Revision of the Apocephalus attophilus-group of ant-decapitating flies. (Diptera: Phoridae)

60 Pages·1997·53.3 MB·English

by  Brian V Brown

Revision of the Apocephalus attophilus-group
of Ant-decapitating Flies
(Diptera: Phoridae)
Brian V. Brown1
CONTENTS
ABSTRACT.................................... 2
INTRODUCTION 2
METHODS.... 3
SYSTEMATICS........... 3
Apocephalus Coquillett. 3
Apocephalus attophilus-group 4
Apocephalus lamellatus-subgroup 5
Apocephalus lamellatus Borgmeier 5
Apocephalus pseudocercus new species 7
Apocephalus peniculatus-subgroup 8
Apocephalus cantleyi new species 8
Apocephalus rionegrensis Borgmeier 10
Apocephalus peniculatus Borgmeier 10
Apocephalus neivat Borgmeier 11
Apocephalus longipes Borgmeier 11
Apocephalus vannus new species 12
Apocephalus octonus new species................................. 12
A. luteihalteratus-subgroup 13
Apocephalus tenuitarsus new species 13
Apocephalus spinosus new species 15
Apocephalus luteihalteratus Borgmeier. 22
Apocephalus onorei new species. 25
Apocephalusguapilensis new species 26
Apocephalus infraspinosus-subgroup 27
Apocephalus rudiculus new species 27
Apocephalus occidentalis new species 28
Apocephalus laselvaensis new species 29
Apocephalus setilohus new species 29
Apocephalus infraspinosus Borgmeier 30
Apocephalus hibbsi new species 30
Apocephalus patulus new species 31
Apocephalus cultellatus-subgroup 31
Apocephalus cultellatus Borgmeier 32
Apocephalus clavicauda new species 32
Apocephalus parallelus new species. 33
Apocephalus ancylus new species .......... 34
Apocephalus completus new species 34
Apocephalus singulus new species .... ..... ...... 35
Apocephalus securis new species ....... ............... 36
Apocephalus quadriglumis-subgroup 37
Apocephalus striatus new species 37
Apocephalus quadriglumis Borgmeier 37
1. Entomology Section, Natural History Museum of
Los Angeles County, 900 Exposition Boulevard, Los An-
geles, CA 90007.
Contributionsin Science,Number468, pp. 1-60
Natural HistoryMuseum ofLos Angeles County, 1997
Apocephalus bilobus new species 38
Apocephalus wallerae-subgroup 38
Apocephalus nigricauda new species 39
Apocephalus ritualis new species 39
Apocephalus wallerae Disney 40
Subgroup Unknown 41
Apocephalus asymmetricus new species 41
Apocephalus extraneus new species 41
Apocephalus tricuspis Borgmeier 42
Apocephalus laticauda Borgmeier 43
Apocephalus sinuosus new species 43
Apocephalus paulus Borgmeier 44
Apocephalus setitarsus new species 44
Apocephalus fads new species 45
Apocephalus attophilus Borgmeier 46
Apocephalus colombicus new species 46
Apocephalus hirsutus new species 47
Apocephalus quadratus new species 47
Apocephalus colobus new species 48
Apocephalus vibrissicauda new species 48
Apocephalus decurvus new species 49
Apocephalus lativentris new species 50
Apocephalus spinilatus new species 50
Apocephalus lunatus new species 51
Apocephalus angusticauda new species 52
Apocephalus dichromatus new species 52
Apocephalus stillatus new species 53
Apocephalus concavus new species 53
Apocephalus diffusus new species 54
Apocephalus oblongus new species 55
Key to Females 55
FUTURE STUDIES 59
ACKNOWLEDGMENTS 59
LITERATURE CITED 59
ABSTRACT.ThegenusApocephalusisdiscussed,with PleuropborinaBorgmeierconsideredajuniorsubjective
synonym(newsynonymy) andP. turgidaBorgmeiertransferredtoApocephalus(newcombination).Sixinformal
groups ofApocephalus, subgenus Apocephalus are proposed: the A. attophilus-group, A. miricauda-group A.
,
mucronatus-group,A.pergandei-group,A.feeneri-group,andA.grandipalpus-group.WithintheA.mucronatus-
group,thefemalesof“Neodohrniphora”arnaudiarefoundtobeconspecificwithApocephalushorridus,known
onlyfroma male (newsynonymy). One oftheproposed subgroupsofApocephalus,theA. attophilus-group,is
diagnosed and revised. Fifty-eight species are recognized, including the following forty-four new to science:
Apocephaluspseudocercus, cantleyi, vannus, octonus, tenuitarsus, spinosus, onorei,guapilensis, rudiculus,occi-
dentals, laselvaensis,setilobus, hibbsi,patulus, clavicauda,parallelus,ancylus, completus,singulus,securis,stria-
tus, bilobus, nigricauda, ritualis, asymmetricus, extraneus,sinuosus,setitarsus,facis,colombicus,hirsutus,quad-
ratus, colobus, vibrissicauda, decurvus, lativentris, spinilatus, lunatus, angusticauda, dichromatus,stillatus,con-
cavus, diffusus, andoblongus. ThenameA. barbicaudaBorgmeierisconsideredajuniorsubjectivesynonymof
A. rionegrensisBorgmeier (newsynonymy). Lectotypes aredesignated forA.peniculatusBorgmeier,A. luteihal-
teratus Borgmeier,A. quadriglumis Borgmeier, andA. laticauda Borgmeier. All specieswith knownlifehistories
areparasitoidsoffungus-gardeningants (Attinae).TheA.attophilus-groupishypothesizedtohaveevolvedfrom
relativelymoreprimitiveApocephalusspeciesthatparasitizeantsofthesubfamilyPonerinae.TheA.attophilus-
group isextremelypoorlysampled, andfurthercollectinginalmostanyareaoftheNewWorldtropicsislikely
to uncoveradditionalnewspecies.
INTRODUCTION ants, subgenus Mesophora, has been revised recently
The genus Apocephalus Coquillett, with the addition (Brown, 1993b, 1994, 1996a, b). The last key to the
of new species described herein, is the second largest ant parasitoids was given by Borgmeier (1971) anda
genus of phorid flies. Commonly referred to as ant- few species were described by Disney (1980a, 1981,
decapitatingflies,themajorityofspeciesinthisgroup 1982).
are parasitoids of ants (Hymenoptera: Formicidae). It has become apparent, however, these works
An exceptional group of species that does not attack have only just begun to describe the incredible di-
2 Contributionsin Science, Number468 Brown: Revision ofApocephalusattophilus-group
versity of this group. Extrapolations from Malaise CNCI Canadian National Collection of Insects, K.
trap catches have predicted that there are about Neatby Building, Agriculture Canada, Ottawa,
150 ormore species ofApocephalus atonetropical Ontario, Canada, K1A 0C6 (J. Gumming).
site alone (Brown and Feener, 1995), of which 86 EMUS Department of Biology, Utah State University,
have so far been collected. Clearly, there is much sLoong)a.n, Utah 84322-5305, U.S.A. (W.J. Han-
work to be done. INBC Institute Nacional de Biodiversidad, A.P. 22-
This paper is a first effort at defining some spe- 3100, SantoDomingo,Heredia,CostaRica (M.
cies-groups within Apocephalus, subgenus Apoce- Zumbado).
phalus andatrevisingone oftheseassemblages,the INPA Institute Nacional de Pesquisas da Amazonia,
A. attophilus-group. Estrada doAleixo, 1756, C.P. 478, 69.011 Ma-
METHODS LACM nEanutso,moBrlaozgily(SJ.ecRtaifoane,l)N.atural History Museum
of Los Angeles County, 900 Exposition Boule-
SPECIMEN TREATMENT. Most specimens were col- v(Ba.rVd.,BLrooswn)A.ngeles, California 90007, U.S.A.
lected into ethanol and critical-point-dried (Brown, MCZC Museum of Comparative Zoology, Harvard
1993a; Gordh and Hall, 1979). To observe characters of
University, Cambridge, Massachusetts 02138,
the femaleterminalia, abdomenswereclearedusinglactic U.S.A. (on indefinite loan to B.V. Brown).
acid (Gumming, 1992). MIUP Museo de Invertebrados Graham B. Fairchild,
DESCRIPTIONS. Species descriptions concentrate on Universidad de Panama, Estafeta Universitaria,
the female sex. Males ofthe A. attophilus-group (see be- Panama (D. Quintero).
low) are relatively similar, are known for only a fewspe- MUCR Museo de Insectos, Universidad de Costa Rica,
cies, and thus provide few taxonomically useful charac- San Pedro, SanJose, Costa Rica (P.S. Hanson).
ters. MUSM Museo de Historia Natural, Universidad Na-
USE OF COLOR. Color is a difficult character to use cional Mayor de San Marcos, Av. Arenales
when specimens have been subjected to differing preser- 1267,Apartado 14-0434,Lima-14,Peru(G.La-
vation and storage procedures. Those collected intoalco- mas).
hol and promptly (within 1-2 years) critical-point-dried QCAZ Quito Catholic Zoology Museum, Deparamen-
are closeto natural color. Specimens thatare killedincy- to de Biologia, Pontificia Universidad Catolica
anideandallowedtoairdrybecomedarkerincolor.Spec- del Ecuador, 12 de Octubre y Carrion, Apto.
imens stored in alcohol for long periods oftime become 2184, Quito, Ecuador (G. Onore).
auniform,bleachedlightbrowncolor.Wheneverpossible, TAMU Department of Entomology, Texas A&M Uni-
I used critical-point-dried specimens to judge color and versity,CollegeStation,Texas77843,U.S.A. (R.
otherwise notedwhen such specimenswerenotavailable. Wharton).
TERMS AND NAMES. The nomenclature ofant spe- USNM United States National Museum, Smithsonian
cies was checked against Bolton (1995). Institution,Washington,D.C.20560,U.S.A.(on
Geographicalcoordinatesarequotedasdecimaldegrees indefinite loan to B.V. Brown).
rather than degrees, minutes, and seconds (e.g., 90.5°W
ratherthan 90°30'W; Crawford, 1983). SYSTEMATICS
BARCODES. In addition to the usual insect labels re-
cordinglocalityinformation,specimenswerelabelledwith Apocephalus Coquillett
barcoded insect labels (Thompson, 1994) and data were
recorded in a database. All barcoded labels that begin Apocephalus Coquillett 1901: 501.Typespecies:A.
with the abbreviation “LACM ENT” indicate that the pergandei Coquillett, by original designation.
NaturalHistoryMuseumofLosAngelesCounty(LACM) Pseudoplastophora Schmitz 1915: 327. Type spe-
is the institution where the data are stored. Specimens cies: P. caudataria Schmitz, by monotypy. Syn-
with barcoded labels beginning “INBIO” have their data onymized by Borgmeier, 1968.
stored at LACM and the Institute Nacional de Biodiver- Pleurophorina Borgmeier 1969: 66. Type species:
sidad in Costa Rica. To make later recognition of holo- P. turgida Borgmeier, by original designation,
types easier, I list their individual barcode numbers in new synonymy
square brackets.
COLLECTING. Most of the specimens examined in Zyziphora Peterson and Robinson 1976: 119.Type
thisstudywerecollectedbyMalaisetraps(Malaise,1937; species: Z. hirtifrons Peterson and Robinson, by
Townes, 1972), including those operated by the Arthro- original designation. Synonymized by Brown,
pod Survey of La Selva Biological Station, Costa Rica 1992.
w(aALsAaSn;alLyoznegdinpor,ev1i9ou9s4l)y.bTyheBrcoatwcnhafnrdomFetehneerAL(1A9S95)tr.aps NOTES ON SYNONYMY. The genus Pleuro-
MATERIAL. Specimens belong to the following insti- phorina Borgmeierwasrecognizedasseparatefrom
tutions (codensfromArnettetah, 1993;curatornamesin Apocephalus based on the setulose anepisternum,
parentheses): swollen costa, andthe shortened frons. Thesechar-
AMNH acters are known to occur in other Apocephalus,
DepartmentofEntomology,AmericanMuseum
of Natural History, Central Park West at 79th however, and are no longer sufficient to justify a
USt.rSe.eAt.,(ND.ewGrimYaolrdki,).New York 10024-5192, gdrisatnidnicptalgpeinsu-sg.roIunppsapretciiceuslairn,tthheeLreAaCrMe csoelvleercatlioAn.,
CASC Department of Entomology, California Acade- including paratype females of A. praedator Borg-
myofSciences, Golden GatePark, San Francis- meier, that have setulae on the anepisternum.
co, California 94118,U.S.A. (P.H. Amaud,jr.). The single species affected by this action is A.
Contributionsin Science, Number468 Brown: Revision ofApocephalusattophilus-group 3
turgida (Borgmeier) (new combination). In the have a distinctive, separate sclerite on the venterof
LACM collection are some A. grandipalpis-group the ovipositor. The group includes the Nearctic Re-
specimens fromPakitza, Peru, thatappearto bethe gion species “Neodohrniphora” arnaudi, which is
females of this species. in fact the female (and junior synonym) ofApoce-
I examined many specimens of A. caudataria, phalus horridus Borgmeier (new synonymy). This
originally described in the genus Pseudoplastopho- is a relatively small group, some species of which
ra Schmitz (1915), fromIndia. Thesespecimensare attack ants of the genus Camponotus.
consistent with the current, working definition of A. pergandei-group. This grouping includes spe-
Apocephalus, butitseems exceedinglyunlikelythat cies with a distinctive lateral expansion of the ovi-
this species, as well as the Australian A. insolitus positor, as well as some distinctivecharactersofthe
Borgmeier and A. niger Malloch, are congeneric male terminalia that need further study. There are
with the New World Apocephalus. For now, how- relatively few species; some are known to attack
ever, they are retained within the genus. ants of the genus Camponotus.
DIAGNOSIS. Small flies, 0.8-4 mm long. Lower A. feeneri-group. The species of this large group
interfrontal setae close together, distantfrommargin have a distinctive group of black spinules in the
ofeye. Notopleuralcleftabsent.Anepisternalfurrow intersegmental membrane between the ovipositor
present (= anepisternum divided), anepisternum and the stylet. Disney’s illustration ofthe female of
with or without setulae, without large setae. Wing A. feeneri (Disney, 1982, fig. 1) shows this char-
vein R2+3 usually, but not always, present. Tibia of acter, although he did not remark upon it. All spe-
all legs without large, isolated setae. Dufour’smech- cies with known life histories attack ants ofthe ge-
anism present, round. Distinctive, sclerotized, para- nus Pheidole.
sitoid-type ovipositor present; anterior margin of A. grandipalpis-group. This is a large assemblage
ovipositor with distinctive darkening (secondarily that is characterized by a short ovipositor whose
lost in some species). Segments posteriorto segment dorsal sclerite is narrower than the ventral sclerite,
7withdrawn inside female segment 7 atrest. Cercus producing a distinct, rounded, lateral concavity in
ofmale usuallyelongate, thin. Larvawithraisedspi- dorsal view (e.g., Borgmeier, 1969, fig. 39). There
racular area on segment 8. are also characters of the stylet that appear to de-
SUBGROUPS WITHIN APOCEPHALUS. fine this group, but theserequire further study.Few
There are two formally recognized subgroupings hosts are known, butsome attackants ofthegenus
within this genus: subgenus Apocephalus and sub- Pheidole.
genus Mesophora. The monophyly of Mesophora NOTES ON IDENTIFICATION. Disney’srecent
has been proposed and its species revised recently key (Disney, 1994) is now the standard reference
(Brown, 1993b, 1994, 1996b). for genus-level identifications of phorid flies. Some
The status ofsubgenus Apocephalus is less clear, species ofApocephalus will notkey outproperlyin
and no statements about its monophyly have been this key, however. For example, males ofA. lamel-
made. Basically, it represents the non-Mesophora latus Borgmeier and A. pseudocercus new species
majority ofthe genus. In mypreliminarystudieson key to Megaselia at couplet 158; females ofA. ten-
this genus, I have found that there are at least five uitarsus new species and A. sinuosus new species
recognizable groupings within subgenus Apoce- key more or less to Microselia at couplet 207; fe-
phalus (there are also a number of species that do males ofA. wallerae Disney, A. ritualisnewspecies
notfall into these groups). Some ofthese mightnot and A. onorei new species key to Phalacrotophora
be monophyletic with respect to other groups, but or Megaselia at couplet 204. Disney’s key is a tre-
for convenience, I propose the following, informal mendousimprovementoverpreviouslyavailablere-
names: sources, but it is overwhelmed by the copious di-
A. attophilus-group. This is a moderate-sized versity of Neotropical phorids.
group that is characterized by a female ovipositor
with a separate apical sclerite, primitively with Apocephalus attophilus-group
heavily sclerotized, blacklateralmargins (e.g.,Figs.
76-79). Males are unknown for most species, but RECOGNITION. The genus Apocephalus is a
all known specimens have distinctly straight cerci large and complicated group, with a diverse array
(Figs. 80-81, 83-84). The few species with known of species. Understandably, subgroupings within
life historiesare parasitoidsofantsofthesubfamily such a group are difficult to diagnose; the A. atto-
Attinae. philus-group is no exception. Species ofthis group
A. miricauda-group. This relatively small group are characterized by a female ovipositor with a
probably is paraphyletic with respect to the A. at- completely separated apical portion, herein termed
tophilus-group. It is characterized by a partially the apicalsclerite,posteriortothemajorsclerotized
separated apical portion of the ovipositor, that portion. As in other Apocephalus, in most A. at-
mightgive rise to the apical sclerite. Males are sim- tophilus-group species, the ovipositor is character-
ilar to those of the A. attophilus-group. All species ized by a heavily sclerotized anteriormargin, form-
with known life historiesattackants ofthesubfam- ing an anteriorly-directed V-shaped darkening. In
ily Ponerinae. some species, this darkening, as well as mostofthe
A. mucronatus-group. The females ofthis group sclerotization of the ovipositor, has been lost (e.g.,
4 Contributionsin Science, Number468 Brown: Revision ofApocephalusattophilus-group
Fig. 41). In other species, the sclerotization of the It is not clear whether all A. attophilus-group
apical sclerite is greatly reduced or lost (Figs. 62- species belong in a single monophyletic taxon. Al-
64). Inallinstances,however, atleasttherudiments though there is no evidence of independent deri-
of one or the other is present, and a definitive di- vation of the apical sclerite, in the absence of fur-
agnosis is possible. ther supporting characters for this group, such a
A key to separate the subgenera ofApocephalus possibility must be entertained. For now, the A. at-
was presented previously (Brown, 1996b), but it tophilus-group can be considered only tentatively
must be modified to include the newly studiedspe- monophyletic.
cies in this revision: Within the A. attophilus-group, there are several
readily diagnosable, monophyletic units: A. lamel-
1 Male 2 latus-subgroup; A. peniculatus-subgroup; A. lutei-
- Female 5 halteratfws-subgroup; A. infraspinosus-subgroup;A.
2 Flagellomere 1 greatly enlarged (Brown, 1996b, cultellatus-subgroup;A. quadriglumis-subgroup;A.
figs. 3-5); lowerand usuallyupperfronto-orbital wallerae-subgroup. There are many species that do
setae absent subgenus Mesophora not fit into any of these subgroups and are treated
- Flagellomere 1 smaller, often round; fronto-or- separately at the end ofthe taxonomy section.
bital setae present 3 At this time, I cannot reconstruct the relation-
3 Wingvein CuA, short, notreachingwingmargin ships among the subgroups. Almost all of the evi-
subgenus Mesophora dence available is from adult female characters
- Wing vein CuAl reaching wing margin 4 only, which is a rich but limited source of infor-
4 Anteroventral row of setulae on hind basitarsus mation. Definitelyassociatingmaleswithfemalesis
enlarged basally (similar to Brown, 1996b, fig. difficult, and males ofthe A. attophilus-grouphave
63); flagellomere 1 pyriform; halter dark brown relativelysimilarterminalia. Furtherprogressinun-
A. Mesophora atavus Brown derstandingthe phylogenyofthisgroupmustawait
( )
- Anteroventral row of setulae on hind basitarsus newevidence frommales, immatures,ormolecules.
not noticeably enlarged; othercharactersvarious
subgenusApocephalus Apocephalus lamellatus-subgroup
5 Female terminalia consistingofproximalovipos-
itor and apicaAl.sclAeproitceep(hmoasltusofaFtitgos.ph4i-l7u9s-)gr.ou.p. conDsIisAtGiNngOSoIfSa.paFiermoafledorwsiatlh, cceorcmupsl-elixketelrombiensalbiea-
- Female terminalia( consisting of) a single structure tween the ovipositor and the apical sclerite (Fig. 4)
although some lateral portions of it may be sep- anda ventral, apparentlyopposablepairofsclerites
used for grasping the host. The ventral complex
arated (e.g., Borgmeier, 1971, fig. 150) 6
6 Abdominal glands near segment 5 dark in color consists ofa more anterior, quadrate structurethat
(Brown, 1993b, fig. 45, g) ifnot, ovipositorwith folds back over a more posterior, sclerotized trian-
ventral notch apically (Brown, 1993b, fig. 51, n) gle (Fig. 5). Both of these structures have large,
subgenus Mesophora thick setae associated with them (Figs. 6-7).
- Abdominal glands white, invisible in cleared PHYLOGENETIC RELATIONSHIPS. Based on
specimens; ovipositor lacking ventral notch 7 the complex and distinctive structure of the ovi-
7 All fronto-orbital setae present positor, the two included species are apparently
subgenusApocephalus closely related.
- Lower fronto-orbital setae absent; in some spe- CHOROLOGICAL AFFINITIES. One species is
cies, upper fronto-orbital setae also absent 8 found in Atlantic Coastal Brazil, the other in mid-
8 Ovipositor apically pointed (Brown, 1996b,.fi.gs. elevation Costa Rica. Given the paucity ofcollect-
50-58) subgenus Mesophora ing in areas between these two sites, their distri-
- Ovipositor short, broad, multidentate (Borg- butions could be much wider.
meier, 1961; figs. 59-60) It is interesting that A. lamellatus is sympatric
. . Apocephalus (Apocephalus) insignis Borgmeier wsiutbhgrAo.up)lutienihBarlatzielr,atwushe(rwehaischA.bpesloenugdsocteorcaunsotnheewr
PHYLOGENETIC RELATIONSHIPS. The A. species is sympatricwithA. luteihalteratusinCosta
attophilus-groupapparentlyhasevolvedfromwith- Rica. If the proposed separate species status forA.
in the A. miricauda-group, which possibly is a pseudocercus is correct, then this possibly repre-
paraphyletic taxon with respect to the A. attophi- sents an instance of one species undergoing differ-
lus-group. Both of these groups have darkened, entiation while a sympatric congener did not.
heavily sclerotized posterolateral margins of the
Apocephalus lamellatus Borgmeier
ovipositor that in the A. attophilus-group become
separated off as the apical sclerite. In the A. miri- (Fig. 7)
dcaaurdkae-ngerdouapr,eatshefrreomistaherarnegset oofftsheepaorvaitpioosnitoofr,thaensde ApocephaluslamellatusBorgmeier, 1926:49,fig. 7.
some species might be more closely related to the LECTOTYPE (examined). 9, BRAZIL: Petro-
A. attophilus-group than to other A. miricauda- polis, 20.xii.1924, T. Borgmeier (MZSP) [LACM
group species. ENT 093426].
Contributions in Science, Number468 Brown: Revision ofApocephalusattophilus-group 5
Figures 1-3. Female head. 1. A. laselvaensis new species. 2. A. bibbsinew species. 3. A. hirsutusnewspecies. Figures4-
5. Female terminalia, Apocepbalus pseudocercus new species. 4. Dorsal. 5. Ventral. Figures 6-7. Ventral triangle of
ovipositor. 6. Apocepbaluspseudocercusnew species. 7. Apocepbalus lamellatus Borgmeier.
6 Contributionsin Science, Number468 Brown: Revision ofApocepbalusattopbilus-group
SPECIES RECOGNITION. This species is ex- nest of the ant Acrornyrmex muticinoda Forel (=
tremely similar to A. pseudocercus, but differs by A. niger (Smith)).
the shape and setation ofthe ventraltriangle (com- PHYLOGENETIC RELATIONSHIPS. Based on
pare Figs. 6 and 7). Examination of additional fe- the extremely distinctive female terminalia, this is
males of A. lamellatus is necessary to determine the sister-species ofA. pseudocercus.
whether the separation ofthese two species can be OTHER MATERIAL EXAMINED. Paralectoty-
maintained. pe d, same data as lectotype.
DESCRIPTION. Body length 1.60-1.71 mm.
Female. Frons dark brown, broad. One pair of Apocephalus pseudocercus new species
supra-antennal setae present. Lowerinterfrontalse- (Figs. 4-6, 80-81)
tae slightly divergent. Upper interfrontal setae nor-
mal-sized. Flagellomere 1 yellow, round. Palpus SPECIES RECOGNITION. This species is ex-
small, yellow; palpal setae normal-sized, pointed. tremely similar to A. lamellatus, differing by the
Dorsumofthoraxbrown. Pleuronbrown. Legsyel- shape andsetationoftheventraltriangleoftheovi-
lowish-brown. Apex of hind femur with abrupt positor (Fig. 6).
darkening on anterior face. Anteroventral setae of DESCRIPTION. Body length 1.75-2.15 mm.
mid femur shorter than width of tibia. Hind tibia Female. Frons dark brown, broad. One pair of
without differentiated anterodorsal row ofsetulae. supra-antennal setae present. Lowerinterfrontalse-
Mid tarsomeres 1-3 with ventral setae subequal to tae slightly divergent. Upper interfrontal setae nor-
those of other legs. Apex oftarsomere 5 ofall legs mal-sized. Flagellomere 1 brown, round. Palpus
normal, blunt; tarsal claws of normal size. Costal small, yellow; palpal setae normal-sized, pointed.
setae normally spaced. Wing vein R2+3 present. Dorsum of thorax light brown. Pleuron light
Halter light brown. Abdominal tergites dark-col- brown. Legs yellowish-brown. Apex ofhind femur
ored. Venter of abdomen gray. Abdominal glands with abruptdarkeningonanteriorface. Anteroven-
ofsegment 5 dark. Lateral margin oftergite 5 lack- tral setae ofmid femur shorter than width oftibia.
ing unusually enlarged setae. Tergite 6 contiguous. Hind tibia without differentiated anterodorsalrow
Abdominal segments 5 and 6 without dense setae of setulae. Mid tarsomeres 1-3 with ventral setae
laterally. Abdominal segment 6 about as long as subequal to those of other legs. Apex oftarsomere
segment 5. Venter of segments 3-5 bare. Venter of 5 of all legs normal, blunt; tarsal claws of normal
segment 5 without sclerite. Ventral setae on seg- size. Costal setae normally spaced. Wing vein R2+3
ment 6 present. Ventral setae long, in straight line, present. Halter brown. Abdominal tergites mostly
without shorter setae medially. Segment 6 without dark, with some yellow markings. Venter ofabdo-
sternite. Venter of intersegment 6-7 bare. Dorsum men yellow. Abdominal glands of segment 5 dark.
of ovipositor with triangular sclerite and long an- Lateral margin of tergite 5 lacking unusually en-
teriorprocess. Dorsal setaeofovipositorshort,few. larged setae. Tergite 6 contiguous. Abdominal seg-
Ovipositor with large, setose, cercus-like lobes be- ments 5 and 6 without dense setae laterally. Ab-
tweentip ofovipositorandapicalsclerite (asinFig. dominal segment 6 about as long as segment 5.
4). Anterodorsal apex of ovipositor composed of Venter of segments 3-5 bare. Venter of segment 5
single, sclerotized process, sclerotized portion of withoutsclerite.Ventralsetaeonsegment6present.
ovipositor without a recognizable “V”-shaped Ventral setae short, in straightline,withoutshorter
darkening. Posteroventral apex of ovipositor with- setae medially. Segment 6 without sternite. Venter
out large medial seta, without lobes. Venter ofovi- ofintersegment6-7bare. Dorsumofovipositorrel-
positor with hinged grasping structure (as in Figs. atively evenly sclerotized, superimposed with dark
4-5). Triangular plate with apex truncate and peg- “V” and anterior projecting process. Dorsal setae
like setae extended to apex (Fig. 7). Venter of ovi- of ovipositor absent. Ovipositor with large, setose,
positorwithshort,medialspineanteriortosternite, cercus-like lobes between tip ofovipositor and api-
withoutlateralgroup ofsetae. Apical scleriteshort, calsclerite (Fig. 4). Anterodorsalapexofovipositor
approximately as long as wide, relatively parallel- composed of single, sclerotized process, sclerotized
sided, dorsoventrally depressed, symmetrical. Dor- portion of ovipositor with a recognizable “V”-
sal and ventral apices of apical sclerite without shaped darkening. Posteroventral apex of oviposi-
large setae. Apical sclerite straight in lateral view. torwithoutlargemedial seta,withoutlobes.Venter
Venter ofapical sclerite withoutmedian, digitiform of ovipositor with hinged grasping structure (Fig.
process. Stylet long. 5). Triangularplatewithapexroundedandpeg-like
Male. Palpusabsentfromspecimen.Flagellomere setae extended only halfway down sides (Fig. 6).
1 round, brown. Pulvilli offore and mid legs small. Venter of ovipositor with short, medial spine an-
I consider it questionable whether this specimen is terior to sternite, without lateral group of setae.
indeed conspecific with the female lectotype. Apical sclerite short, approximately as long as
GEOGRAPHICAL DISTRIBUTION. Known wide, relatively parallel-sided, dorsoventrally de-
from a single site in Brazil. pressed, symmetrical. Dorsal and ventral apices of
WAY OF LIFE. According to Borgmeier (1926), apical sclerite withposterolateral tuftofsetae. Api-
the specimens were collected at the entrance of a cal sclerite straight in lateral view. Venter ofapical
Contributionsin Science, Number 468 Brown: Revision ofApocephalusattophilus-group 7
sclerite without median, digitiform process. Stylet tae subparallel. Upper interfrontal setae normal-
long. sized. Flagellomere 1 brown, round to oval. Palpus
Male. Palpus small, yellow; setulaenormal-sized, small, yellow; palpal setae normal-sized, pointed.
pointed. Flagellomere 1 round, brown. Pulvilli of Dorsum of thorax light brown. Pleuron light
fore and mid legs small. Terminalia as in Figs. 80- brown. Legs yellowish-brown. Apex ofhind femur
81. with abrupt darkeningonanteriorface. Anteroven-
GEOGRAPHICAL DISTRIBUTION. Middle el- tral setae ofmid femur shorter than width oftibia.
evations in central Costa Rica. Hind tibia without differentiated anterodorsalrow
WAY OF LIFE. Unknown. It is an extremely of setulae. Mid tarsomeres 1-3 with ventral setae
common species at Zurqui de Moravia, where Ac- subequal to those of other legs. Apex oftarsomere
romyrmex coronatus is a common attine ant. 5 of all legs normal, blunt; tarsal claws of normal
PHYLOGENETIC RELATIONSHIPS. See A. la- size. Costal setae normally spaced. Wing vein R2+3
mellatus. present. Halter brown. Abdominal tergites 1-4
DERIVATION OF SPECIFIC EPITLIET. The light brown, 5-6 dark. Venter of abdomen light
name, derived from Greek, means false cercus, re- gray, segments5-6 almostblack.Abdominalglands
ferring to the unusual processes between the ovi- of segment 5 white, invisible in cleared specimens.
positor and apical sclerite. Lateral margin of tergite 5 lacking unusually en-
HOLOTYPE. 9, COSTA RICA: San Jose: Zur- larged setae. Tergite 6 contiguous. Abdominal seg-
qui de Moravia, 10.05°N, 84.02°W, vii.1991, P. ments 5 and 6 without dense setae laterally. Ab-
Hanson, Malaise trap, 1600 m (LACM) [LACM dominal segment 6 about as long as segment 5.
ENT 009389]. Venter of segments 3-5 bare. Venter of segment 5
PARATYPES. COSTA RICA: Cartago: La Can- with rectangular sclerite. Ventral setae on segment
greja, 9.8°N, 83.97°W, 2$, iv.1991, 31d, 2$, 6 present. Ventral setae short, instraightline,with-
vii.1991, 19, viii—ix.1991, 156, 19, xi.1991, 66, out shorter setae medially. Segment 6 without ster-
iii—v.1992, P. Hanson, Malaise trap, 1950 m nite. Venter of intersegment 6-7 bare. Dorsum of
(LACM); SanJose: Braulio Carrillo National Park, ovipositor largely unsclerotized, except formargin-
Id, iv.1989, 46, x-xii.1989, 19, iv-v.1990, P. al darkening. Dorsal setae of ovipositor extremely
Hanson, Malaise trap, 1000 m (LACM), Zurquide long, numerous. Ovipositor without cercus-like
Moravia, 10.05°N, 84.02°W, 2d, iii.1989, Id, lobes. Anterodorsalapexofovipositorcomposedof
vi.1989, 4d, 29, vii.1990, 15d, 19, ix-x.1990, single, sclerotized process, sclerotized portion of
6d, 89, x-xii.1990, 19, i.1991, 52d, 109, ovipositor with a recognizable “V”-shaped dark-
U.1991, 16d, 39, vi.1991, 97d, 329, vii.1991, ening. Posteroventral apex of ovipositor without
5d, 69, ix.1991, 19, xii.1991-ii.1992, 19, large medial seta, without lobes. Venter of ovipos-
hi.1992, 3d, 209, v.1992, 3d, 189, vi.1992, 29, itor without hinged structure. Venter of ovipositor
vii.1992, 59, vi.1993, Id, 39, v.1994, P. Hanson, lacking medial spine, with group ofsetae lateral to
Malaise trap, 1600 m (INBC, LACM, MCZC, sternite. Apical sclerite long, relatively parallel-sid-
MUCR, USNM). ed, dorsoventrally depressed, symmetrical. Dorsal
and ventral apices of apical sclerite without large
Apocephalus peniculatus-subgroup setae. Apical sclerite straight in lateral view. Venter
DIAGNOSIS. Dorsal setae of ovipositor long of apical sclerite without median, digitiform pro-
cess. Stylet long.
(Figs. 8-14).
PHYLOGENETIC RELATIONSHIPS. Three Male. Palpus small, yellow; setulaenormal-sized,
species,A. peniculatus, A. rionegrensis, andA. can- pointed. Flagellomere 1 round, brown. Pulvilli of
tleyi new species have a distinct lateral seta (in A. forGeEaOndGRmiAdPHleIgsCsAmLall.DISTRIBUTION. Known
cantleyi there is actually a bundle of setae) on the
only from a single site in Costa Rica.
underside of the ovipositor. Potentially, this char- WAY OF LIFE. Unknown.
actCerHcOoRulOdLgOrGouIpCtAhLeseAFtFhrIeNeITspIeEcSie.s.Most of these PHYLOGENETIC RELATIONSHIPS. Un-
species are known only from Atlantic Coastal Bra- known. Based on the lateral setae ventrally on the
ovipositor, this species might be related to A. ri-
zil.
WAY OF LIFE. All known hosts are species of oneDgEreRnIsVisATanIdONA. pOeFnicSulPaEtCuIs.FIC EPITHET. This
Acromyrmex.
species is named for Mr. Jesse J. Cantley, scientific
Apocephalus cantleyi new species illustrator for the Entomology Section of the
LACM.
(Fig. 13) HOLOTYPE. 9, COSTA RICA: San Jose: Zur-
SPECIES RECOGNITION. The females of this qui de Moravia, 10.05°N, 84.02°W, ii.1991,
species are easily identified by the long, numerous, P.Hanson, Malaise trap, 1600m (LACM) [LACM
lateral setae on the ovipositor (Fig. 13). ENT 009421].
DESCRIPTION. Body length 1.88-2.30 mm. PARATYPES. COSTA RICA: San Jose: Zurqui
Female. Frons dark brown, broad. One pair of de Moravia, 10.05°N, 84.02°W, 19, x-xii.1990,
supra-antennal setae present. Lowerinterfrontalse- Id, ii.1991, 29, iv.1991, 19, vii.1991, 56, 39,
8 Contributionsin Science, Number468 Brown: Revision ofApocephalusattophilus-gvonp
Figures 8-14. Femaleterminalia, dorsal. 8.Apocephalusoctonusnewspecies. 9.ApocepbaluspeniculatusBorgmeier. 10.
Apocephaius rionegrensisBorgmeier. 11.ApocepbalusneivaiBorgmeier. 12.Apocepbaluslongipesnewspecies. 13.Apo-
cepbalus cantleyi new species (note ovipositor is shown partially withdrawn into intersegment 6-7). 14. Apocepbalus
vannus new species. Figures 15-16. Apical sderite, dorsal. 15. Apocepbaluspeniculatus. 16. Apocepbalus rionegrensis.
Contributions in Science, Number 468 Brown: Revision ofApocephalusattopbilus-group 9
vi.1992, 3d, 59, vii.1992, 19, v.1994, P. Hanson, composed of single, sclerotized process, sclerotized
Malaise trap, 1600 m (INBC, LACM, MCZC, portion of ovipositor with a recognizable “V”-
MUCR, USNM). shaped darkening. Posteroventral apex of oviposi-
torwithoutlarge medialseta,withoutlobes.Venter
Apocephalus rionegrensis Borgmeier of ovipositor without hinged structure. Venter of
(Figs. 10, 16) ovipositor lacking medial spine, with single lateral
seta. Apical sclerite long, relatively parallel-sided,
Apocephalus rionegrensis Borgmeier, 1928: 122.
dorsoventrally depressed, symmetrical. Dorsal and
Apocephalus barhicauda Borgmeier, 1931: 218, pi.
24, fig. 24 new synonymy. ventral apices ofapical sclerite without large setae.
Apical scleritestraightinlateralview.Venterofapi-
NOTES ABOUT SYNONYMY. Borgmeier cal sclerite withoutmedian, digitiformprocess. Sty-
(1958) stated that A. barhicauda differed from A. let long.
rionegrensis bythe followingcharactersofthewing Male. Unknown.
venation: (1) costa much shorter, (2) fork (formed GEOGRAPHICAL DISTRIBUTION. Atlantic
MbyjwairnisginvgeifnrsoRm2b+a3saenodfRfo4r+k5,) sahnodrt(e4r),w(i3)ngwivneginveMin CoaWsAtaYl BOrFaziLlI.FE. The holotype of A. rionegrensis
s
more concave. was collected with Acromyrmex subterraneus var.
Withourincreasedknowledgeoftheintraspecific brunneus Forel, whereas that ofA. barhicaudawas
variation of wing venation (e.g., Disney, 1980b), collected with A. lundii (Guerin-Meneville).
little reliance is placed on these characters for rec- PHYLOGENETIC RELATIONSHIPS. See A.
ognizing species. I examined the terminalia of ho- cantleyi, above.
lotype specimens of both species and found them OTHER MATERIAL EXAMINED. BRAZIL:
to be nearly identical. One other similar species,A. Santa Catarina: BomRetiro, 19,24.i.1929,C.Pra-
peniculatus, has a relatively short apical sclerite, de [holotype ofA. barhicauda] (MZSP),NovaTeu-
and therefore I continue to recognize it as a sepa- tonia, 27.18°S, 52.38°W, 39, F. Plaumann, 300-
rate species. 500 m (MZSP, USNM).
HOLOTYPE (examined). 9, BRAZIL: Parana:
Rio Negro, 26.1°S, 49.8°W, 13.ih.1924, W. Frey Apocephalus peniculatus Borgmeier
(MZSP) [LACM ENT 029267]. (Figs. 9, 15)
SPECIES RECOGNITION. This species closely
ApocephaluspeniculatusBorgmeier, 1925: 193,fig.
resembles A. peniculatus, but has a longer apical
23, pi. VIII, fig. 37.
sclerite.
DESCRIPTION. Body length 1.50-2.10 mm. LECTOTYPE (here designated). 9, BRAZIL:Pe-
Female. Frons yellow, broad. One pair ofsupra- tropolis, 23.ii.1924, C. Prade (MZSP) [LACM
antennal setae present. Lower interfrontal setae ENT 046378].
slightly divergent. Upper interfrontal setae normal- SPECIES RECOGNITION. As discussed above,
sized. Flagellomere 1 yellow, round. Palpus small, this species is extremely similar to A. rionegrensis,
yellow; palpal setae normal-sized, pointed. Dorsum but has a relatively shorter apical sclerite (compare
of thorax light brown. Pleuron light brown. Legs Figs. 15 and 16).
yellowish-brown. Apexofhindfemurofevencolor DESCRIPTION. Body length 1.45 mm.
anteriorly. Anteroventral setaeofmidfemurshorter Female. Frons light brown, broad. One pair of
than width of tibia. Hind tibia without differenti- supra-antennal setae present. Lowerinterfrontalse-
ated anterodorsal row of setulae. Mid tarsomeres tae subparallel. Upper interfrontal setae normal-
1-3 with ventral setae subequal to those of other sized. Flagellomere 1 light brown, round. Palpus
legs. Apex oftarsomere 5 ofall legs normal, blunt; small, brown; palpal setae normal-sized, pointed.
tarsal claws of normal size. Costal setae normally Dorsum of thorax light brown. Pleuron yellow.
spaced. Wing vein R2+3 present. Halter yellow. Ab- Legs yellowish-brown. Apex ofhind femur ofeven
dominal tergites dark-colored. Venter of abdomen color anteriorly. Anteroventral setae of mid femur
gray. Abdominal glands of segment 5 white, invis- shorter than width oftibia. Hind tibia withoutdif-
ible in cleared specimens. Lateral margin oftergite ferentiated anterodorsal row of setulae. Mid tar-
5 lacking unusually enlarged setae. Tergite 6 con- someres 1-3 with ventral setae subequal to those
tiguous. Abdominal segments 5 and 6 without of other legs. Apex of tarsomere 5 of all legs nor-
dense setae laterally. Abdominal segment 6 about mal, blunt; tarsal claws ofnormal size. Costalsetae
as long as segment 5. Venter ofsegments 3-5 bare. normally spaced. Wing vein R2+3 present. Halter
Venter of segment 5 without sclerite. Ventral setae brown. Abdominal tergites dark-colored. Venterof
onsegment6 present. Ventral setae long,instraight abdomen yellow. Abdominal glands of segment 5
line, without shorter setae medially. Segment 6 white, invisible in cleared specimens. Lateral mar-
without sternite. Venter of intersegment 6-7 bare. gin of tergite 5 lacking unusually enlarged setae.
Dorsum of ovipositor with triangular sclerite and Tergite 6 contiguous. Abdominal segments 5 and 6
long anteriorprocess. Dorsalsetae ofovipositorex- without dense setae laterally. Abdominal segment6
tremely long, numerous. Ovipositor without cer- about as long as segment 5. Venter ofsegments 3-
cus-like lobes. Anterodorsal apex of ovipositor 5 bare. Venter of segment 5 without sclerite. Ven-
10 Contributionsin Science, Number 468 Brown: Revision ofApocephalusattophilus-group