a revision of the genus aleochara gravenhorst of ...

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A REVISION OF THE GENUS ALEOCHARA GRAVENHORST OF AMERICA NORTH OF MEXICO (COLEOPTERA: STAPHYL|NtDAE, ALEOCHARTNAE) JAN KLIMASZEWSKI Biosystematics Research Institute Agriculture Canada Ottawa, Canada K1A 0C6

MEMOIRS OF THE ENTOMOLOGICAL SocIETY oF CANADA-No, 129 I

I

Stephen M. Smith, Editor

ENToMoLoclcAL soctETy oF OANADA 1320 carting Avenue ottawa K1z7Kg

1984

(613) 725_2619

CONTENTS PecE

Abstract/R6sum6

J


Introduction

A

Methods and materials.

4

Explanation of terms

6

Genus AZEO C H ARA Gravenhorst

7

Key to subgenera of Aleochara

T4

l.

l4

Subgenus Coprochara Mulsant and Rey. Key to species of Coprochara . .....

16

Bimaculata Group 1. Aleochara (C oprochara) 2. Aleochara (Coprochara) 3. Aleochara (Coprochara) 4. Aleochara (C oprochara) 5. Aleochara (Coprochara) 6. Aleochara (Coprochara) Sulcicollis Group J . Aleochara (Coprochara)

n

bimaculata Gravenhorst notula Erichson. verna Say ... densis sima Bernhauer

20 22

26

suffusa (Casey)

21

bilineata Gyllenhal

29

sulcicollis Mannerheim 2. Subgenus Xenochara Mulsant and Rey Key to species of Xenochara Zrislis Group 8. Aleochara (Xenochara) tristis Gravenhorst .. Lacertina Grotp 9. Aleochara (Xenochara) lacertina Sharp . Sculpliveturis Group 10. Aleochara (Xenochara) sculptiventrls (Casey). ll. Aleochara (Xenochara) sallaei Sha.p . 72. Aleochara (Xenochara) taeniata Erichson. 13. Aleochara (Xenochara) puberula Klug . . Lanuginosa Group I 4 . Ale o c har a (Xe no chara) lanugino sa Gravenhorst 15 . Aleochara (Xenochara) inexspectata sp. nov. Fumata Group | 6 . A I e o c har a (X e no c har a) fumat a Gravenhorst. 17 . Aleochara (Xenochara) castaneipennis Mannerheim 18. Aleochara (Xenochara) quadrata Sharp. 19. Aleochara (Xenochara) arizonica sp. nov.

3. Subgenus Calochara Casey. Key to species of Calochara

)L

34 36

JI 39

42 44 44 46 48

5l

.....

51

54 57 58 59

60

Speculicollis Group 20 . AI e o c har a (C al o c hara) sp e c uli c o lt is B ernhauer Laramiensis Grotp 21 . Aleochara (Calochara) laramiensis (Casey). Villosa Group 22. Aleochara (Calochara) villosa Mannerheim

61

62 63


Rubripennis Group 23. Aleochara (Calochara) 24 . Al e o c har a (C al o c har a) 25. Aleochara (Calochara) Rubricalis Group 26. Aleochara (Calochara)

nidicola sp. nov.

65 66 61

rubricalis (Casey)

69

cavernicola sp' nov. rubr ip e nn i s (Casey)

.

10

4. Subgenus Aleochara Mulsant and Rey

-1 tl

Key to species of Aleochara Lustrica Grottp . Aleochara (Aleochara) 28. Aleochara (Aleochara) 29. Aleochara (Aleochara) 30. Aleochara (Aleochara) Gracilicornis Group 3 l . Ale ochar a (Al e o chara) 32. Aleochara (Aleochara) 33. Aleochara (Aleochara) 34. Aleochara (Aleochara) 35. Aleochara (Aleochara) 36. Aleochara (Aleochara)

12

Iustrica Say . . . centralis Sharp.

2'7

curtula (Goeze) lata Gravenhorst

15 '76 .

78

.

81 83 85

ornis Bernhauer tahoensis Casey. thoracica Casey. rufobrunnea sp. nov.

gr acilic

81 87 89

rufonigra sp. nov. unicolor sp. nov.

89

5. Subgenus Echochara Casey.

90

Key to species of Echochara Lobata Group 31. Aleochara (Echochara) lobata sp. nov.

9l

Lucifuga Grotp 38. Aleochara (Echochara) lucifuga (Casey) 39. Aleochara (Echochara) ocularis sp. nov. 40. Aleochara (Echochara) fenyesi Bernhauer .. ..

6. Subgenus Emplenota Casey

.

Key to species of Emplenota 41. Aleochara (Emplenota) littoralis (Maklin) 42. Aleochara (Emplenota) pacifica (Casey) 43. Aleochara (Emplenota) curtidens sp. nov.

.

94 95 96

96 100 101

t02

7. Subgenus Maseochara Sharp. Key to species of Maseochara. .. Valida Group

92 93

.

.

44. Aleochara (Maseochara) valida LeConte 45. Aleochara (Maseochara) opacella (Sharp) 46. Aleochara (Maseochara) depressa (Sharp)

Wickhami Group 4'/ . Aleochara (Maseochara) wickhami (Casey) Checklist of Nearctic and introduced or Holarctic species of Aleochara

103

to4 107

108 111

References.

tt2 lr4

Illustrations (Figs. 5-51 1) . .. ..

r20

Index

209

.

.

Ftc. 1 . Frontisp tece Aleochara

(C

oprochara) bimaculata Gravenhorst


A REVISION OF THE GENUS ALEOCHARA GRAVENHORST OF AMERICA NORTH OF MEXICO (COLEOPTERA: STAPHYLINIDAE, ALEOCHARINAE)


JaN Kr-rlreszEwsKrl

Abstract A revision of the

of America north of Mexico is presented. Seven subgenera are redefined with 47 cunently recognized species, of which nine are genus Aleochara

described as new (A. arizonica, A. cavernicola, A. curtidens, A. lobata, A. ocularis, A. nidicola, A. rufobrunnea, A. rufonigra, A. unicolor). The species are arranged

according to their relationships into species groups within each subgenus. sixty-one synonymies are newly established (the first specific name is valid): Subg. COPROCHARA, A. bimaculata Gravenhorst : B. deserticolaCasey : B. innocuaCasey : B. obsolescensCasey : B. rectaCasey;,4. notulaErrchson: A. duplic,c/aErichson : B. nanella Casey : B. nitidicollis Casey; A. suffusa (Casey) : B. acuminata Casey : A. verna brundini Bernhauer; A. vernd Say : 4. alticola Sharp. Subg.

: B. salicola Casey : B. imbricata Casey : B. idonea Casey : A. densiventrisBemhauer; A. sallaei Sharp : A. torquatasharp; A. puberulaKlug : B. bipartitaCasey;A. fumataGravenhorst: A. languidasachse : B. defecta Casey : B. ffiuens Casey; A. castaneipennis Mannerherm : A. glenorana Casey : B. rotundicollis Casey : B. mannerheimi Casey : B. acomanaCasey: B. insulanaCasey: B. concurrensCasey: B. eludensCasey : A. oregona Hatch; A. quadrata Sharp : B. uvidulaCasey : A. sparsicollisBemhauer. Subg. CALOCHARA, A. rubripennls Casey : A. rubripes Blatchley : A. wallawallaeHatch. Subg.ALEOCHARA,A. centalis Sharp : A. srygialisSharp; A. gracilicornls Bernhauer : A. kansana Casey -- A. pleuralls Casey A. americana Casey : A. ellipsicollis Casey;A. lustrica Say : a. pauper Sharp : A. serrata Shu.p : A. texana Casey : A. fusicornis Casey : A. sternalis Casey : XENOCHARA, A. lacertina Sharp

A. algonquina Casey : A. medialis Casey;A. tahoensis Casey : A. postpictaCasey : A. montanica Casey; A. thoracica Casey : A. collusor Casey. Subg. ECHOCHARA, A. lucifuga (Casey) : A. cavicola (Garman). Subg. EMpLENOTA, A. littoralis (Miiklin) : A. maritima (Casey) : P. arenaria Casey : E. quadrifer Casey : E. trilimbata Casey : E. longiceps Casey. Subg. MASEOCHARA, A. valida LeConte : M. ponderosa Casey : M. robusta Sharp : M. ruficauda Casey; A. opacelLa (Sharp) : M. hogei Sharp : M. decipiens Casey; A. depressa (Sharp) : M. puberulaCasey : M. basalis Casey : M. mustaCasey. Ninety-three lectotypes and three neotypes are designated (neotypes are designated for'.

C. caviola Garman, A. lustrica Say, and A. verna

Say).

Keys to subgenera and species are provided. All species are described, each illustrated with line drawings and scanning electron microscope photomicrographs, and all available bionomic and distributional data are given. Emphasis was put on host records of those Aleochara species which may be used in biological control of some pest specles of Diptera. The systematics and phylogeny of the subgenera and zoogeography of species are discussed.

A checklist of Nearctic Aleochara and a table listins the known host records for some species is added.

R6sum6 L'auteur r€vise le genre ALeochara en Am6rique du Nord. Il red6finit et reconnait 7 sous-genreset4Tespdcesdont9nouvelles(A. arizonica,A. cavernicola,A. curtidens, A. lobata,A. ocularis,A. nidicola,A. rufobrunnea,A. rufonigraetA. unicolor),les espdces 6tant regroup6es en groupes d'espdces i l'int6rieur de chaque sous-genre. Curent address: Entomology Department and Lyman Entomological Museum, Macdonald Cmpus of McGill University, Ste. Anne de Bellevue, Qu6bec H9X 1C0.

]\4EMOIRS OF THE ENTOMOLOGICAL SOCIETY OF CANADA


Soixante et un nom d'espbces deviennent synonymes, 93 lectotypes et 3 n6otypes ont 6t6 d6signds. Les sous-genres et les espdces peuvent 6tre d6termin6es d I'aide de cl6s d'identification. Toutes les descriptions d'espdces sont illustr6es de dessins et de photographies au microscopes 6lectronique d balayage et compl6t6es par des informations sur la biologie et la r6partition g6ographique de celles-ci. L'emphase a notamment port6 sur les espbces d'Aleochara susceptibles d'Otre utilis6es dans la lutte biologique contre certains diptdres nuisibles. Enfin, l'auteur discute de la syst6matique et de la phylogdnie des sous-genres ainsi que de la zoogiographie des espbces.

INTRODUCTION

This is my third revision of Nearctic genera and species of the poorly known subfamily Aleocharinae. The genus Aleochara, in spite of its importance in the control of many pest species of Diptera, is poorly known and greatly in need of a modern revision. The numerous undesignated synonyms, lack of adequate keys for specific identification, usage of superlicial and often useless external characters (e.g. body length) and total lack of illustrations, made species identification difficult, if not impossible. Furthermore the genus had been divided into numerous, poorly defined genera. The study of the Nearctic fauna of Aleochara is also complicated by the presence of some introduced Palearctic species, Holarctic species formerly described from Europe and other species described from Mexico. Central or even South America which reach their northern distribution limits in the southern part of the Nearctic region. The only available source of information on the Nearctic species of Aleochara has been Casey's (1906) review of the species and genera of the tribe Aleocharini. Unfortunately, Casey did not use many of the most important characters for species recognition i.e. shape of the median lobe of the aedeagus and its internal structures, shape of the spermatheca, and shape of the terminal abdominal segments. As a result, Casey described 61 species of which 49 I have placed in synonymy. His concept of species having limited distributional ranges and his ignorance of the European, Mexican and Central American fauna, substantially decreased his contribution to the knowledge of this group. However, his paper markedly contributed to the knowledge of the genus as it was the first attempt to treat the Nearctic species of Aleochara systematically. The numerous synonyms proposed in this paper were based on careful analysis of the type specimens, including external and genitalic characters. Among extetnal characters the most reliable prove to be the shape of pronotum, the pattern of pubescence on pronotum and elytra, the size and pattem of punctation on pronotum, the color of upper surface of body, and the pattern of microsculpture on pronotum and elytra if applicable. The genitalic characters prove to be surprisingly consistent and were considered as the ultimate criteria for species recognition. The most reliable were the shape of median lobe in lateral and dorsal views, the shape and location of structures of internal sac of median lobe, and the shape of spermatheca (for details see discussion under aedeagus and spermatheca in "Explanation of terms"). The present study was conducted as a Postdoctoral Fellowship project at the Biosystematics Research Institute, Ottawa, under the supervision of Dr. A. Smetana. It in-

corporates all available information about the systematics, phylogeny, life history and distribution of the adults, and includes numerous scanning electron photomicrographs of external features and line drawings of terminalia and genitalia. This paper is far from perfection, but I hope it will ease the existing problem of identification, contribute to a better understanding ofthis group, and possibly encourage some entomologists to conduct further, comprehensive studies of the life histories of the Nearctic spectes' METHODS AND MATERIALS This study was based exclusively on examination of about 10,000 adults from North America, and some from Mexico, Central and South America, and Europe. Therefore,

THE GENUS

ALEOCHARA

5


whenever a "species" is compared with another "species" or a "subgenus" with another "subgenus", it is understood that actually the adults of the two particular taxa are being

compared. Lectotypes were designated for 92 species, whenever holotypes rvere not clearly fixed in original descriptions. The type locality on original labels often included only general data, like states or provinces, while more specific data, like towns, counties, etc., were given in original descriptions. Both data are given under the "type locality" with a clear division between those on the original labels and those in original descriptions. Types of 102 published names were studied, and three neotypes were dJsignated for types

presumably missing or destroyed. Comparisons were made between external charailers and also the terminalia and genitalia of both sexes for each species. The terminalia and genitalia of both sexes were illustrated by line drawings, and the external characters by scanning electron photomicrographs. Most of the specimens, including types, were dijsected, and their terminalia and genitalia mounted in Canada balsam on plastic plates as described by Smetana (1982) . A list of localities is given for each species, with references to the name of the collector, name of collection or individual from whom specimens were borrowed, and the number of specimens studied except for very common species where only the localities are listed to save space. The types of mosi species deicribed from Europe have not been examined; I have followed the synonymy established bv Bernhauer

"to TT:;t":tiirt,?,?uL"o,

was borrowed from u nu-r". or,^,uu,,ons and some privare collections. The abbreviations listed below refer to the material in the text. The assistance of the curators responsible for these collections and the assistance of a few individuals who sent the specimens for study is gratefully acknowledged.

AMNH

American Museum

york,

of Natural History, New Dr. L.H. Herman, Jr. - British Museum (Narural History), London, Mr. p.M. Hammond CAS - California Academy of Sciences, San Francisco, Dr. D.H. Kavanaush CNC - Canadian National Collection, Ottawa DBC - Mr. D.B. Conn, University of Cincinnati, Ohio, personal collection DEI - Deutsches Entomologisches Institut, Eberswatde, Dr. L. Dieckmann FG - Mr. F. G6nier, Ste. Anne de Bellevue, personal collection FMNH - Field Museum ol Narural Hisrorry. Chicaso. Dr. E.H. Smith JHF - Dr. J.H. Frank, Vero Beach, peisonal collection - Dr. L. LeSage, Ottawa, personal collection LL MCZ - Museum of Comparative Zoology, Harvard University, Cambridge, Mass., Dr. A.F. Newton MZH Museum Zoologicum Universitatis, Helsinki, Mag. phil. H. Silfverberg NMP - National Museum of Natural History, paris. Mlle. M.N. Berti OSU - Oregon State University. Corvallis, Mr. G.L. peters PUI - Purdue Univeisity, Lafayette, Ind., Mr. A. provonsha RS - Mr. R. Sexton, Ottawa. Dersonal collection SM - Dr. S. Miller, Santa Barbara, Calif., personal collection SP - Dr. S. Peck, Ottawa. personal collection usNM - United States Nationai Museum, now known as National Museum of Natural History, Washington, D.C. Dr. T.L. Erwin zMB Zoologisches Museum der Humboldtuniversitdt zu Berlin, Berlin, Dr. M. Uhlig I am most grateful to Dr. A. smetana who supervised this project, and whose en-

BMNH

couragement and substantial help, including editorial work on the manuscript and help with English, enabled me to finish it in a relatively short time. Without his contribution the project could not have been completed. I heartily thank Dr. J.M. Campbell for his helpful suggestions and assistance with English, Dr. J.H. Frank for sending many rare specimens together with bionomic data, Mr. E. Rickey for his assistance in pieparing the scanning electron photomicrographs, and Mr. G. Sato, who made the habitus d.awingr. I also thank Dr. Z. Likovsk! from Prague, czechoslovakia, who sent me many Europein

MEMoIRS oF THE ENToMoLoGIcAL socIETY oF CANADA

6 specimens and

Dr. D.K. McE. Kevan who allowed me to complete this project after

b^eginning my tenure as a postdoctoral fellow at the Lyman Entomolo-gical Museum' The


utJirtun.i of nt. G.R. Lohse from Hamburg, Germany, is also gratefully acknowledged' EXPLANATION OF TERMS !: in bibliographic citation indicates that the holotype has been examined. Terminalia (1 ierminal segments): terga and sterna VIII and IX, also tergum X. Genitalia: aedeagus (aedeagi), spermatheca(e), or both combined'

wider than base of setae. Punctures coarse -no wider than base of setae' Punctures fine

Aedeagus-(Figs. 1G-13): the male copulatory organ consisting of the median lobe t*o pa.ameres. The median lobe is a tubular sclerotized, intromittent organ 1: with an enlarged basal portion called the bulbus and a narrow apical portion called the tubus. The bulbus has a small opening (foramen mediale) in the ventral part, which is an entrance for the ejaculatory duct (ductus ejaculatorius) and an external carina in the anterior part, which serves is an attachment for muscles connecting the bulbus with the condylites of the parameres (see further in the text). The internal space of bulbus contains mainiy a complei of muscles which is directly or indirectly responsible for eversion and retraciion of ihe internal sac and its associated structures. The dorsal part of the median lobe is covered by membranous, mobile lamina (: compressor plate' Seevers 1978), which is sometimes partially weakly sclerotized. The vertical muscles of the bulbus are apically attached to tiris lamina. The apical portion of the median lobe (tubus) is slender, subcylindrical and narrowed towards tlie apex. The shape of the apical part of median lobe varies from species to species or from species group to species group, and is a good penis) and

taxonomic character. The internal sac and its structures are the most complex parts of the aedeagus, and are the most useful characters for species identification in this genus. The surface of the internal sac may be smooth or with numerous spinules, granulations, or other fine sculptures which pro6ably aide in retention of the sac in the female genital chamber' The internal sac is retracied in repose. There are several accessory structures inside the internal sac: sclerites and sclerotized flagellum ( : virga) ' The flagellum is a narrow' tubular structure with an expanded, funnel-shaped basal part which is in contact with the ejaculatory duct. The flagelium serves to introduce sperm into the female ductus receptaculi (Peschke l978d1 It is sometimes very long with several coils in the bulbus (A. bimaculata, A. tristis), or may be short, at most as long as half the length of the median lobe (e.g. species of subgenus Echochara, and some of subgenus Aleochara). The sclerites of the internal ,u. u." very diverse and usually have a complex shape. These sclerites serve as accessory srrucrures, along with the microsculpture of the internal sac membrane, which retain the intemal sac in ihe female genital chamber during copulation (Seevers 1978; peschke 1918a). Peschke (1918a) described the shape and function of the structures of the internal sac in A. curtuLa, and found a correlation between their shape and the structures of the female genital chamber. Because of the great diversity of the internal sac structures in this genus, iwas usually unable to establish homology between the structures in different speciesl. I was able to establish certain homology between internal sac structures only in species of the subgenus Coprochara and within some species groups of the other subgenera of Aleochara. In cases whLere the homology was evident, homologous structures are in-

dicatedindrawingsbythesameletters(e.g. x,y,z).Veryoftenlreferonlytotheapical or subapical struciures (sclerites) to avoid ionfusion. The shape, number, and position of the struitures in the internal sac are the most useful criteria for species, species group, and sometimes subgeneric identification. The shape of the structures is surprisingly consistent regardless of zoogeographic distribution of specimens'


parameres,.,r,

';:i;;"'";.

ffi" ffiil;;:;r'.h

enabre

tt'"

-4" tJ

find the tip of the female abdomen just before copulation, using the setal pattern on the female abdomen as a guide (Peschke 1919). Each paramere is composed of two parts the condylite and the paramerite, articulately connected. The condylite is a slender, -elongate structure, which articulates with the bulbus of the median lobe and also with the condylite of the other paramere. The paramerite is a complex structure which articulates with the condylite. The apical part of the paramerite consists of a narrow lobe bearing 3-4 prominent setae and a semimembranous velar sac. The basal part of the paramerite is a slat-shaped structure, delimited from the apical portion by a weakly sclerotized strip. The basal part bears a fragma on its internal surface for the attachment of muscles. The shape of the paramere is used here as a generic character. The spermatheca (receptaculum seminis) (Fig. 14) is the female genital organ for

storing and perhaps selecting sperm (Dybas et Dybas 198 1). The posterior part of the spermatheca is fused with a thin, weakly sclerotized, seminal canal(ductus spermathecae), which connects it with the vagina and serves to transport the sperm. The seminal canal is thin and very long. The seminal canal is not considered here as a morphological part of the spermatheca. The spermatheca in most species of Aleochara consists of three clearly defined divisions: a more or less spherical capsule, an enlarged chamber and a narrower duct (Fig. l4) but there are many modifications of these divisions. Sometimes all three divisions are fused together to form a single unit (see discussion under female genitalia in generic description). The shape of the spermathecal divisions and also the way they are connected with each other is ofa great value for separating species ofthis genus.

SYSTEMATICS

Genus ALEOCHARA Gravenhorst Aleochara Gravenhorst, 1802: 61; Mannerheim, 1830: 480; Erichson, 1831-39 (1837): 353, 1839-40 (1839): 158; Redtenbacher, 1849: 668, 1858: 154,1814: 136;Lacordaire, 1854: 40,59 Kraatz,1856-58 (1856): 82; Jacquelin du Val, 1857-59 (1856): 12; Kraatz, 1859a: l0; Fauvel, 1864-65 (1866): 285; Harold, 1884: 124, Fauvel 1884: 16,304; Olliff, 1886: 455; Ganglbauer, 1895: 25; Fowler, 1888: 10; Everts, 1898: 165; Bernhauer, 1901b:436; Dubois, 1906-07:22; Reitter, 1909:22; Blatchley, 1910: 360, 363; Johansen, 1914: 15 Fenyes, 1920:396; Cameron, 1920:361, 397; Kr6sa, 1922: 8V82; Everls, 1922: 74, 75; Bernhauer and Scheerpeltz, 1926: 755; Portevin, 1929 234; Scheerpeltz, 1930: 80, 1934: 1708; Blackwelder, 1943: 559. 1952:43; Hatch, 1957 136; Horion, 1961: 368 Strand and Vik, 1968: 105; Palm,1972'.418; Likovsky,l914:293; Moore and Legner, 1915 321; Seevers, 1978: 135. Life history: Westwood, 1839: 166; Sprague, l810a,b: 302, 310 Coquillett, 1891: 318; Riley, 1893: 405-410; Slingerland, 1894:481-5'17; Ganglbauer, 1895: 26; Xambeu, 1898: 184, 1900 22,1902:95,1909: 11; Blatchley, 1910: 360, 363; Bickhardt. l9l2 181.188;Wadsworth, 1915: 1-27; Schoene, 1916: 99-160; Scott, 1916:206,1920: 148; Lesne and Mercier, 1922:351-58; Kemner, 1926: 133-110; Boving and Craighead, 1930: 29; Lindquist , 1936: 816-11; Burks, 1952: 379; Colhoun, 1953: 1-8; Hatch, 1957 384 Wisharl, 1957:450-54; Read, 1962:411-24; Wingo, 1963: 54; Drea, 1966: 1368-73 White and Legner, 1966: 513-7'7; Jones, 1961: 816-11; Wingo et al. 1967 l5l4-17 Moore and Legner, l91l | 184; Peschke and Fuldner, 1971:242-262.; Peschke, 1919: 155-159. Type-species staphylinusfuscipesLinn6.FixedbyLeach,1819:lll,bysubsequent designation.

Fungicola Zetterstedt, 1840: 78. - As synonym of Aleochara: Blackwelder, 1952: 166; Moore and Lesner. 1915: 321 .

o rro.-,0..'",'

rffi;,ffiI"l",l"rlo'*;;;i#.,,

1838: 78, rhrough

objective synonymy with Aleochara, of which fuscipes had already been fixed

as

type-species.

Solier, 1849 : 341 . - As synonym of Aleochara: Bernhauer and Scheerpeltz, 1926: 11 5; Blackwelder, 1952: 232; Moore and Legner, 191 5: 327 . Type-species: Mecorhopalus elongatus Solier. Fixed by Chenu and Desmarest, 1857: 18, by subsequent designation.


M e corhop alus

Copiata des Gozis, 1886: 12. - As synonym of ALeochara: Blackwelder, 1952: 105; Moore and Legner, 1975:32'7. Type-species: StaphyLinus fuscipes Linn6. Fixed by des Gozis, 1886: 12, by original designation.

Opiochara Bernhauer, 1901b:439.- Assynonym ofAleochara: Blackwelder,1952:216; Moore and Legner, 1915: 327 . Type-species'. Aleochara breiti Ganglbauer. Fixed by Fenyes, l9l8:24, by subsequent designation.

Description. Antenna with 11 segments, scape somewhat clavate, usually longer or as longaspedicel(Figs. 1,3,4);segment3slightlyshorterthanpedicel(Figs. 1,3,4),and segment 4 usually shortest of the 4 basal segments (Figs. 1 , 3, 4). Maxilla with palpus 4-segmented with minute apical pseudosegment (Figs. 243-245); lacinia wide, multispinose (Figs. 253, 259); galea wide, as long as lacinia, with tuft of apical setae (Figs. 253, 259). Labium with palpus 3-segmented with a more or less defined apical pseudosegment

(Figs. 243, 253,255,257,261,263); ligula small, bilobed apically (Fi9s.322,324); mentum trapezoidal, usually slightly emarginate apically (Figs. 258, 260,262); submentum as mentum reversed in shape (Figs. 258, 260, 262); gula narrow (Figs. 258, 260, 262); gena expanded (Figs. 258, 260,262). Procoxae conically elongate. prominent; mesocoxae broadly oval; metacoxae transverse, triangularly elongate apically, contiguous near base (Figs.249-25 1); tarsal formula 5,5,5. Mesosternum variable, ranging from completely carinate through morphocline to lacking carina (Figs. 248-251 , 349-351 , 502-505). Body pubescent. Larvae (as far as known) are ectoparasitoids within puparia of cyclorrhaphous Diptera where they undergo hypermetamorphosis (although the life history of many Aleochara species is still unknown, it seems likely that this is a generic character).

Male. Tergum and sternum VIII pubescent, with prominent setae distributed in apical part (Figs. 4446, 50-51 , 53-54,91-95). Tergum IX divided, bearing ventral struts, and with both parts separated by base of tergum X (Figs. 5, 6). Tergum X disc-shaped, often shallowly emarginate apically and with stout apical setae (Figs. 5, 6, 4749). Aedeagus (Figs. 10-13) with median lobe having enlarged bulbus portion bearing anterior carina or small projection serving as attachment of muscles; internal sac mostly with complex structures, flagellum unusually long to moderately short; paramere consisting of two parts (condylite and paramerite, Figs. 12, 13) with 3-4 apical setae. Female. Tergum and sternum VIII usually broadly rounded apically, pubescent, with prominent setae distributed in apical parl. Tergum IX divided (Figs. 7, 8), with both parts separated by base of tergum X, unlike in male without ventral struts. Tergum X similar to that of male, usually with shallow apical emargination (Figs. 7, 8). Sternum IX not apparent, possibly fused with ventral parts of tergum IX. Spermatheca (Figs. 14, 17-19, 63-65, 68-11, 140, 142,145) consists of three morphological divisions (capsule, chamber, duct), the duct is fused with thin, seminal canal. There is a considerable number of modifications of spermathecal divisions . In subgenus C oprochara the capsule and chamber are fused together and are connected with a sclerotized, basally coiled duct (Figs. l7-19, 24,25,28,29); inthe Lacertina and Lustrica groups the capsule, chamber, and duct are


KLIMASZEWSKI: REVISION OF THE GENUS ALEOCHARA

A

/,A\/,a\ / (:.:) \r e

tr=-=--'-7 \-....-.----, t-_,,-7 srJ

+ \'/ 1 \/V 1u V o o o

--)

3

'

As Ji

/'{j/

| l---*l | t--'-{ L-r\l

2 []i \ll

A-1

\ l

.^-m ',A-4\ z--\ ,1 / \\ /1 / \ \

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/i iljl\ Li: rlil

. \l/ \Y

I

ffi*h.l vv

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a

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2A C. Some ol the possrble phylogenetic trees of Aleochara based on the character states listed in Table (solid cirr:les: apomorphic character states; open circles: plesiomorphic character states). The character state marked with an asterisk has plesiomorphic state in subgenus Emplenota. A : subgenus Aleochora, CA subgenus Calochara, CO : subgenus Coprochara, EC - subgenus Echochara, Em : subgenus Emplenota,

FIG.

II

M:

subgenus Maseochara.

X:

subsenus Xenochara.

l0

MEMorRs oF THE ENToMoLoclcAL soclETY oF cANADA


fused together (Figs. 59,140,142,145);inthe Fumata, Sculptiventrls, and Valida groups the capsule is fused with the chamber (Figs. 63-65, 81,206,207 ,212). Bionomics. All species of which the life history is known, are active predators of larvae and eggs of cyclorrhaphous Diptera as adults, and ectoparasitoids of the pupae within the puparium as larvae. Both states are frequently found in decaying plant substances, dung,

or carrion infested with dipteran larvae. (See Table I.) " Females lay their eggs in sites infested with fly larvae, and the newly emerged first instar larvae of Aleochara hunt for dipteran pupae (Burks 1952). The larva gnaws a hole in the puparium, crawls inside, seals the entrance behind it, and feeds on the pupa (Burks 1952; Feichke and Fuldner 1911). The most recent descriptions of larvae of some species of Aleochara are given by Fuldner (1960), Peschke and Fuldner (1911), and White and Legner (1966). Geographic distribution (Maps 1-32). Aleochara is a worldwide genus of approximately 300 ipelies, of which none is known from Antarctica. The majority of species occur in the Niarctic and Palearctic regions. The Nearctic fauna contains 47 currently recognized species; some of them are introduced from Europe (A. bilineata, A. curtula, A. Jumata, i. Ianuginosa, A. lata, A. tristis, and A. villosa); two possibly have a genuine Holarctic distribuiion (A. verna and A. suffusa); and one is a cosmopolitan species (A. puberula). Distributions of North American species are of six major types: I, northern and western Nearctic; II, western Nearctic; III, eastern Nearctic; IV, transcontinental, common throughout Nearctic; V, southern elements, with northern range in southern United States; VI, both coastal regions or only western coast. Phylogenetic relationships. The phylogeny presented here is limited to the Nearctic fauna lecause species of other zoogeographic regions have not yet been revised' Seven subgenera are recognized in the Nearctic fauna and their phylogenies are shown in Fig. zA--C. The morphological characters (Table II) were recognized as apo- or plesiomorphic : primitive") and outstates according to thJwell known criteria of in-group ("common group comparisons (Watrous and Wheeler 1981). - The subgen".u of the genus Aleochara (present sense) were used for in-group comparisons and the genera of the tribe Hoplandriini (sensu Seevers 1978) fbr out-group comparisons. I have-followed Seevers'concept in considering the tribe Hoplandriini as the most closely related to Aleocharini (sensu Seevers 1978). The latter is approximately equivalent to the genus Aleochara (present sense)' The most important characters for the reconstruction of phylogeny and for classification of the genis are those associated with the mesosternum: absence or presence and length of carina (Table II, characters 1, 1a). These characters proved to be invariable within the subgenera, but are polarized within the genus from a completely carinate state of the mesosternum through partly carinate to non-carinate. Within the genus Aleochara the carinate mesosternum occurs in three subgenera (Coprochara,Xenochara, Calochara) and is absent in the other four (ALeochara, Echochara, Emplenota, Maseochara). The out-group comparison proved the presence of the mesosternal carina in the genus Tinotus2 Strarp and the absence of the carina in three other genera of Hoplandriini Fenyes (Hoplanctria Kraatz, Lophomucter Notman, and Platandria Casey; specimens of Nosora Casiy and GenosemaNotman of the same tribe were not available for study). The presence of the mesostemal carina is considered here to be an apomorphic character state because it occurs in fewer taxa than does the non-carinate condition (in-group comparison: carinate mesosternum in 3 subgenera and absent in the remaining 4; out-group comparison: carinate mesosternum in I genus (Tinotus)2, absent in 3 others). The mesostemal characters are probably less likely to be convergent than others such as body-shape or length of the last 2The association

of the

genus Tinotus with Hoplandriini is controversial, Lohse (1974) associated

it with the tribe Aleocharinl

KLIMASZEWSKI: REVISION OF THE GENUS

Table

I.

Host records for A/eo

Host (DIPTERA)

Parasitoid

A.

ALEOCHARA

bimaculataGrav.

1I

chala in North America (only recent records are taken into consideration) Reference

subg. COPROCHARA

MUSCIDAE:

HaematobiairritansL.(hom MooreandLegner, l97l;

sp.,

fly), Musca autumnalis De Geer, Orthellia rthe llia cae sarion Meig.

Peschke and Fuldner, 1977


O

SARCOPHAGID AE: Sarcophaga sp., Ravinia querula \'/alker

A.bilineataGyll.

ANTHOMYIIDAE:Hylemyabrassicae Bouch6 MooreandLegner, 1971; (cabbage maggot), H. platura Meig. (seedcom Peschke and Fuldner, 1977 maggot), H. antiqua Meig. (onion maggot), H. floralis Fall. (tumip maggot), H. planipalpus Stein, Pegomya hyoscyami Curlis (spinach leafminer)

MUSCIDAE: Musca domesticzr L. (house fly) CALLIPHORID AE: C al I iphora e ry throc ep hala Meig.

A. verna

Say

ANTHOMYIID AE: Hylemya brassicae Bouch6 Peschke and Fuldner,

H.

(cabbage maggot), H. planipalpus Stein, ,platura Meig. (seedcom maggot), Pegomya

19771, an