, 62, USNM.
Other material examined. DOMINICAN REPUBLIC. Pedernales, 26 km N Cabo Rojo, 18°06’N, 71°38’W, 730 m, 13-25 July 1990, L. Masner, J. Rawlins & C. Young, wet deciduous forest, intercept trap, 22, CMNH; RD-095 Rodeo, ~0.5 km E Presa de Blanco, Bonao, Monsenor Nouel Prov., 20 March 2003, D. Perez, R. Bastardo, B. Hierro, 2, USNM.
Comments. The paratypes from Florida were collected on grapefruit and guava trees (Sabrosky and Steyskal 1974).
Sobarocephala cruciger Sabrosky & Steyskel, 1974 (Figs. 2, 24-26; Map 4) Sobarocephala cruciger Sabrosky & Steyskal, 1974: 376.
Redescription (Fig. 2)
Male. Body length 3.5-5.4 mm. Bristles yellow. Two dorsocentral bristles. Acrostichal bristle absent. Two pairs of well-developed lateral scutellar bristles. Ocellar bristle absent. Arista densely plumose. Sides of frons converging posteriorly. Scutum mostly yellow; lateral margin with one pair of brown spots behind transverse suture and one pair of spots abutting scutellum; postpronotum light yellow. Scutellum brown laterally. Anatergite brown below scutellum. Pleuron and legs yellow with fore tarsi brown. Coxae white. Head mostly yellow, with gena and parafacial white, and first flagellomere brown on dorsal half; gena silvery tomentose. Tergites 1-5 with central brown stripe; posterior half of tergite 4 brown; tergite 6 with small central spot; surstylus white; remainder of abdomen yellow. M,,, ratio 4.0-4.8. Wing clouded on distal 1/3. Cell bm open. Face convex on dorsal half below antennal bases.
Female. Externally as described for male except anatergite and tergite 6 with brown medial stripe, tergite 7 light brown, and cerci brown.
Male terminalia. (Figs. 24-26) Sternite 5 evenly setose. Annulus reduced to thin band ventrally. Width of epandrium 2/3 height and length half height. Surstylus short, acutely triangular, and curved inwards; tubercle-like bristles terminal only. Cerci projecting and bifid. Hypandrial arm longer and thicker than ventral lobe, projecting at 90° basally (arcuate medially); ventral lobe with one minute and two long distal bristles. Pregonite long and thin with one distal bristle. Postgonite and basiphallus small. Epiphallus thin and weakly sclerotized. Distiphallus short and bent medially; lateral lobe truncate and minutely toothed distally with thumb half length of lobe.
Distribution. Florida (Map 4).
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Holotype. UNITED STATES, FL: Manatee Co., Cedar Hammock, 16 September 1966, D. C. Chancey, Steiner trap, ¢, USNM.
Allotype. UNITED STATES, FL: Dade Co., Miami Springs, 3 January 1972, G. R. Searls, 9, USNM. | :
Paratypes examined. UNITED STATES, FL: Sarasota, McPhail trap, C. K. Hickman, 13 March 1972, 34, USNM; Bradenton, Steiner trap, D. C. Chancey, 10 September 1965, 2, USNM; 29 July 1966, 2, USNM; Volusia Co., S. Dayton, McPhail trap, J. N. Pott, 18 October 1966, 3, 22, USNM; 5 October 1965, 2, USNM; 10 March 1972, 2, USNM; 14 May 1965, 3, USNM; grapefruit tree, 29 October 1971, 9, USNM; 12 October 1972, 9, USNM; 10 March 1972, 29, USNM; Allendale, McPhail trap, 3 November 1966, 2°, USNM; Volusia Co., Harbour Oakes, McPhail trap, J. N. Pott, 28 October 1966, 29, USNM; 3 November 1966, 2, USNM; Dade Co., Miami Springs, G. R. Searls, 3 January 1972, 39,
MAP 4. Map of Florida showing distribution of Sobarocephala cruciger Sabrosky & Steyskal.
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USNM; 22 October 1971, 9, USNM; C. F. Dawling Jr., McPhail trap, 20 February 1968, 2, USNM; Pinellas Co., High Point, McPhail trap, grapefruit tree, K. Hickman, 17 November 1971, 2, USNM; Sarasota Co., Osprey, McPhail trap, S. V. Hiatt, 24 October 1966, °, USNM; Hillsborough Co., Tampa, McPhail trap, T. J. Flavoroso, 1 August 1968, 9°, USNM; 9 October 1967, 2, USNM; 20 October 1969, 29, USNM; Manatee Co., Palmetto, McPhail trap, 23 September 1966, C. J. Bickner, 9, USNM; J. R. McFarlin, McPhail trap, 8 August 1971, 22, USNM; McFarlin & Bickner, 9 March 1972, 29, USNM.
Additional material examined. UNITED STATES, FL: Gainesville, Doyle Conner Bldg., Malaise trap, 23 September 1973, H. V. Weems Jr., 9, USNM; Alachua Co., Pierce’s Homestead, Malaise trap, W. H. Pierce, 13 October 1973, 2, USNM.
Sobarocephala dreisbachi Sabrosky & Steyskal, 1974 (Figs. 14, 54-56, Map 5) Sobarocephala dreisbachi Sabrosky & Steyskal, 1974: 382.
Redescription (Fig. 14)
Male. Body length 3.3 mm. Bristles brown to dark brown. Two dorsocentral bristles with one minute bristle in front of anterior dorsocentral. Acrostichal bristle absent. Two pairs of well-developed lateral scutellar bristles. Arista sparsely plumose. Sides of frons parallel. Scutum yellow, with notopleuron brown to light brown and postsutural stripes brown (broadly connected at base). Scutellum and laterotergites beside scutellum brown (sometimes also with large spot below scutellum). Pleuron and legs light yellowish- white with fore tarsi brown and anepisternum sometimes yellow. Head yellow with orange tint (sometimes darker laterally), with ocellar tubercle brown and gena light yellow and pilose. Abdomen yellow. M.,,, ratio 4.8. Wing lightly clouded around distal half of R,,.. Cell bm open. Face flat.
Female. Similar to male except tergites 3-6 and posterior margin of tergite 2 brown; posterior corners of tergite 5 yellow. Females from Québec with notopleuron yellow or light brown and fore tibia brown (tip lighter).
Male terminalia. (Figs. 54—56) Fifth sternite with posteromedial comb. Sclerites of annulus well-developed. Epandrium slightly wider than high; length approximately 3/5 width. Surstylus as high as epandrium, with outline subtriangular and broadly rounded; tubercle-like bristles absent. Cerci sunken below distal margin of epandrium, and all bristles short. Hypandrial arm atrophied; ventral lobe with one minute distal and two long medial bristles. Phallapodeme carinate and atrophied distally. Postgonite small. Pregonite membranous with one distal setula. Basiphallus well-developed. Epiphallus long, thin, and strongly projecting from basiphallus. Distiphallus 2/5 length of phallapodeme; thumb of lateral lobe short and truncate.
Distribution. Canada: ON, PQ. United States: MI, NC, TN, TX (Map 5).
Holotype. UNITED STATES, MI: Ogemaw Co., 1-2 August 1959, R. R. Dreisbach, °, USNM.
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Paratypes. CANADA, ON: Ottawa, 26 July 1959, J. R. Vockeroth, ¢, CNCI. UNITED STATES, NC: Macon Co., Wayah Gap, 3500 ft, 10 August 1957, J. G. Chillcott, 4, CNCI.
Additional material examined. CANADA, ON: Algoma Dist., Hilton Twp., Hilton Beach, Malaise trap at edge of field and hardwood forest, 20 August 1992, J. E. Swann, 2, DEBU; Bruce Co., Inverhuron P. P., back dunes, 44°17°50”N, 81°35’27”W, 20 July—20 August 2003, Malaise, S. A. Marshall, 9, DEBU. PQ: Duncan Lk., nr. Rupert, J. F. McAlpine, 6 August 1970, 9, CNCI; 21 July 1971, 2, CNCI; 16 July 1971, 29, CNCI; 27 July 1971, 2, CNCI. UNITED STATES, TN: Union Co., 9 mi SE La Folette, 9 June 1973, A. O. Lavallee, 9, EMUS. TX: Jasper Co., Bouton Lake Rec. Area, Angelina Nat’! Forest 11 miles SE of Zavaila, off Texas Hwy 63, 15 May 1993, I. Yarom, 4, TAUI.
Comments. The genitalia of Sobarocephala dreisbachi are similar to those of S. setipes and S. lachnosternum, in that there is a comb of bristles on the fifth sternite and the cerci are sunken, but the surstylus is much longer.
MAP 5. Distribution of Sobarocephala dreisbachi Sabrosky & Steyskal.
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Sobarocephala flava Melander & Argo, 1924 (Figs. 8, 33-35, Map 6)
Sobarocephala flava Melander & Argo, 1924: 40. Sabrosky & Steyskal, 1974: 383. Sobarocephala populi Steyskal, 1951: 129.
Redescription (Fig. 8)
Male. Body length 2.8-5.3 mm. Bristles dark brown to black. Two dorsocentral bristles plus one minute bristle in front of anterior dorsocentral. Acrostichal bristle present. Lateral scutellar bristles weak to well-developed. Arista sparsely plumose. Sides of frons parallel. Notum yellow with notopleuron light brown to brown and lateral margin of scutum behind notopleuron sometimes brown. Pleuron, coxae, tibiae, and basal 1/3 of femora white, with fore tarsi light brown or yellow and anepisternum sometimes light yellow. Head yellow with ocellar tubercle brown and sclerites below antenna white; gena pilose. Abdomen yellow. M,,, ratio 4.0. Wing dusky along R,,, and costa (more so on distal 4); occasionally clear. Cell bm open. Face flat.
Female. Similar to male except as follows: notopleuron rarely yellow (yellow in holotype); face, parafacial, and gena sometimes light yellow to white; first flagellomere with light infuscation along anterior margin; tergites 6-7 brown (tergite 7 usually only pigmented anterolaterally); specimens from Mississippi with tergites 2-5 (sometimes also tergites 7 and 8) brown laterally; female from Texas mostly yellow, with notopleuron light brown, and tergites 6 and 7 dark brown anteriorly and laterally. Female from Rondeau Provincial Park (Ontario) with entire lateral margin of scutum and scutellum dark brown.
Male terminalia. (Figs. 33—35) Externally similar to S. atricornis (Figs. 30, 31) except apex of surstylus slightly broader. Internally similar to S. atricornis (Fig. 32) except as follows: pregonite clavate with five bristles along length; only two hypandrial bristles usually present; lateral lobe strongly bent medially with thumb short, pointed, and projecting at 90°.
Distribution. Canada: ON. United States: FL, GA, IL, IN, LA, MD, MI, MO, MS, NC, NY, PA, TX, UT, VA (Map 6).
Holotype. UNITED STATES, VA: Fairfax Co., Dead Run, ISS 15 April [year unknown], 2, USNM.
Paratypes examined. UNITED STATES, LA: Opelousas, April 1917, 2, USNM. MI: Plummer Isl., 29 June 1913, 9, USNM. VA: Dead Run, R. C. Shannon, 15 July 1915, 39, USNM; ex. Maple log, 13 March 1915, ISS 19 April 1915, 2, USNM; 11 July 1915, 9, USNM; Falls Church, N. Banks, June 1920, 2, USNM; 4 July 1913, F. Knab, 29, USNM.
Additional material examined. CANADA, ON: 7 mi E Griffith, 1 July 1990, B. E. Cooper, 22, CNCI; Hamilton, 13-19 July 1980, Malaise trap, M. Sandborne, 22, DEBU; Burlington, Royal Bot. Gardens, 16 July 1997, K. N. Barber, sweeps, trail-side Poa, Agrostis, Phleum, Festuca, Juncus, 43°17.5’N, 79°52.4’W, 2°, DEBU; Burlington, Bronte Crk. Prov. Pk., 17-20 August 1983, Brown & Marshall, Malaise trap, 9, DEBU; Dundas, May 1980, E. A. Menard, reared, larva in dead Elm, 2, DEBU; Algonquin Prov. Pk., Swan Lk. Res. Sta.,
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reared from rothole in dead Maple tree, emerged 30 May 1995, D. C. Caloren, 9, DEBU; Five Points, 29-30 July 1983, J. G. Thompson, 2, DEBU; Fergus (yard), 9 August 1985, S. A. Marshall, floodplain, Malaise trap, 2, DEBU; Fergus, Malaise trap, S. A. Marshall 8-14 June 1982, 2°, DEBU; 16 July 1990, 29, DEBU; Port Credit, 15 July 1918, M. C. Van Duzee, 2, CASC; Bells Corners, E. C. Becker, 31. May 1954, reared from under Beech, 2, USNM; 21 June 1954, reared ex. puparia on rotten log, ¢, USNM; Bells Corners, reared ex. puparia on rotten log, 21 June 1954, E. C. Becker, 54, 62, CNCI; Ottawa, J. R. Vockeroth, damp second-growth Acer-Betula wood, 4 July 1991, 32, CNCI; 2 July 1992, 22, CNCI; 11 July 1989, 2°, CNCI; 13 August 1991, 22, CNCI; 6 July 1991, 29, CNCI; 28 July 1991, 29, CNCI; 9 August 1992, 9, CNCI; McDonald Isl., St. Lawrence Isl. N. P., A. Carter, Malaise trap, 19 July 1976, 2, CNCI; 20 July 1976, 2, CNCI; 5 August 1976, 29, CNCI; Metcalfe, B. E. Cooper, 24 July 1984, 22, CNCI; 22 July 1984, 2, CNCI; 4 August 1984, 2°, CNCI; 1 August 1984, 9, CNCI; Essex Co., Point Pelee, 18 July 1978, J. M. Cumming, 2, DEBU; Windsor, Ojibway Prairie, S. Paiero unburnt forest, yellow pans, 3-6 July 2001, 2, DEBU; 31 July—3 August 2001, 29, DEBU; burnt savannah, yellow pans, 25-29 June 2001, 9, DEBU; Kent Co., Rondeau P. P., Spicebush Trail, 42°18°09"N, 81°51°06”W, Carolinian forest, Malaise, 16-29 July 2003, S. A. Marshall, 22, DEBU. UNITED STATES, FL: Gainesville, Pine Hill Estates, 4 October 1973, H. V. Weems Jr., Malaise trap, 9, USNM. GA: Forsyth, 2 June 1970, 22, CNCI. IL: Champaign Co., Brownfield Woods, 2 mi NE Urbana, 29 June 1976, C. T. Maier, 2, EMUS; Mason Co., Sandridge St. Forest, 4 mi NW Forrest City, 23 June 1976, C. T. Maier, 9, EMUS; Tazewell Co., 3 mi N Macking along Panther Cr., Webb & Marlin, 10 June 1969, 4¢, EMUS; Carlinville, 9 May 1952, M. R. Wheeler, 4, AMNH. IN: La Fayette, 8 July 1916, ¢, USNM; July 1912, ¢, USNM; J. M. Aldrich, 3 July 1915, 9, USNM; August 1914, ¢, USNM; July 1921, ¢, USNM. MD: Plummers Isl., 11 July 1915, R. C. Shannon, 2, USNM; Colesville, W. W. Wirth, 11 July 1974, 42, USNM; 4 July 1976, 2, USNM; 18 June 1977, 22, USNM; 14 June 1977, 9, USNM; 28 July 1976, 2, USNM; Montg’y Co., Colesville, W. W. Wirth, Malaise trap, 30 June 1977, ¢, USNM; 26 June 1977, 4,22, USNM; Montg’y Co., Dickerson, G. A. Foster, 14 July 1974, 24, USNM; Montg’y Co., Rockville, W. W. Wirth, Malaise trap, 4 September 1977, 2, USNM; 3 August 1979, ¢, USNM; 26 July 1979, 9, USNM; Montg’y Co., Chevy Chase “woodend”, 3 August 1974, G. F. Hevel, 2, USNM; Prince Georges Co., Patuxent Wildlife Res. Centre, W. W. Wirth, 8 July 1978, 2, USNM; J. F. Reinert 31 July 1977, , USNM;; Glen Echo, J. R. Malloch, 23 July 1921, 9, USNM; Laurel, 11 June 1965, marsh edge, 32, CNCI; Prince George Co., Beltsville, 15 July 1979, A. Freidberg, 9, TAUI. MI: Grosse Ile, 30 June 1949, G. Steyskal, 9, USNM; Berrien Co., St. Joseph, D. D. Wilder, 14 July 1971, 2, USNM; Monroe Co., 20 July 1965, J. Truchan, ex. Malaise trap, ¢, USNM; Wayne Co., Grosse Isle, G. C. Steyskal, 1 July 1949, 9, USNM; 14 June 1949, 2, USNM; 4 June 1962, 29, USNM; 4 August 1956, 22, USNM; 3 August 1951, 22, USNM; 7 August 1951, 29, USNM; 8 August 1956, 2, USNM; 22 June 1949, 29, USNM; 24 July 1951, 69, USNM. MO: Columbia, Malaise trap, 24 June 1967, 4pm—7am, F. D. Parker, 2, USNM. MS: Forrest Co., 6 mi W Wiggins, Sweet Bay Bog, dung trap, 5-8 May 1994, sphagnum, S. A. Marshall, 32, DEBU. NC: Columbus Co., Lk. Waccamaw, 6 July 1985, W. Stein & A. Gerberich, blacklight in Oak & Pine scrub sand barriers nr lake, 9, USNM; Bladen Co., Singletary Lk. St. Pk., 34°35’0N, 78°27°30W, Malaise, 19-22 May 2003, Marshall & Paiero, 2, DEBU: Wake Co., 7 air mi SW of Raleigh off rd., 1 September 1985, C. S.
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MAP 6. Distribution of Sobarocephala flava Melander & Argo.
Parron, Malaise trap, 29, NCSU; Raleigh, late July, F. Sherman, 2, NCSU. NY: Brainard, Rensselaer Co., 11-22 July 1966, P. & B. Wygosinsky, 9, AMNH. PA: State College, Centre Co., 4 July 1972, D. J. Shetlar, 9, CASC. TX: Kerrville, 23 April 1959, Becker & Howden, 2, CNCI. UT: Garden City, 15 July 1951, F. C. Harmaton, 9, USNM. VA: Alexandria, 24 June 1951, W. W. Wirth, 3, USNM; St. Falls, 9 July 1926, A. L. Melander, 2, USNM; Falls Church, Holmes Run, 6 August 1960, light trap, 4, USNM; Dead Run, 28 July 1915, 2, USNM; 29 June 1915, 9, USNM; R. C. Shannon, 15 July 1915, 29, USNM; 28 July 1915, 2, USNM; Scott Run, July 1954, M. R. Wheeler 3, 2, USNM:; Marina National Airport, 28 August 1994, A. Freidberg, 2, TAUI.
Comments. Sobarocephala flava is a commonly collected species throughout much of eastern North America; specimens have also been collected in central Texas and Utah.
Sobarocephala flaviseta (Johnson, 1913) (Figs. 3, 18—20; Map 7)
Heteromeringia flaviseta Johnson, 1913: 99. Heteromeringia convergens Malloch, 1922: 50. Sobarocephala flaviseta, Sabrosky & Steyskal 1974: 380.
Redescription (Fig. 3) Male. Body length 3.0-5.0 mm. Bristles brown. Two dorsocentral bristles plus one minute bristle in front of anterior dorsocentral. Acrostichal bristle present. Two pairs of 83
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well-developed lateral scutellar bristles. Arista sparsely plumose. Sides of frons converging posteriorly. Scutum yellow with postpronotum and notopleuron white (at least centrally). Pleuron light yellow. Legs light yellow with coxae and base of femora white. Head yellow to white, with ocellar tubercle brown, and with parafacial, gena, sides of face, and scape white. Abdomen yellow. M__,, ratio 3.1-3.3. Wing clear (sometimes clouded distally in first radial cell). Cell bm closed. Face, parafacial, and anterior margin of frons uniformly bulging.
Female. As described for male except as follows: tergite 3 with faded central stripe; tergites 4 and 5 light brown with anterior-lateral corners widely yellow; abdominal pattern sometimes reduced to central stripe on tergite 5. Cercus light brown.
Male terminalia (Figs. 18—20). Sternite 5 evenly setose. Sclerites of annulus well- developed. Epandrium as wide as high and length 3/4 height. Surstylus short, acute, and curved inwards; tubercle-like bristles distal only. Cerci projecting and slightly emarginate with one pair of slightly longer central bristles. Hypandrial arm stout and projecting at 90°; ventral lobe thin, slightly longer than arm, and with one minute and two long distal bristles. Phallapodeme well-developed and slightly sinuate. Pregonite narrow basally, wide distally and with five apical bristles. Postgonite absent. Distiphallus % length of phallapodeme and curved at base; lateral lobe wide, truncate, minutely serrate, and with long thumb.
MAP 7. Distribution of Sobarocephala flaviseta (Johnson).
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Distribution. NB, ON, PQ and eastern U.S. to BC, AB, ID, CO, and TX (Map 7). Holotype. UNITED STATES, NJ: New Brunswick, 28 May, J. B. Smith, 2, MCZC.
Paratypes examined. UNITED STATES, IN: Lafayette, July 1927, 9, USNM. VA: Chainbridge, 20 August 22, J. R. Malloch, 9, USNM. NJ: same collection as holotype, °, MCZC.
Additional material examined. 192, 1982 [AMNH, CNCI, DEBU, TAMU, TAUI, USNM].
Comments. Sobarocephala latifrons and S. flaviseta are two of the most commonly encountered Sobarocephaila in northeastern North America, although neither is as frequently collected as species of Clusia or Clusiodes. Sobarocephala flaviseta is common in eastern North America, and it has been occasionally collected in Texas and the northwest (Sabrosky and Steyskal 1974). ;
The apparent female-biased sex ratio of over 10:1 is unusual, although females of other species are generally more often collected than males. We suspect that this apparent female-biased sex ratio is a collecting artifact, and is more likely to reflect the clumping of males at mating sites than a population-level ratio. Too few reared specimens are available to assess actual sex ratios of any clusiid species.
Sobarocephala interrupta Sabrosky & Steyskal, 1974 (Figs. 13, 51-53; Map 8) Sobarocephala interrupta Sabrosky & Steyskal, 1974: 384.
Redescription (Fig. 13)
Male. Body length 4.5 mm. Bristles light brown. Two dorsocentral bristles. Acrostichal bristle present. Arista sparsely plumose. Anterior lateral scutellar bristle minute or absent. Sides of frons parallel. Notum yellow, with notopleuron brown, scutum with one pair of basal spots (often serrate), and lateral margin of scutellum brown. Pleuron and legs light yellow with fore tarsi brown; Mississippi specimens with pleuron and basal half of femora white. Head yellow, with gena and parafacial dirty white to white, ventral margin of face sometimes orange, and ocellar tubercle brown; gena pilose. Abdomen yellow with tergites 2-5 brown (excluding anterior corners). M,_, ratio 3.6-5.5. Wing clear. Cell bm open.
Female. Similar to male except tergites 3 and 4 brown and tergite 6 with posterior margin light brown centrally. Scutellum entirely yellow in South Carolina paratype.
Male terminalia. (Figs. 51-53) Sternite 5 evenly setose. Sclerites of annulus well-developed. Epandrium as high as wide and length half height. Surstylus bifid with anterior lobe well bristled and posterior lobe sparsely setose with small conical tubercle- like bristles; lobes subequal in length. Cerci concave medially, flush with distal margin of epandrium, and with all bristles short. Hypandrial arm very short; ventral lobe with one minute distal and two long medial bristles. Phallapodeme thinned medially, wide and
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truncate distally, and not extending past dorsal margin of hypandrium. Pregonite large and circular with four medial bristles. Postgonite thin and flat with four bristles. Basiphallus well-developed. Epiphallus small and well-sclerotized. Distiphallus nearly half length of phallapodeme; lateral lobe small and finger-like with thumb absent.
Distribution. United States: FL, GA, IL, LA, MS, SC, TX (Map 8).
Holotype. UNITED STATES, FL: Orange Co., Rock Springs, 21 April 1970, W. W. Wirth, 7, USNM.
Allotype. UNITED STATES, FL: Alachua Co., Chantilly Acres, 25 April 1970, W. W. Wirth, Malaise trap, 9, USNM.
Paratypes examined. UNITED STATES, LA: Kilbourne, 3 May 1959, W. W. Wirth, °, USNM. SC: Greenwood, Long Cane Lake, 21 July 1957, W. R. Richards, 9, CNCI.
Additional material examined. UNITED STATES, FL: Gainesville, Doyle Conner Bldg., H. V. Weems Jr., Malaise trap, 16 October 1973, 2, USNM; 8-9 September 1973, 2, USNM; Highlands Hammock St. Pk., H. V. Weems Jr., 15 July 1956, 9, USNM. GA:
MAP 8. Distribution of Sobarocephala interrupta Sabrosky & Steyskal (dot) and S. texensis Sabrosky & Steyskal (circle).
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Liberty Co., St. Catherine’s Isl., 18-21 September 1972, F.C. & B. J. Thompson, 3, AMNH. IL: Champaign Co., Brownfield Woods, 2 mi NE Urbana, 29 June 1976, C. T. Maier, 9, EMUS. MS: Forrest Co., 6 mi W Wiggins, Sweet Bay Bog, 5-8 May 1994, sphagnum, dung trap, S. A. Marshall, ¢ 32, DEBU. SC: Georgetown Co., Hobcaw Barony, Belle Baruch Marine Field Lab, on slime mold, May 2004, S. A. Marshall, 9, DEBU [in alcohol]. TX: Montg. Co., Jones St. Forest, 8 mi S Conroe, 28 June—13 July 1987, Wharton, Steck, Carroll, ¢, TAMU; 1-7 June 1987, Wang, Wharton, Praetorius, Malaise trap, 2, TAMU.
Comments. The medially yellow scutellum, characteristic notal stripes, and bifid surstylus most readily diagnose Sobarocephala interrupta.
Sobarocephala lachnosternum Melander & Argo, 1924 (Figs. 16, 62-64; Map 9)
Sobarocephala lachnosternum Melander & Argo, 1924: 42. Sabrosky & Steyskal, 1974: 382. .
Redescription (Fig. 16)
Male. Body length 3.9-4.9 mm. Bristles dark brown. Two dorsocentral bristles plus one minute bristle in front of anterior dorsocentral. Acrostichal bristle present. Two pairs of well-developed lateral scutellar bristles. Arista sparsely plumose. Sides of frons parallel. Scutum yellow with notopleuron dark brown (notopleuron sometimes faded, often in newly emerged adults). Scutellum and laterotergites yellow, although anatergite sometimes with one pair of light brown stripes lateral to scutellum. Pleuron light yellow to white excluding yellow anepisternum and katepisternum. Coxae white. Legs yellow with fore tarsi brown. Head yellow with parafacial and occiput white, gena white and silvery tomentose, ocellar tubercle brown, and anterior margin of frons with dirty yellow tint continuing onto dorsal margin of face. Abdomen light yellow, occasionally with light brown to brown bands on posterior margin of tergites 3-5 (faded medially). M,,, ratio 3.1- 3.5. Wing clear. Cell bm open. Face flat.
Female. Similar to male except as follows: posterior margin of tergite 2 sometimes brown; tergites 3 and 4 brown with anterior corners yellow; tergite 5 brown centrally and on posterior margin (sometimes reduced to light posterior spot); tergite 6 dark brown; remainder of abdomen yellow; pedicel brown in one female (Maryland).
Male terminalia. (Figs. 62—64) Sternite five with comb of bristles on posteromedial margin. Sclerites of annulus well-developed. Epandrium large and barrel-shaped (as wide and high as pre-genitalic abdominal segments). Surstylus small and broadly rounded; tubercle-like bristles absent. Cerci small, rounded, slightly projecting, and with all bristles short. Ventral lobe of hypandrium elongate and poorly-defined with three minute bristles. Phallapodeme elongate and well-developed. Basiphallus well-developed. Epiphallus small. Postgonite large and ovate with several minute setulae. Pregonite absent. Distiphallus very short and curved; lateral lobe ovate.
Distribution. Canada: ON, PQ, SK. United States: IL, IN, KS, MD, MI, NC, TX, UT, VA (Map 9).
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Holotype. UNITED STATES, VA: Rosslyn, R. C. Shannon, found larva 25 November 1912, 3, USNM.
Paratypes. UNITED STATES, VA: Rosslyn, R. C. Shannon, found larva 25 November 1912, ISS 10 January 1913, 4, USNM; bred specimen coll. 25 November 1912, ISS 26 January 1913, °, USNM; bred specimen coll. 25 November 1912, ISS 24 January 1913, , USNM; ex. rotten log, 2 May 1913, ISS 20 May 1913, 9, USNM; ex. rotten log, 2 May 1913, ISS 17 May 1913, 4, USNM.
Additional material examined. CANADA, ON: Fergus, 10 July 1990, Malaise trap, S. A. Marshall, ‘°°, DEBU; Guelph, 21 July 1978, W. Ralley, °, DEBU; Dundas, 29 June 1981, J. Kircher, °, DEBU; Algonquin Prov. Pk., Swan Lk. Survey, 45°29°15”"N, 78°43°20°W, Malaise trap, hemlock/hardwood, 16-28 July 1993, Larson, Marshall & Barr, ‘°, DEBU; Shakwa Lk., Oulette Twp., 21 July 1996, A. Applejohn, °°, DEBU; Hamilton, 19-28 August 1980, M. Sandborne, Malaise trap, ‘°°, DEBU; Elgin Co., Springwater Cons. Area, 24 June 1996, D. C. Caloren, °, DEBU; Bruce Co., Dunk’s Bay, Malaise trap, S. A. Marshall, 17 July-18 August 1996, 9°, DEBU; Icewater Crk., 12.7 km NNE Searchmont, mil0.5 Whitmore Dam Rd., 24 June 1986, K. N. Barber, mixed forest, ¢, DEBU; [Nippising] Hwy#1l7, 7 km W Mattawa, 12 June—2 July 2004, opaque mini sticky traps, mixed forest, 46°17.3°N, 78°49.0°W, K. N. Barber, YY, DEBU; Windsor, Malaise trap, S. A. Marshall, 7-16 August 1982, Y°, DEBU; Sault Ste. Marie, Bristol Pl. Pk., 2 July 2001, K. N. Barber, sweeps, mostly sedges, 46°30.8’°N, 84°16.6°W, 2°, DEBU; 40 km SSW White River, K. N. Barber, boreal mixedwood, Malaise trap, 48°14.08’N, 85°22.02’W, 7-22 July 2003, °, DEBU; multi-colour sticky trap, 48°14.05’N, 85°21.97’W, 10-23 July 2003, 4,39, DEBU; 16-26 June 2003, 4, DEBU; 48°14.14°N, 85°22.02’W, 26 June—10 July 2003, 7, 6%, DEBU; Ottawa, 24 July 1972, J. R. Vockeroth, damp secondary growth Acer-Betula wood, ©, CNCI; Innisville, 12 July 1963, W. R. M. Mason, Y°, CNCI; 7 mi E Griffith, 5 July 1990, B. E. Cooper , 2, CNCI. PQ: Gatineau, King Mtn., S. A. Marshall; 23 August 1977, ¢, DEBU; Kirk’s Ferry, 23 August 1924, G. S. Walley, 4, CNCI; Duncan Lk. Nr. Rupert, J. F. McAlpine, 20 July 1969, 2, CNCI; 28 July 1971, 2, CNCI; 24 July 1971, 9, CNCI. SK: Beaver Crk. Cons. Area, ~13 km S Saskatoon, 12 July 1999, K. N. Barber, sweeps, mostly grasses under Betula/Populus, 51°58.6°N, 106°43’W, 3°, DEBU. UNITED STATES, IL: Tazewell Co., 3 mi N Mackinaw along Panther Crk., Webb & Marlin, 10 June 1969, 2, EMUS. IN: Posey Co., Harmonie St. Pk., 24-26 June 1998, Wharton et al., ‘°°, DEBU. KS: Pottawatomie Co., 12 mi W Wamego, 24 July 1966, G. F. Hevel, 9, USNM. MD: Montg’y Co., Colesville, W. W. Wirth, Malaise trap, 26 June 1977, , USNM; 15 August 1975, 2, USNM; Colesville, W. W. Wirth, 6 August 1976, 2, USNM; 28 July 1976, 2°, USNM; Malaise trap, 13 August 1977, °, USNM; Montg’y Co., Bethesda, G. C. Steyskal, 9 August 1972, 29, USNM; Laurel, 11 June 1965, marsh edge, ¢, CNCI. ME: Dryden, 7 July 1959, G. H. Heinrich, 9, CNCI. MI: Wayne Co., Grosse Isle, G. C. Steyskal, 8 August 1956, 2, USNM; 15 June 1949, °, USNM; 4 August 1956, 2, USNM; 23 June 1956, 4, USNM; 19 June 1955, 4, USNM; Monroe Co., 26 July 1956, J. Truchan, ex. Malaise trap, 3, USNM. NC: Wake Co., 7 air mi SW of Raleigh off rd., 25 July 1985, C. S. Parron, Malaise trap, 9, NCSU. TX: Salmon, Anderson Co., 6-19 October 1974, H.R. Burke, 2, TAMU. UT: Utah Co., Provo, Malaise trap, 6-9 July 1985, 4, 9, EMUS; 16-23
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et ogi te asm ee
' i ' ! i ! i
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MAP 9. Distribution of Sobarocephala lachnosternum Melander & Argo.
July 1985, 22, EMUS; 20-27 July 1985, 9, EMUS; 7-14 August 1985, 32, EMUS; 14-21 August 1985, 9, EMUS; 6-16 July 1985, 32, EMUS; 18-26 June 1985, 4, 149, EMUS; 25 June—6 July 1985, 112, EMUS. VA: Warsaw, 26 July 1952, W. W. Wirth, 2, USNM.
Comments. While male Sobarocephala lachnosternum superficially resemble other pale North American Sobarocephala, they can be readily identified by a large barrel-shaped epandrium and small surstyli. Females can be identified by their characteristic abdominal pattern.
See comments for Sobarocephala setipes.
Sobarocephala latifacies Sabrosky & Steyskal, 1974 (Figs. 4, 21—23; Map 10) Sobarocephala latifacies Sabrosky & Steyskal, 1974: 380.
Redescription (Fig. 4)
Male. Body length 3.0-5.6 mm. Bristles black. Two dorsocentral bristles plus one minute bristle in front of anterior dorsocentral. Acrostichal bristle present. Two pairs of well-developed lateral scutellar bristles. Arista short-plumose. Sides of frons converging posteriorly. Notum predominantly yellow with notopleuron brown to dark brown; scutum sometimes brown on lateral margin and laterotergites yellow or brown. Pleuron light yellow. Coxae white. Legs yellow with fore tarsi light brown (at least apically). Head yellow with gena and parafacial white, face light yellow and ocellar tubercle brown. Abdomen yellow.
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M, ,, ratio approximately 3.0. Wing clouded in cell R,, around distal 1/5 of R,,., and around R,,,. Cell bm open. Face, parafacial, and anterior margin of frons uniformly bulging.
Female. Scutum as described for male. Abdomen yellow with tergites 6 and 7 brown and tergites 2-5 with small spot on anteromedial margin (tergites 2-5 sometimes with light brown medial stripe and light brown border, or only with posterior margins brown).
Male terminalia. (Figs. 21—23) Sternite 5 evenly setose. Sclerites of annulus well-developed. Epandrium as wide as long and almost as high as wide. Surstylus short, rounded, thick, and slightly pointed apically; tubercle-like bristles along entire length of apical and posterior margins, somewhat encroaching upon posterior surface. Cerci projecting, somewhat emarginate and with one pair of longer central bristles. Hypandrial arm curved, widest distally, as long as ventral lobe and projecting at 60-70° basally; lobe with one minute and two long distal bristles. Phallapodeme long and relatively thin. Pregonite long, tapered at both ends and with five medial bristles. Postgonite minute. Basiphallus large with epiphallus at tip. Distiphallus approximately 2/3 length of phallapodeme and bent at midpoint; lateral lobe well-developed, curved, and toothed distally; thumb well- developed.
Distribution. Canada: NB, ON, PQ. United States: MA, MI, NC, NH, NY, SC, UT (northeastern North America and Utah) (Map 10).
Holotype. UNITED STATES, MI: Grosse Ile, Wayne Co., 29 August 1948, G. Steyskal, 3, USNM.
Paratypes examined. CANADA, ON: Bells Corners, ex. Rotten log, 21 June 1954, E. C. Becker, 2, 2, USNM; Bells Corners, 12 June 1954, E. C. Becker, reared ex. puparia on rotten log, 2¢, 22, CNCI; Burke Falls, 13 July 1926, F. P. Ide, 29, CNCI; nr. Picton, 9 July 1970, J. F. McAlpine, 2, CNCI. PQ: Hull, 10 August 1965, ex. Malaise trap, 2, CNCI; Laniel, 28 July 1933, 2, CNCI; Duncan Lk. Nr. Rupert, 1 August 1969, J. F. McAlpine, 9, CNCI. UNITED STATES. White Mts., Morrison, ¢, USNM.
Additional material examined. CANADA, ON: Tenby Bay, Malaise at lake edge, 19 July 1992, 9, DEBU; Hamilton, 10-13 July 1980, Malaise trap, M. Sandborne, 3’, 32, DEBU; Dundas, E. A. Menard, 27 June 1980, 3, 22, DEBU; 13 June 1980, 4, 9, DEBU; 17 June 1980, d, 2, DEBU; 1 July 1980, 24, DEBU; 23 June 1980, 9, DEBU; 8 July 1980, 3, DEBU; 15 July 1980, 4, DEBU; reared in dead Elm, May 1980, ¢, DEBU; Dornoch, fen, 20 July 1996, S. A. Marshall, 4, DEBU; Port Franks, Watson property nr. lake, pans, 8-12 July 1996, J. Skevington, 9, DEBU; Sault Ste. Marie, Bristol Pl. Pk., sweeps, mostly /mpatiens, Clematis, Rubus, grasses, 46°30.8’N, 84°16.6’W, 11 June 1999, K. N. Barber, 2, DEBU; Fergus, Malaise trap, S. A. Marshall, 5 July 1990, 2, DEBU; 4 August 1990, 2, DEBU; 9 July 1990, 2, DEBU; (yard) flood plain, Malaise trap, 27 June 1985, S. A. Marshall, °, DEBU; Guelph, K. N. Barber, 11 July 1979, 29, DEBU; 9 July 1979, 2, DEBU; pan traps, 5-18 July 1980, 22, DEBU; Brown & Marshall, University of Guelph Arboretum, Malaise head, 28 June4 July 1983, 9, DEBU; Arkell, E. A. Innes, 7 May 1979, 2, DEBU; 7 May 1979, (emerged 22 May 1979), 2, DEBU; 8 May 1979 (emerged 22-30 May 1979), 416, 222, DEBU. UNITED STATES, MD: Colesville, W. W. Wirth, 4 July 1976, 2, USNM; 11
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{ : Jos? : : ; ’ Sas os : 4 \ a y ‘ XS 7A Os WV ¢, at Aine fi 3S Id
MAP 10. Distribution of Sobarocephala latifacies Sabrosky & Steyskal.
July 1974, 3, USNM; 18 June 1977, 3, USNM; Montgomery Co., Carderock Pk., 13 June 1970, L. V. Knutson, 2, USNM; Prince Georges Co., Patuxent Wildlife Res. Centre, W. W. Wirth, 14 July 1978, 9, USNM. MI: Wayne Co., Grosse Ile, 30 June 1949, G. Steyskal, 22, USNM. NC: Cumberland Co., Ft. Bragg, 28 May-3 June 1967, J. D. Birchim, 9, CASC; Wake Co., 7 air mi SW of Raleigh off rd., 30 August 1985, C. S. Parron, Malaise trap, ¢, NCSU. NY: Rensselaer Co., 11-22 July 1966, P. &.B. Wygodzinsky, 2, AMNH. SC: Georgetown Co., Hobcaw Barony, Belle Baruch Marine Field Lab, on slime mold, May 2004, S. A. Marshall, 29, DEBU [in alcohol]. UT: Summit Co., Coalville, 1710 m, 30 July 1973, P. H. Arnaud Jr., 2, CASC.
Comments. The abdominal pattern of the female is characteristic, but the male abdomen is entirely yellow, similar to that of several other North American species. Males can be best diagnosed by a frons that narrows posteriorly, an absence of white shoulder patches, and an open cell bm.
Sobarocephala latifrons (Loew, 1860) (Figs. 10, 48-50; Map 11) Heteroneura latifrons Loew, 1860: 82.
Heteromeringia latifrons, Johnson, 1913: 99. Malloch, 1918: 8. Sobarocephala latifrons, Melander & Argo, 1924: 42. Sabrosky & Steyskal, 1974: 381.
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Redescription (Fig. 10)
Male. Body length 3.3-4.5 mm. Bristles black. Two dorsocentral bristles plus one minute bristle in front of anterior dorsocentral. Acrostichal bristle present. Small, weak presutural intra-alar bristle sometimes present. Lateral scutellar bristles weak. Arista sparsely plumose. Sides of frons parallel. Scutum predominantly yellow with notopleuron brown and lateral margin behind suture light brown. Scutellum yellow with median brown stripe (darkest apically and sometimes faded basally). Laterotergites sometimes light brown lateral to scutellum. Pleuron and legs white with fore tarsi light brown. Head predominantly yellow, with face light yellow, ocellar spot large and triangular, and mouthparts, parafacial and gena white; gena pilose. Abdomen light yellow, often light brown on tergite 5 and laterally on tergites 2-4, and sometimes with small spot at base of epandrium. M.,, ratio 3.8-4.4. Wing clear. Cell bm open. Face flat.
Female. Similar to male except as follows: lateral brown margin on scutum wider; fore tarsi brown; scutellar stripe and lateral stripes on laterotergites strong and always present; laterotergites with thin stripe below scutellum; tergites 2-4 brown laterally, medially, on posterior margin and laterally on anterior margin; tergite 5 sometimes with wide median stripe (tapering anteriorly); tergite 6 and anterior margin of tergite 7 brown; wing dusky along anterior margin.
Male terminalia. (Figs. 48-50) Sternite 5 evenly setose. Sclerites of annulus well-developed. Epandrium as wide as high and length 4/5 height. Surstylus 4/5 height of epandrium and with rounded emargination from midpoint of posterior margin to apex
MAP 11. Distribution of Sobarocephala latifrons (Loew) (dot), S. wirthi spec. nov. (circle) and S. pengellyi spec. nov.
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(appearing “hook-like”’); tubercle-like bristles along posterior and apical margins. Cerci small and rounded with one slightly longer central bristle. Hypandrial lobe with three short distal bristles. Phallapodeme and postgonite well-developed. Pregonite elongate and ovate with four basal bristles, and with thin elongate projection on inner-medial surface. Basiphallus and epiphallus small. Distiphallus 2/3 length of phallapodeme; thumb and lateral lobe well-developed with lobe half length of distiphallus.
Distribution. Canada: ON, PQ. United States: DC, GA, IL, MA, MD, MI, NC, NJ, NY, PA, TX, VA, WV (Map 11).
Holotype. UNITED STATES, DC: “Osten Sacken”, 2, location unknown.
Paratype. UNITED STATES, VA: Fairfax Co., Dead Run, 22 June 1915, R. C. Shannon, 3, USNM.
Additional material examined. 1034, 1532 AMNH, [CASC, CNCI, DEBU, EMUS, TAMU, TAUI, USNM].
Comments. Sobarocephala latifrons is a commonly collected species in northeastern North America south of Ottawa and east of the Mississippi. Specimens have also been found in North Carolina, Georgia, and Texas. The only similar species with an overlapping range is the closely related S. wirthi, which is more southeastern in distribution. Males of these two species are distinct (see couplet 13), but females are more difficult to separate.
Sobarocephala muesebecki Sabrosky & Steyskal, 1974 (Figs. 7, 36-38; Map 1) Sobarocephala muesebecki Sabrosky & Steyskal, 1974: 378.
Redescription (Fig. 7)
Male. Body length 2.6-4.1 mm. Bristles brown. Two dorsocentral bristles plus one minute bristle in front of anterior dorsocentral. Acrostichal bristle absent. Two pairs of well-developed lateral scutellar bristles. Arista densely plumose. Sides of frons parallel. Body yellow with fore tibia and tarsi light brown, ocellar spot and infuscation at base of arista brown, epandrium dark brown, and coxae, gena, parafacial, and anterior (or lateral) margin of frons usually white; sometimes lateral half of postpronotum brown (Alabama), or postpronotum and notopleuron brownish (some North Carolina specimens). Occiput usually pilose. M,_,, ratio 4.0-4.5. Wing lightly clouded along R,,, on distal 1/3. Cell bm open. Face flat.
Female. Externally similar to male except as follows: fore tibia light brown; hind tibia and tarsi sometimes browned (Alabama, Florida, North Carolina); tergites 3, 4, and posterior margin of tergite 2 with central stripe; tergites 5-7 brown; tergite 8 brown or with anterior half light brown.
Male terminalia. (Figs. 36-38) Similar to S. atricornis (Figs. 33-35), except surstylus broader at base, tubercle-like bristles densely arranged along surstylus, pregonite
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widest basally, and lateral lobe strongly bent medially with small swelling at base of thumb; thumb short, pointed, and projecting at 90°.
Distribution. United States: AL, FL, NC, NJ, PA, TX (Map 1).
Holotype. UNITED STATES, FL: Levy Co., 7 mi NE of Cedar Key, 1 June 1970, D. L. Bailey, 3, USNM.
Allotype. UNITED STATES, FL: same collection as holotype, 2, USNM.
Paratypes examined. UNITED STATES, FL: Levy Co., 7 mi NE of Cedar Key, | June 1970, D. L. Bailey, 3, USNM; ex. Malaise, 18 May 1970, 2, USNM.
Additional material examined. UNITED STATES, AL: Baldwin Co., Bon Secour Nat. Wildlife Refuge, 5-7 May 1994, S. A. Marshall, 22, DEBU. FL: Santa Rosa Co., Blackwater R. St. Forest, 23 June 1973, W. W. Wirth, 2, USNM; Levy Co., 7 mi NE Cedar Key, | June 1970, D. L. Bailey, 4, 2, USNM; Cedar Key, ex Malaise trap, 18 May 1970, D. L. Bailey, 9, USNM; Highlands Co., Archbold Biol. Stn., 11 October 1964, P. H. Arnaud Jr., 42, CASC; 23 April 1967, B. V. Peterson, 2, CNCI. NC: Columbus Co., Lk. Waccamaw, 6 July 1985, W. Steiner & A. Gerberich, 9, USNM; Cumberland Co., Fort Bragg, J. D. Birchim, 23-25 August 1967, 2°, CASC; 16 August 1967, 2, CASC; Bladen Co., Singletary Lk. St. Pk., 34°35’0N, 78°27°30W, Malaise, 19-22 May 2003, Marshall & Paiero, 62, DEBU. NJ: Oswego Lk., Burlington Co., 30 August 1974, Menke & Miller, 4°, USNM. PA: State College, Centre Co., 4 July 1972, D. J. Shetlar, 9, CASC. TX: Salmon, Anderson Co., 22 June 1974, H. R. Burke, Malaise trap, 4, TAMU; 1-8 July 1974, 3,22, TAMU; | June 1974, 4,52, TAMU; Anderson Co., 10 mi SW Elkhart, H. R. Burke, modified Malaise, 5-6 June 1976, 29, TAMU; Brazos Co., College Stn., Wharton, Malaise trap, 20 September-4 October 1974, 4, TAMU; 22 July—2 August 1974, 49, TAMU; 14- 21 July 1974, 54, 22, TAMU: Montg. Co., Jones St. Forest, 8 mi S Conroe, 28 June—13 July 1987, Wharton, Steck & Carroll, 22, TAMU; 21-27 June 1987, Wharton, Steck & Carroll, 2, TAMU; 1-7 June 1987, Wang, Wharton & Praetorius, Malaise trap, 2, TAMU; 28 April—13 July 1987, Steck, Wharton & Carroll, 329, TAMU.
Comments. See comments for Sobarocephala affinis.
Sobarocephala pengellyi spec. nov. (Figs. 12, 42-44; Map 11)
Description (Fig. 12)
Male. Body length 2.9-3.8 mm. Bristles brown. Two dorsocentral bristles plus one minute bristle in front of anterior dorsocentral. Acrostichal bristle present. Two pairs of well-developed lateral scutellar bristles. Arista sparsely plumose. Sides of frons parallel. Scutum yellow with lateral margin brownish behind light brown notopleuron; scutellum with thin light brown central stripe extending onto base of scutum. Pleuron light yellow with anepisternum and anterior face of katepisternum yellow. Coxae white. Legs light yellow
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with fore tarsi light brown. Head yellow with face light brown centrally, gena and parafacial white and pilose, occiput white and ocellar tubercle brown. Abdomen predominantly yellow with wide light brown central stripe on tergites 2-4 (tapering anteriorly on each tergite) and epandrium brown (lighter on distal half). M,,, ratio 3.0. Wing clear or with light apical cloud around R,,,. Cell bm open. Face flat.
Female. As described for male except as follows: scutum with one pair of light brown lateral spots at base; scutellar stripe most pronounced apically (sometimes continuing onto base of scutum); laterotergite sometimes with light brown stripe lateral to scutellum and one stripe below; fore tarsi brown; face yellow (sometimes with orange tint); central stripe on tergites 2-4 brown, thin, and of equal width along length; tergites 5-7 brown and sternites 6 and 7 brown.
Male terminalia. (Figs. 42-44) Sternite 5 evenly setose. Sclerites of annulus well-developed. Length, height, and width of epandrium subequal; tapered to base. Surstylus 7/10 height of epandrium and broadly rounded; tubercle-like bristles along apical and posterior margins. Cerci small, rounded, and slightly projecting. Hypandrium with arm and ventral lobe subequal in length, and ventral lobe with two short medial bristles and one minute distal bristle. Phallapodeme and basiphallus well-developed. Epiphallus minute. Pregonite large and ovate with five central bristles. Postgonite absent. Distiphallus 7/10 length of phallapodeme; lateral lobe strongly bent, spinulose distally, with broad flat medial lobe, and with thumb absent.
Distribution. United States: AL, GA (Map 11).
Holotype. UNITED STATES, GA: Forsyth, 2 June 1970, 3, CNCI.
Paratypes. UNITED STATES, AL: Baldwin Co., Raft River Tributary, 16 October 2001, 30.52.261N, 87.56.934W, J. W. McCreadie, 22, DEBU. GA: same collection as holotype, 102, CNCI.
Comments. Sobarocephala pengellyi can be distinguished from congeners by a black epandrium, one pair of lateral subbasal spots on the female scutum, a light facial spot, a relatively short surstylus, and a characteristic lateral lobe on the distiphallus.
Etymology. The specific name is a patronym dedicated to the late D. H. Pengelly, a great entomologist whose spirit lives on in the community of entomologists associated with “his” insect collection, the University of Guelph Insect Collection.
Sobarocephala quadrimaculata Sos, 1963 (Figs. 1, 27-29; Map 3)
Sobarocephala nitida Sos, 1963: 393. Sobarocephala quadrimaculata Sos, 1963: 394. Sabrosky & Steyskal, 1974: 374.
Redescription (Fig. 1) Male. Body length 2.9-4.4 mm. Bristles yellow with lateral scutellar bristles dark
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brown. Two dorsocentral bristles. Acrostichal bristle absent. Two pairs of well-developed lateral scutellar bristles. Ocellar bristle absent. Arista densely plumose. Sides of frons converging posteriorly. Scutum yellow with one pair of short transverse presutural spots behind postpronotum and one pair of large, quadrate, postsutural spots on lateral margin; small triangular spot sometimes in front of scutellum. Scutellum and laterotergites dark brown. Pleuron yellow. Coxae white. Legs yellow with fore tibia and tarsi brown; fore tibia and basal 2/3 of fore tarsomere 1 yellow in lighter specimens; hind tibia sometimes light brown. Head yellow with parafacial and (sometimes) gena white, first flagellomere with brown dorsal stripe, and ocellar tubercle brown. Abdomen yellow with epandrium and tergites 2 (medially), 3, 4, 5 (medially) and 6 brown. M ,, ratio 4.0-4.2. Wing lightly clouded in cell R, and around bm-cu, and with dark infuscation distally on R,,, (appearing “smudged” to R,,.). Cell bm closed. Face convex on dorsal half below antennal bases.
Female. Externally as described for male except tergite 1 with large oval spot and abdomen past tergite 6 yellow. Female from Alabama with scutellum yellow medially.
Male terminalia. (Figs. 27—29) Sternite 5 evenly setose. Annulus with sternites 6 and 7 reduced to thin but well-sclerotized band ventrally. Epandrium as wide as high and almost as long as high. Surstylus rectangular; tubercle-like bristles long, curved, and terminal. Cerci flat with one longer central bristle. Hypandrial arm relatively long and sharply angled with base projecting at 90° basally; ventral lobe wide, slightly shorter than arm, and with one minute and two long distal bristles. Phallapodeme well-developed with head reduced. Pregonite long, thin, and setulose medially and apically. Postgonite small, rounded, and setulose. Basiphallus small. Epiphallus long and thin. Distiphallus *% length of phallapodeme and wide distally (appearing “spoon-shaped”’); lateral lobe large, broad, and serrate distally, with thumb absent.
Distribution. Bahamas, Colombia, Costa Rica, Mexico, Nicaragua, Panama, United States (AL, FL, GA, TX) (Map 3, in part).
Holotype. UNITED STATES, FL: 2°, USNM.
Additional material examined. COSTA RICA. Guanacaste: 14 km S Canas, F. D. Parker, 19-28 February 1990, 2, EMUS; 5-10 August 1990, 4, EMUS; 20-30 October 1989, 3, EMUS; 28 July 1991, 4, EMUS; 1-7 April 1991, 3, 52, EMUS; 8-15 February 1991, 24, 32, EMUS; 20-24 March 1989, 2, EMUS; 16-22 February 1990, ¢, 32, EMUS; 1-5 August 1992, 9, EMUS; 19-28 February 1990, ¢, 22, EMUS; 1-15 July 1991, d, EMUS; 16-26 January 1990, ¢, EMUS; 24-31 May 1990, 2¢, EMUS; 23-28 February 1990, 73, EMUS; 5-11 January 1991, ¢, EMUS; 1-10 July 1991, 2, EMUS; 2-4 June 1991, 29, EMUS; 9-14 February 1989, 2, EMUS; 3 km SE R. Naranjo, F. D. Parker, 15-25 January 1993, 9, EMUS; 21 July 1993, 2, EMUS; 11-20 December 1992, 2, EMUS; S Canas, F. D. Parker, 7-10 March 1989, 9, EMUS; 28 July 1991, 2, EMUS; 21-25 January 1989, 22, EMUS; 9-14 February 1989, 3, EMUS; Santa Cruz, P. N. Marino Las Baulas, 0 m, 14 December 2000, Malaise, Y. Cardenas, 2, INBC. Heredia: LaSelva Res. Sta., 11-17 June 1986, W. Hanson & G. Bohart, ¢, EMUS. Limon: Cuatro Esquinas, P. N. Tortuguero, 0 m, September 1989, J. Solano, 2, INBC. Puntarenas: Pen. De Osa, Puerto Jimenez, 10 m, P. Hanson, January 1991, 2, DEBU; May 1991, ¢, DEBU; November 1991, 9,
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DEBU; May 1992, 4, DEBU; 5 km N Puerto Jimenez, 10 m, May 1991, P. Hanson, 4, 29, DEBU; Malaise trap, June-July 1990, 9, USNM. COLOMBIA. Tol. Armero, Malaise trap, 30 January—5 February 1977, E. L. Peyton, 2, USNM. HONDURAS. Roatin Isl., 27 February 1979, G. E. Bohart, 9, EMUS, Roatin Isl (west), 1 January 1980, 29, EMUS. MEXICO. Jalisco: Puerto Vallarta, 1 January 1971, sea level, P. H. & N. Arnaud, 9, EMUS; Puerto Vallarta, G. E. Bohart, 10 December 1988, 29, EMUS; 8 December 1984, 3, EMUS; Quint. Roo, sweeping, F. Carillo, 10-14 October 1986, 24, EMUS. NICARAGUA. Puerto Rabezas, July 1971, J. Maldonado, 2, USNM. UNITED STATES, AL: Mobile Co., Chunchula (site 20), Malaise, 25 October 2004, 30.90N, 88.20W, E. Benton, 2°, DEBU. FL: Royal Palm Pk., 29 January 1933, A. L. Melander, 9, USNM; Gainesville, 8- 22 December 1986, hardwood forest, Malaise trap, W. Mason, 3, 9, CNCI. GA: Georgia: Liberty Co., St. Catherine’s Island, 18-21 September 1972, F. C. & B. J. Thompson, &, AMNH. TX: Austin, 28 October 1951, M. R. Wheeler, 2, AMNH.
Comments. Sobarocephala quadrimaculata is a relatively common species from Florida and Mexico to Colombia. .
Sobarocephala setipes Melander & Argo, 1924 (Figs. 15, 59-61; Map 12) Sobarocephala setipes Melander & Argo, 1924: 47. Sabrosky & Steyskal, 1974: 384.
Redescription (Fig. 15)
Male. Body length 2.4-2.9 mm. Bristles light brown. Two dorsocentral bristles plus one minute bristle in front of anterior dorsocentral. Acrostichal bristle absent. Two pairs of well-developed lateral scutellar bristles. Arista sparsely plumose. Sides of frons parallel. Scutum yellow with posterior corner of notopleuron and (often) one pair of wide basal stripes brown. Scutellum yellow (sometimes with brownish central tint). Laterotergites brown lateral to scutellum; yellow in specimens from Florida and holotype with posterior notal stripes also absent. Pleuron yellow, with meron and posterior half of katepisternum white. Legs yellow, with coxae and basal half of femora white and fore tarsi light brown to brown. Head predominantly yellow, with gena, parafacial, and occiput white and pilose, ocellar tubercle brown, and light infuscation sometimes around base of arista. Abdomen yellow with wide stripe on tergites 2-5 (also tergite 6 in one FL male). M__,, ratio 3.2-3.7. Wing clear. Cell bm open. Face flat.
Female. Similar to male except as follows: anterior half of notopleuron and postpronotum sometimes brown; notopleuron with central light brown spot; tergite 6 with wide central stripe. One female from Florida (CNCI) with distal 2/3 of fore tibia brown, and several females from Québec with fore tibia light brown. Ontario specimens sometimes with corners of scutellum brown.
Male terminalia. (Figs. 59-61) Sternite five with comb of bristles on posteromedial margin. Annulus reduced to thin band ventrally. Epandrium as high as wide and length 3/5 height. Surstylus short and rounded; tubercle-like bristles absent. Cerci small, rounded, slightly sunken below distal margin of epandrium, and bristles short and subequal in length. Hypandrial arm short, thin and projecting distally; three minute distal bristles on ventral
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lobe. Phallapodeme thin and shorter than length of hypandrium. Pregonite clavate with one distal bristle. Postgonite minute. Basiphallus 1/3 length of phallapodeme. Epiphallus absent. Distiphallus “4 length of phallapodeme; lateral lobe small and ovate with thumb absent.
Distribution. Canada: ON. United States: FL, IL, IN, LA, MD, MI, MS (Map 12). Holotype. UNITED STATES, MD: Marlboro, 19 June 1916, R. C. Shannon, ¢, USNM.
Additional material examined. CANADA, ON: Wellington Co., University of Guelph Arboretum, ex. dung, 8 August 2004, J. Klymko, ¢, DEBU [in alcohol]; 9 August 2004, O. Lonsdale, 22, DEBU [in alcohol]; 12 August 2004, O. Lonsdale, 34, 2, DEBU [in alcohol]; 19 July 2005, O. Lonsdale, 4, DEBU [in alcohol]; Essex Co., Point Pelee N. P., forested area by west beach, Malaise trap and pans, 10-21 July 1999, O. Lonsdale, ¢, DEBU; Carleton Place, 27 July 1959, J. G. Chillcott, 9, CNCI; Ottawa, J. R. Vockeroth, 13 August 1974, &, CNCI; 28 July 1964, 29, CNCI; damp second-growth Acer-Betula wood, 11 July 1991, 2, CNCI; 16 July 1991, 29, CNCI; 3 August 1992, 29, CNCI; Perth Rd., Rideau Tr., 14 July 1981, H. J. Teskey, 9, CNCI; North Gower, 10 August 1984, D. Bell, 2, CNCI. UNITED STATES, FL: Gulf Hammock, 23 April 1952, G. Peck, 2, CNCI; Sarasota Co., Myakka R. St. Pk., 21 May 1973, W. W. Wirth, Malaise trap, 2, USNM; Alachua Co., Pierce’s homestead, W. H. Pierce, 13 October 1973, Malaise trap, 9, USNM; Alachua Co., Chantilly Acres, 25 April 1970, W. W. Wirth, Malaise trap, 9, USNM; Gainesville, Doyle Corner
MAP 12. Distribution of Sobarocephala setipes Melander & Argo.
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Bldg., 23 September 1973, H. V. Weems Jr., Malaise trap, 29, USNM. IL: Champaign Co., Brownfield Woods, 2 mi NE Urbana, 26 June 1976, C. T. Maier, 29, EMUS; Equality, 11 September 1952, M. R. Wheeler, 22, AMNH. IN: La Fayette, 2 July [year not given] , 2, USNM; La Fayette, J. M. Aldrich, 5 August [year not given], 2, USNM; 31 July [year not given], 2, USNM; 23 July [year not given], 9, USNM; | August [year not given], 2, USNM. LA: Chicot S. Pk. nr. Ville Platte, September 1954, M. Wheeler, 2, USNM. MD: Montg’y Co., Dickerson, 14 July 1974, G. A. Foster, 4, 2, USNM: Glen Echo, J. R. Malloch, 16 July 1922, d, 2, USNM; 9 July 1922, 2, USNM; 2 July 1922, 23, USNM. MI: Wayne Co., Grosse Isle, 21 July 1957, G. C. Steyskal, 9, USNM; St. Joseph, Berrien Co., 17 April 1972, D. D. Wilder, ¢, CASC. MS: Forrest Co., 6 mi W Wiggins, Sweet Bay Bog, dung trap, 5-8 May 1994, sphagnum, S. A. Marshall, 2, DEBU.
Comments. Sobarocephala setipes is largely sympatric with the similar S. Jachnosternum, but S. lachnosternum occurs west into Saskatchewan, Lake Superior, Texas, and Utah (Map 9), and is not known from the southeastern United States.
Sobarocephala setipes is sometimes attracted to dung, and we have observed copulating pairs on dung baits in mid August in the University of Guelph Arboretum.
Sobarocephala texensis Sabrosky & Steyskal, 1974 (Figs. 17, 57, 58; Map 8) Sobarocephala texensis Sabrosky & Steyskal, 1974: 381.
Redescription (Fig. 17)
Male. Body length 2.6 mm. Bristles brown. Two dorsocentral bristles. Acrostichal bristle absent. Two pairs of well-developed lateral scutellar bristles. Arista sparsely plumose. Sides of frons parallel. Thorax yellow with narrow light brown spot from base of scutum to center of scutellum. Legs yellow with fore tibia and tarsi brown. Head predominantly yellow, with occiput white, gena and parafacial white and silvery tomentose, first flagellomere with infuscation at base of arista and ocellar tubercle brown. Abdomen yellow with wide central stripe on tergites 2-5. M_,, ratio 2.8-3.7. Wing lightly clouded along anterodistal margin. Cell bm open. Face flat.
Female. Externally as described for male.
Male terminalia. (Figs. 57,58) Sternite five with comb of bristles on posteromedial margin. Sclerites of annulus well-developed. Epandrium as wide as high and length 2/3 height. Surstylus rounded and 2/5 height of epandrium; tubercle-like bristles absent. Cerci small, rounded, slightly sunken below distal margin of epandrium, and with all bristles short. Hypandrium with one minute distal and one short and one long medial bristle on ventral lobe; arm atrophied, distal, and projecting at acute angle to long axis of phallapodeme. Phallapodeme relatively thin. Pregonite rectangular with one distal bristle. Basiphallus well-developed. Epiphallus and postgonite small. Distiphallus 2/3 length of phallapodeme with lateral lobe well-developed.
Distribution. United States: TX (Map 8).
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Holotype. UNITED STATES, TX: Rio Frio, Leakey, Real Co., 23 May 1972, W. W. Wirth, ¢, USNM [not examined].
Allotype. UNITED STATES, TX: same collection as holotype, 9, USNM.
Paratypes examined. UNITED STATES, TX: same collection as holotype, 24, 9°, USNM. ;
Comments. Sobarocephala texensis differs from the similarly coloured S. setipes (Fig. 15) in having a yellow thorax with a light median stripe on the scutellum that extends onto the base of the scutum.
Sobarocephala wirthi spec. nov. (Figs. 11, 45-47; Map 11)
Description (Fig. 11)
Male. Body length 3.3-5.0 mm. Bristles dark brown. Two dorsocentral bristles plus one minute bristle in front of anterior dorsocentral. Acrostichal bristle present. Lateral scutellar bristles weak. Arista sparsely plumose. Sides of frons parallel. Notum yellow with notopleuron and (sometimes) postpronotum brown; supra-alar spot sometimes present (Georgia) and confluent with notopleural spot. Pleuron, coxae, and femora white, with yellow spot on anepisternum. Legs yellow (entirely yellow in Georgia specimens) with fore tarsi brown and fore tibia usually light brown to brown. Scutellum with brown apical spot (highly reduced in some North Carolina specimens) that is sometimes wide and attaining anterior margin of scutellum. Laterotergites variably coloured. Head yellow with face light yellow, ocellar tubercle brown, and parafacial, gena, and occiput white; gena pilose. Abdomen yellow, sometimes with lateral margins of tergites (2)3-5 lightly infuscated (Georgia). M,,, ratio 3.3. Wing clear, but if scutellar stripe entire, wing dusky in cell R, and around distal 1/3 of R,,,. Cell bm open. Face flat.
Female. Externally similar to male except as follows: central stripe on scutellum weak if entire; abdomen partly yellow, with tergite 6 brown, tergite 7 brown basally and laterally, and posterolateral margins of tergites 2-5 with light brown spots.
Male terminalia. (Figs. 45-47) Sternite 5 evenly setose. Sclerites of annulus well-developed. Epandrium as wide as high and with length 4/5 height. Surstylus as high as epandrium and subtriangular; tubercle-like bristles along posterior and apical margins. Cerci small and rounded with one pair of longer central bristles. Hypandrium with one minute distal and two long medial bristles on ventral lobe, sometimes with two minute distal bristles and three long medial bristles. Phallapodeme well-developed with head thin and recurved. Pregonite thin and elongate with four distal bristles. Basiphallus and postgonite small. Epiphallus as large as basiphallus in outline. Distiphallus half length of phallapodeme; lateral lobe well-developed with thumb short, thin, and strongly projecting.
Distribution. United States: DE, FL,GA, MD, NC, NJ (Map 11).
Holotype. UNITED STATES, MD: Montg’y Co., Colesville, W. W. Wirth, 11 July 1974, 3, USNM. 100
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Paratypes. UNITED STATES, DE: Rehoboth, W.W. Wirth, Malaise trap, 18 July 1972, 3, 2, USNM. FL: Jackson Co., Florida Caverns State Pk., 26 May 1973, W. W. Wirth, Malaise, 3, USNM; Liberty Co., Torreya State Pk., 13 June 1974, H.'V. Weems Jr. & C. R. Artaud, Malaise trap, 2, USNM; 5 July 1965, H. V. Weems Jr., Malaise trap, 2, USNM; Alachua Co., Gainesville, Austin Cary Forest, G. B. Fairchild, 30 August 1976, 2, USNM; 30 July 1976, 2, USNM; 11 June 1976, insect flight trap, 6, 9, USNM. GA: Athens, 8- 11 July 1969, R.