.AJft

Number 507
8 June 2006

Contributions

IN Science

Revision of the Melaloncha ungulata-
Group of Bee-Killing Elies
(Diptera: Phoridae)

Brian V. Brown and Giar-Ann Kung

j^atural

A /rHistory
Museum ^

of Los Angeles County

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Museum ol
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John Heyning, Deputy Director
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John M. Harris, Committee Chairman
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Natural History Museum
OF Los Angeles County
900 Exposition Boulevard
Los Angeles, Calieornia 90007

Printed at Allen Press, Inc., Lawrence, Kansas
ISSN 0459-8113

Revision of the Melaloncha ungulata-Grou? of
Bee-Kieeing Flies (Diptera: Phoridae)

Brian V. Brown^ and Giar-Ann Kung^

ABSTRACT. The Melaloncha ungulata-group is a hypothesized monophyletic lineage based on the
presence of a 3-pronged dorsal process, here called the trident, of intersegment 7-8 of the female
abdomen. The group is revised and, exclusive of the M. imgulata-senes, consists of 3 described species,
M. flava Borgmeier, M. nigrifrons Borgmeier, and M. nigrita Borgmeier, plus the following 17 new to
science: M. a coma, M. adusta, M. Candida, M. caligula, M. claviapex, M. cristula, M. culmena, M.
debilis, M. diastata, M. fuscipalpis, M. juxta, M. platypoda, M. setitibialis, M. spina, M. tambopatensis,
M. titparroensis, and M. ustulata. A further described species, M. iingidata Borgmeier, and the following
8 new to science are classified in the M. ungnlata-series: M. atrilingula, M. borgmeieri, M. castanea, M.
curtibrachia, M. inicita, M. laticlava, M. pegmata, M. strigosa, and M. trita. Phylogenetic relationships
within this group are partially resolved, with M. acoma, M. adusta, M. Candida, and M. nigrita
considered closely related, the M. ungulata-senes forming a second monophyletic group, and the
majority of the rest of the species belonging, although with unresolved relationships, within a larger
monophyletic group containing the M. ungulata-stnes. Hosts and behavior of several species of these
Neotropical bee-killing flies are recorded.

INTRODUCTION

The genus Melaloncha Brues is a group of
colorful, bee-parasitizing phorid flies (Fig. 1)
found almost exclusively in the New World
tropics. Before recent attention this was a relative-
ly small group, with 32 described species (Borg-
meier, 1968, 1971b), but recent revisions (Brown,
2004a, 2004b, 2005; Gonzalez and Brown, 2004)
have described many new forms, such that the
final number of species is now expected to be
between 200 and 300.

Besides their diversity, this group is of interest
for their parasitoid habits and the interesting
behaviors of adult females as they attack their
hosts. A greater knowledge of the species in this
genus, both in the laboratory and in the field, will
allow us to better understand the history and
function of their diverse body forms and struc-
tures.

In his first paper about Melaloncha, Brown
(2004b) hypothesized some monophyletic groups,
including 2 subgenera, Udaniochiras Enderlein
and Melaloncha s.s. Within Melaloncha s.s., there
is a distinctive group of species that we here term
the M. ungulata-group, with a 3-pronged struc-
ture at intersegment 7-8 (Fig. 5). This paper
describes the known species of the group and
proposes some preliminary ideas about their
relationships. The phylogeny of this group is also
being studied in a combined morphological and
molecular project by B. Brown and P. Smith (in
preparation).

1-2. Entomology Section, Natural History Museum
of Los Angeles County, 900 Exposition Boulevard, Los
Angeles, California 90007, USA. E-mail: Tbrown@
nhm.org; ^gkung@nhm.org

MATERIALS AND METHODS

This revision is based almost exclusively on female
specimens. Males of Melaloncha are extremely similar
to each other (at least within subgroups), are often highly
divergent in structure and color from females, and show
too few differences to be useful in characterizing species.
Only the few known males that were collected in copula
with females are described here. If mating pairs could be
collected reliably, there would be the possibility of
making further progress in the taxonomy of males, but
unfortunately, in our experience, it is extremely rare that
such pairs are encountered.

All specimens have a bar-coded insect label, and their
data are stored at the LACM. Bar-code data for
holotypes is presented in square brackets for their easy
identification.

Material is deposited in the following collections (for
more details on collections, see Arnett et ah, 1993):

AMNH Department of Entomology Collection, Amer-
ican Museum of Natural History, New York,
New York, USA

CBEC Coleccibn Boliviana de Eauna, La Paz, Bolivia
CMNH Section of Invertebrate Zoology, Carnegie
Museum of Natural History, Pittsburgh,
Pennsylvania, USA

EMUS Department of Biology, Utah State University,
Logan, Utah, USA

EMNH Insect Collection, Eield Museum of Natural
History, Chicago, Illinois, USA
INBC Instituto Nacional de Biodiversidad, Heredia,
Costa Rica

INPA Insituto Nacional de Pesquisas da Amazonia,
Manaus, Brazil

LACM Entomology Section, Natural History Muse-
um of Los Angeles County, Los Angeles,
California, USA

MACN Division Entomologia, Museo Argentino
de Ciencias Naturales, Buenos Aires, Argen-
tina

Contributions in Science, Number 507, pp. 1-31
Natural History Museum of Los Angeles County, 2006

2 ■ Contributions in Science, Number 507

Brown and Kung: Revision of M. ungulata-%vo\i^

Figure 1 Melaloncha acoma n. sp. (right) attacking Plebia aff. jatiformis (Cockerell) in the field at Zona Protectora El
Rodeo, Costa Rica (photo by B. Brown)

MCZC Entomology Department, Museum of Com-
parative Zoology, Harvard University, Cam-
bridge, Massachusetts, USA

MIUP Museo de Invertebrados, Universidad de

Panama, Panama

MPEG Departamento de Entomologia, Museu Para-
ense Emilio Goeldi, Belem, Brazil
MUCR Museo de Insectos, Universidad de Costa

Rica, San Jose, Costa Rica

MUSM Museo de Historia Natural, Universidad

Nacional Mayor de San Marcos, Lima, Peru
MZSP Museu de Zoologia, Universidade de Sao

Paulo, Sao Paulo, Brazil

NHRS Naturhistoriska Riksmuseet, Sektionen fur

Entomologi, Stockholm, Sweden
QCAZ Quito Catholic Zoology Museum, Departa-
mento de Biologia, Quito, Ecuador
UNCB Museo de Historia Natural, Instituto de

Ciencias Naturales, Universidad Nacional de
Colombia, Bogota, Colombia
USNM United States National Museum, Smithsonian
Institution, Washington, DC, USA

Color images of specimens of each species are posted at
B. Brown’s Web site, currently at www.phorid.net/
phoridae/Melaloncha/products.html. These images should
be consulted in conjunction with the key herein when
identifying specimens.

Structural terms are those of the Manual of Nearctic
Diptera (McAlpine, 1981), except for some new terms
for Melaloncha derived by Brown (2004b). Erontal
width was measured at midlength of the frons.

Most specimens in this revision were collected by
spraying a mixture of honey and water to attract bee
hosts, which in turn attracted parasitic phorid flies. In
earlier collections we sprayed honey on a white sheet,
termed a bee screen (Brown, 2001), but later collections
were made directly from honey-sprayed undergrowth
vegetation.

SYSTEMATICS

PHYLOGENETIC ANALYSIS

There are relatively few structural characters that
we have identified for hypothesizing relationships

within this group. Most are associated with the
female intersegment 7-8 and foretarsomeres.
These characters are listed below, with the
presumed primitive state labeled (0) and the
derived states labeled (1).

1. Intersegment 7-8 of female abdomen with
sclerotized apex forming a simple elongate
process (0), formed into a 3-pronged pro-
cess, and herein referred to as the trident
(Figs. 4-43) (1). The structure of the
trident is a dorsomedial process and 2
more ventral lateral arms. It is situated
dorsally on the intersegment (Fig. 5) and
apparently is used to help guide the female
terminalia for deposition of eggs in the
host.

2. Foretarsomere 1 parallel-sided (0), broad-
ened at base (Fig. 44) (1).

3. Lateral setae of foretarsomeres small, un-
modified (0), lateral setae, especially those
of tarsomeres 2-4, modified, with inner
(anterior) seta elongate and outer (posterior)
seta thickened, curved (Figs. 51, 54, 59)
(1).

4. Venter of foretarsomeres 1-2 without differ-
entiated setae (0), venter of foretarsomeres
1-2 with thickened setae along posterior
margin (Figs. 45, 50, 53, 58) (1). The form
of the differentiated setae varies among
groups, and the setae are more differentiated
in some than in others.

5. Dorsomedial process of trident not enlarged
or laterally flattened (0), dorsomedial pro-
cess enlarged, laterally flattened (Figs. 15-
23) (1).

6. Dorsomedial process of trident, although
enlarged, not with large dorsoventral ex-
pansion (0), dorsomedial process greatly
dorsoventrally expanded (Figs. 18-23)
(1).

7. Foretarsal claws subequal to somewhat un-
equal (Fig. 47) (0), foretarsal claws greatly

Contributions in Science, Number 507

unequal, with posterior (outer) claw 2.5 X
length of anterior (inner) claw (Figs. 54-55,
57) (1).

8. Dorsum of foretarsomere 5 without large,
differentiated seta (0), dorsum of foretar-
somere 5 with large apical seta (Fig. 56) (1).

9. Apex of dorsomedial process without dark
margins (0), apex of dorsomedial process
with dark lateral margins (Figs. 41-43) (1).

The characters above are all consistent within
groups and thus represent a conservative sam-
pling of attributes used for analysis. Based on
these characters, phylogenetic trees (Figs. 2-3)
were manually constructed, and we have derived
the following preliminary classification for this
group:

M. iingulata-gvou^

M. flava Borgmeier
M. nigri ta-suhgroup
M. acoma new species
M. adusta new species
M. Candida new species
M. nigrita Borgmeier
M. imgulata-suhgrovip
M. nigrifrons Borgmeier
M. platypoda new species
M. i/77gw/c//c/-infragroup
M. diastata new species
M. fuscipalpis new species
M. juxta new species
M. ustidata new species
M. ccdigida-SQnQS
M. claviapex new species
M. debilis new species
M. setitibialis new species
M. caligida-suhsQYiQS
M. Caligula new species
M. cristula new species
M. culmena new species
M. spina new species
M. tambopatensis new species
M. tiiparroensis new species
M. imgulata-SQriQS

M. genitcdis Borgmeier, incertae sedis
M. glabrifrons Borgmeier, incertae sedis
M. borgmeieri new species
M. curtibraehia new species
M. 7//7g7//c//‘r/-subseries
M. eastanea new species
M. latielava new species
M. strigosa new species
M. trita new species
M. ungulata Borgmeier
M. pegmata infraseries
M. atrilingula new species
M. inicua new species
M. pegmata new species

Brown and Kung: Revision of M. ungulata-groxxp ■ 3

TAXONOMY

Melaloncha Brues, 1904
Melaloncha, subgenus Melaloncha Brues, 1904
Melaloncha ungulata-gvoup

DIAGNOSIS. Wing vein R2+3 absent. Coxal
setae black (except yellow in M. platypoda).
Claws of forelegs slightly bifid (Fig. 47), except
those of M. ungulata-sents, which are otherwise
modified (Fig. 55); those of mid- and hind legs not
bifid. Hind tibia without anterodorsal rows of
setae (although with irregular rows of longitudi-
nal setal palisades). Abdominal tergites without
long setae. Ovipositor (segment 7) relatively
uniform, short, slightly upturned, setose (except
bare dorsally in M. acoma and M. adusta). Venter
of ovipositor without cercuslike lobes. Apex
of intersegment 7-8 with elaborate, 3-pronged
process herein termed the trident, consisting of
single dorsomedial process and 2 lateral arms
(Fig. 4).

INCLUDED SPECIES. Melaloncha flava, plus
species of the M. nigrita-suhgroup and the M.
ungulata-suhgroup.

NOTE ON SPECIES KNOWN ONLY
EROM MALES. Females of Melaloncha have
a number of taxonomically useful characters,
found mostly in the head, legs, and ovipositor.
Males, on the other hand, are extremely similar in
structure, as well as being highly divergent in
structure and color from females. Therefore, it is
lamentable that Borgmeier described several
species of Melaloncha from male specimens only
(he later realized the problems he had created;
Borgmeier, 1971a:125). Species based on males
are extremely problematic and cannot be
matched with females with any confidence,
although Borgmeier tried to do so on several
occasions. Additionally, it is not possible to
segregate males that belong in the M. ungulata-
group from those of other subgroupings of
the subgenus Melaloncha (except those of the
M. furcata-group, which are highly distinctive).
We therefore treat the male-only species as
unknowns and describe new species based on
females, even though some synonyms possibly
will be produced.

The following male-only species could poten-
tially fall within the group treated herein: M.
genitalis Borgmeier, M. glabrifrons Borgmeier,
M. luteipleura Borgmeier, and M. zikani Borg-
meier. Of these, M. genitalis and M. glabrifrons
>have been associated with the M. ungidata-
subgroup and are discussed later in this paper.

Melaloncha flava Borgmeier

(Fig. 4)

Melaloncha flava Borgmeier, 1959:182-184, fig.
77.

4 ■ Contributions in Science, Number 507

HOLOTYPE. ?, BRAZIL: Santa Catarina:
Nova Teutonia, 27.18°S, 52.38°W, v.1952, F.
Plaumann (MZSP; examined but not bar-coded).

RECOGNITION. This species is easily recog-
nized by the overall yellow color and the
abdominal tergites that lack any black markings.

Males are similar in color to females, and were
keyed by Borgmeier (1971a).

DESCRIPTION. Female. Body length 1.3-
1.7 mm. Frons yellow, except ocellar triangle
black; with fine, reticulate sculpturing. Mean
frontal width 0.44 head width. Flagellomere 1
yellow. Palpus yellow, palpal setae yellow. Posto-
cular setae black dorsally, yellow ventrally. Genal
setae yellow. Scutum brownish-yellow. Pleuron
brownish-yellow. Legs brownish-yellow. Venter of
foretibia lacking enlarged setae. Foretarsomeres
relatively elongate, narrow. Posteroventral margin
of tarsomeres 1-2 without enlarged setae. Large,
lateral setae of foretarsomeres relatively small,
inconspicuous. Posterior (outer) tarsal claw sub-
equal to anterior claw. Mean costal length 0.45
wing length, range 0.43-0.47. Halter yellow.
Abdominal tergites completely yellow. Ovipositor
yellow basally, otherwise dark brown, with short
setae, slightly dorsally curved. Lateral arms of
trident narrow. Dorsomedial process of trident
subequal in length to lateral arms.

Male. Frons yellow, except ocellar triangle
black. Flagellomere 1 yellow. Palpus yellow; palpal
setae black, well developed. Postocular and genal
setae black. Scutum and pleuron yellowish-brown.
Legs yellowish-brown, except foretarsomeres dark
brown. Foretarsomere 1 about twice as long as
broad; other foretarsomeres about as long as
broad; pulvilli enlarged, yellow. Abdominal ter-
gites yellowish-brown. Venter of abdomen yellow.
Cercus apparently dark brown, although the entire
body is darkened from poor preservation. Surstylus
with short apical setae.

HOST. Borgmeier (1959) recorded Tetragonisca
angustula fiebrigi Schwarz (as Tetragona [Tetra-
gonisca] jaty fiebrigi) as the host of this species.

GEOGRAPHICAL DISTRIBUTION. South-
eastern Brazil.

MATERIAL EXAMINED. BRAZIL: Santa
Catarina: Nova Teutonia, 27.18°S, 52.38°W,
1(5, 89, v.1952, 19, xi.l959, 29, vii.1972, F.
Plaumann (FMNH, LACM, MCZC, MZSP).

Melaloncha ^z/gr/Y<3-subgroup

DIAGNOSIS. Foretarsomere 1 broad, enlarged
subbasally, elongate (Fig. 44).

INCLUDED SPECIES. Melaloncha acoma, M.
adiista, M. Candida, and M. nigrita.

Melaloncha acoma n. sp.

(Figs. 1, 5, 44)

RECOGNITION. The females of this species
are easily recognized by the yellow color, bare

Brown and Kung: Revision of M. ungulata-grovrp

dorsum of the ovipositor, and enlarged tarso-
mere 1.

Three male specimens were collected in the field
while they were copulating with females. These
males were extremely different in body color and
form from the associated females. Males key to
M. glabrifrons in couplet 16 (the last couplet) in
Borgmeier’s (1971a) key to males of Melaloncha,
but the surstyli of M. glabrifrons are much longer
than those of M. acoma, and it is highly unlikely
that they are the same species.

DESCRIPTION. Female. Body length 1.7-
2.5 mm. Frons yellow, except ocellar triangle
black; with fine, reticulate sculpturing. Mean
frontal width 0.45 head width. Flagellomere 1
white basally, yellow apically. Palpus yellow,
palpal setae yellow, thin. Postocular setae black
dorsally, yellow ventrally. Genal setae yellow.
Scutum anteriorly light brown, posteriorly dark
brown. Pleuron brownish-yellow. Legs yellow.
Venter of foretibia lacking enlarged setae. Fore-
tarsomeres 2-5 relatively elongate, narrow; tar-
somere 1 greatly elongate, somewhat widened.
Posteroventral margin of tarsomeres 1-2 without
enlarged setae. Large, lateral setae of foretar-
someres relatively small, inconspicuous. Posterior
(outer) tarsal claw slightly longer than anterior
claw and directed more outwards. Mean costal
length 0.49 wing length, range 0.44-0.53. Halter
yellow. Abdominal tergites anteriorly yellow,
with black posterior margins. Ovipositor yellow
basally, otherwise dark brown to black, bare
dorsally, with short lateral and ventral setae,
slightly dorsally curved. Lateral arms of trident
broad at apex. Dorsomedial process of trident
subequal in length to lateral arms.

Male. Frons orange to darker brown medially,
except ocellar triangle black. Flagellomere 1
basally white, apically brown. Palpus whitish-
yellow; palpal setae black, well developed. Posto-
cular and genal setae black. Scutum anteriorly
light brown, posteriorly dark brown, with well-
defined anteromedial projection of dark color.
Pleuron dark brown, except proepisternum and
anterior portion of anepisternum light brown.
Legs yellowish-brown, except forefemur and apex
of hind femur dark brown. Foretarsomere 1 twice
as long as broad, yellowish-brown; other fore-
tarsomeres about as long as broad and dark
brown; pulvilli enlarged, brown. Abdominal
tergites black, anteriorly with silvery pollinosity.
Basal one-third of cercus brown, apical two-thirds
yellow. Surstylus with short apical setae.

HOST. We have observed females attacking
a wide variety of bees, including Plebeia frontalis
(Friese), P. aff. jatiformis Cockerell (Fig. 1),
Plebeia spp., Tetragona clavipes (F.), Trigona
corvina Cockerell, T. fulviventris Guerin, and T.
silvestriana Vachal. Flies approach bees on foot,
often from behind, and gradually move around to
face the front of the bee at a 45-degree angle. At
all times during an attack the abdomen is curled

Contributions in Science, Number 507

forward under the body (as in Fig. 1). After
attaining the proper position, the fly then dashes
in and attacks the bee, possibly at the mandibular
suture on the head.

GEOGRAPHICAL DISTRIBUTION. This is
one of the most widely distributed species of
Melaloncha^ being found at mid- to low eleva-
tions from Mexico to Argentina.

DERIVATION OE SPECIFIC EPITHET. From
Greek akomos for “bald,” referring to the bare
dorsum of the ovipositor.

PHYLOGENETIC RELATIONSHIPS. This
species is possibly most closely related to M.
adusta, as both have an unusual dorsally bare
ovipositor.

HOLOTYPE. ?, COSTA RICA: Heredia: La
Selva Biological Station, 10.43°N, 84.02°W,
24.vii.2000, G. Rung, bee screen #1 [LACM
ENT 152935] (LACM).

PARATYPES. ARGENTINA: Misiones: Lor-
eto, Ruinas Jesuiticas, 27.77°S, 57.28°W, 1?,
24.viii.2000, P. Fidalgo, yellow pan trap
(LACM). BOLIVIA: Beni: 5 km N Rurrenabaque,
14.43°S, 67.5UW, 1?, 23.iv.2003, B. Brown, S.
Marcotte, E. Zumbado, honey spray (LACM), La
Paz: Coroico, Hotel Don Quixote, 16.19°S,
67.72°W, 1$, 6.iv.2001, B. Brown, G. Rung,
Phoenix palm flowers, 1750 m (LACM). CO-
LOMBIA: Choco: PNN Utria, Sendero Boroboro,
6.03°N, 77.32°W, 1?, 4.vii.2000, honey-sprayed
leaves, 2?, 5.vii.2000, B. Brown, G. Rung, bee
screen (LACM, UNCB); Magdalena: PNN Tayr-
ona, Zamo, 11.33°N, 74.03°W, 1$, 29.ix-
17.X.2000, R. Henriquez, Malaise trap, CAP-
793, 50 m (UNCB). COSTA RICA: Heredia: La
Selva Biological Station, 10.43°N, 84.02°W, 15?,
15.ii-2.iii.l993, ALAS, Malaise trap, M/06/021,
M/07/022, M/08/023, M/12/027, M/13/028, M/
15/030, 1(5, 15.iii.l993, ALAS, Malaise trap M/
03/034, 1 ?, I.iv.l993, ALAS, Malaise trap M/05/
052, 1?, 14.vi.l993, ALAS, Malaise trap M/07/
129, 3?, 15.vii.l993, ALAS, Malaise trap, M/04/
155, 1$, 3.viii.l993, ALAS, Malaise trap M/13/
175, 1?, 15.iii.l994, ALAS, Malaise trap M/09/
379, 1?, 30.vi.l995, ALAS, Malaise trap M/08/
393, 2$, l.viii.l995, ALAS, Malaise trap M/08/
417, 1$, 16.viii.l995, ALAS, Malaise trap M/12/
433, 1?, 14.ix.l995, ALAS, Malaise trap M/03/
449, 1?, 16.X.1995, ALAS, Malaise trap M/04/
474, 1?, 30.ix.l998, ALAS, light trap L/09/478
(INBC, LACM), 1?, 21.vii.2000, G. Rung, bee
screen #6 (LACM); Puntarenas: Las Alturas,
8.95°N, 82.83°W, 1?, xi.l991, P. Hanson,
Malaise trap, 1500 m (LACM), 24 km W Piedras
Blancas, 8.77°N, 83.40°W, 1$, x.l991, P. Han-
son, Malaise trap, 200 m (LACM), 3 km SW
Rincon, 8.68°N, 83.48°W, 1?, iii-iv.l991, P.
Hanson, Malaise trap, 10 m (LACM), 5 km SW
Rincon, Tropical Youth Center, 8.70°N,
83.51°W, 1?, 31.v-7.vi.l998, B. Brown, V.
Berezovskiy, Malaise trap #5, 40 m (LACM),
1$, 10.viii.2001, B. Brown, V. Berezovskiy, E.

Brown and Kung: Revision of M. ungulata-gcoup ■ 5

Zumbado, honey-sprayed leaves (LACM), Wilson
Botanical Garden, Las Cruces, 8.79°N, 82.95°W,
2?, 12. vi. 1998, B. Brown, V. Berezovskiy,
flowers of Syagrus coronata (palm) (LACM),
1?, 18.V.2000, G. Rung, flowers of Syagrus
coronata^ 1?, 19.V.2000, G. Rung, fruits of
Syagrus coronata (LACM); San Jose: University
for Peace, 9.92°N, 84.28°W, 1?, 2.viii.2001, G.
Rung, honey-sprayed leaves (LACM), Zona Pro-
tectora El Rodeo, 9.91°N, 84.28°W, 5?, 1-
5.viii.2001, 5?, 3-8. viii. 2001, B. Brown, V.
Berezovskiy, G. Rung, Malaise trap (LACM),
1(5, 1?, 5.viii.2001, 2(5, 2?, 6.viii.2001, B.
Brown, G. Rung, in copula on leaves (LACM),
2?, l.viii.2001, 17?, 2.viii.2001, 14?,

3. viii.2001, 3?, 4.viii.2001, 3?, 5.viii.2001,
36?, 8. viii. 2001, B. Brown, G. Rung, honey-
sprayed leaves (LACM, MUCR, MCZC, NHRS,
USNM). MEXICO: Veracruz: 33 km NE Cate-
maco, Los Tuxtlas Biological Station, 6?, l.vii-
1. viii. 1983, S. & J. Peck, flight intercept trap, rain
forest (LACM). PERU: Madre de Dios: Tambo-
pata Research Center, 13.14°S, 69.6UW, 2?,
19.vii.2001, 1?, 20.vii.2001, 3?, 23.vii.2001,
1?, 24.vii.2001, B. Brown, G. Rung, honey-
sprayed leaves (LACM, MUSM), 2?, 19.vii.2001,
B. Brown, G. Rung, attacking host bee [Tetragona
clavipes (F.)] (LACM), 1?, 20-23.vii.2001, B.
Brown, G. Rung, Malaise trap #5 (LACM); San
Martin: Estacion “Biodiversidad”, 19 km NE
Tarapoto, 6.46°S, 76.29°W, 1?, 6.vii.2004, B.
Brown, E. Corona, honey-spray, 950 m (LACM),
San Antonio de Cumbaza, 6.40°S, 67.41°W, 1?,

4. vii.2004, 1?, 5.vii.2004, B. Brown, E. Corona,
honey-spray, 400 m (LACM, MUSM).

Melaloncha adusta n. sp.

(Fig. 6)

RECOGNITION. This species is similar to M.
acoma but is much darker in color and has the
trident with longer lateral arms.

DESCRIPTION. Eemale. Body length 1.8-
2.4 mm. Frons orange, darkening to brown
medially, ocellar triangle black; with fine, re-
ticulate sculpturing. Mean frontal width 0.4 head
width. Flagellomere 1 orange. Palpus yellow,
palpal setae black. Postocular setae black. Genal
setae black. Scutum anteriorly light brown,
posteriorly dark brown, with diffuse anteromedial
projection of dark color; scutellum apically
light brown. Pleuron light brown, with darker
markings. Legs brownish-yellow (hind femur
darker and with dark brown apex). Venter of
foretibia lacking enlarged setae. Foretarsomeres
2-5 relatively elongate, narrow; tarsomere 1
greatly elongate, somewhat widened. Posteroven-
tral margin of tarsomeres 1-2 without enlarged
setae. Large, lateral setae of foretarsomeres
relatively small, inconspicuous. Posterior (outer)
tarsal claw slightly longer than anterior claw
and directed more outwards. Mean costal length

6 ■ Contributions in Science, Number 507

0.49 wing length, range 0.47-0.53. Halter yellow.
Abdominal tergites black to blackish-brown, with
dull silver pollinosity and lateral triangular areas
of light brown. Ovipositor yellow basally, other-
wise dark brown, bare dorsally, with short lateral
and ventral setae, slightly dorsally curved. Lateral
arms of trident narrow. Dorsomedial process of
trident slender, shorter than lateral arms.

Male. Unknown.

HOST. We observed females of this species at
Las Cruces attacking Apis mellifera, Paratrigona
ornaticeps (Schwarz), and Partamona cupira (F.
Smith).

GEOGRAPHICAL DISTRIBUTION. Costa
Rica, Panama.

DERIVATION OE SPECIEIC EPITHET. Latin
for dark, referring to the dark color of this species
relative to the similar M. acoma.

PHYLOGENETIC RELATIONSHIPS. This is
possibly the sister-species of M. acoma, sharing
the dorsally bare ovipositor.

It is interesting to note that the dark coloration
of females of this species is similar to that of males
of M. acoma, whose females are bright yellow.
Males of M. adusta are not yet recognized.

HOLOTYPE. ?, COSTA RICA: Puntarenas:
Wilson Botanical Garden, Las Cruces, 8.79°N,
82.95°W, 14. vi. 1998, V. Berezovskiy, flowers of
Syagrus coronata palm [LACM ENT 116411]
(LACM).

PARATYPES. COSTA RICA: Heredia: Chila-
mate, 10.45°N, 84.08°W, 1$, v.1989, P. Hanson,
Malaise trap, 75 m (LACM), La Selva Biological
Station, 10.43°N, 84.02°W, 1$, 4.X.1993, ALAS,
Malaise trap M/15/223 (INBC); Puntarenas:
Wilson Botanical Garden, Las Cruces, 8.79°N,
82.95°W, 1?, 12.vi.l998, 10?, 13.vi.l998, 5?,
14.vi.l998, 5?, 16.V1.1998, B. Brown, V. Bere-
zovskiy, flowers of Syagrus coronata palm
(LACM, MCZC, MUCR, NHRS, USNM), 4?,
17.V.2000, G. Kung, bee screen (LACM), 1?,
16.V.2000, 4?, 17.V.2000, 1?, 18.V.2000, G.
Kung, on Syagrus coronata flowers (INBC,
LACM). PANAMA: San Bias: Nusangandi Re-
serve, 9.33°N, 79.00°W, 1?, 16-23.iv.l994, J.
Pickering, Malaise trap #2862 (MIUP).

Melaloncha Candida n. sp.

(Fig. 7)

RECOGNITION. Females of this species can
be recognized by the elongate, relatively straight
dorsomedial process of the trident, the dark
brown color, and the white legs, palpus, and
flagellomere.

Males key to M. zikani in the latest key to
males (Borgmeier, 1971a); however, the sculptur-
ing of the frons of M. Candida is strongly
reticulated, with distinctive, raised lines (as in
Figs. 48-49), whereas in M. zikani the frons is
smoother (similar to Fig. 52). Unfortunately, there
are so few males of small, dark Melaloncha, and

Brown and Kung: Revision of M. ungulata-gr ouip

there are so few distinguishing characters of
males, that identification of M. zikani males
cannot be verified at this time.

DESCRIPTION. Eemale. Body length 1.5-
1.8 mm. Frons dark brown, lighter at dorsolateral
and anterior extremes (setal bases dark brown),
with reticulate sculpturing, appearing almost like
leather grain. Mean frontal width 0.47 head width.
Flagellomere 1 white. Palpus white, palpal setae
yellow. Postocular setae black dorsally, yellow
ventrally. Genal setae yellow. Scutum dark brown
(anterolateral corner light brown). Pleuron light
brown, with darker markings (ventrally white).
Legs white, apex of hind femur dark brown. Venter
of foretibia lacking enlarged setae. Foretarsomeres
relatively elongate, narrow. Posteroventral margin
of tarsomeres 1-2 without enlarged setae. Large,
lateral setae of foretarsomeres relatively small,
inconspicuous. Posterior (outer) tarsal claw sub-
equal to anterior claw. Mean costal length 0.47
wing length, range 0.46-0.49. Halter whitish-
yellow. Abdominal tergites black, with dull silver
pollinosity. Ovipositor brown basally, darker
brown apically, setose, relatively straight. Lateral
arms of trident narrow. Dorsomedial process of
trident slender, apically pointed, longer than
lateral arms, slightly expanded apically.

Male. Frons dark brown, except orange ventral-
ly and dorsolaterally. Flagellomere 1 yellow.
Palpus white; palpal setae black, well developed.
Postocular and genal setae black. Scutum blackish-
brown. Pleuron dark brown, except proepisternum
and anterior portion of anepisternum light brown.
Legs yellowish-brown, except foretarsomeres 4-5
and apex of hind femur dark brown; coxae
whitish-yellow. Foretarsomere 1 about twice as
long as broad, other tarsomeres about as long as
broad; pulvilli enlarged, white. Abdominal tergites
black, anteriorly with silvery pollinosity. Basal
one-third of cercus brown, apical two-thirds white.
Surstylus with short apical setae.

HOST. One specimen was attracted to an
aggregation of Trigonisca atomaria (Cockerell),
whereas others were collected attacking Trigona
setentrionalis Almeida on Attalea palm flowers.

GEOGRAPHICAL DISTRIBUTION. Amazon
Basin.

DERIVATION OE SPECIEIC EPITHET. Latin
Candidas for “white,” referring to the color of the
ventral part of the body.

HOLOTYPE. ?, BRAZIL: Para: Cai^ara,
1.78°S, 51.43°W, 7.X.2001, B. Brown, A. Rodri-
gues, flowers of Attalea maripa palm [LACM
ENT 075922] (MPEG).

PARATYPES. BRAZIL: Amazonas: Manaus,
Reserva Ducke, 3.13°S, 60.02°W, 1?, 8-

15.iv.l992, J. Vidal, Arm-Cola. 1-B-lOm (INPA);
Para: 2?, same data as holotype, 2$, 1?, same
data except 6.x. 2001 (LACM, MPEG). COLOM-
BIA: Amazonas: PNN Amacayacu, 3.82°S,

70.26°W, 1?, ll.iii.2000, B. Brown, G. Kung,
attacking Trigonisca atomaria (UNCB), 1?,

Contributions in Science, Number 507

22 km NW Leticia, 4.04°S, 69.99°W, 1$, 4-
7.ix.l997, M. Sharkey, Malaise trap #2 (UNCB).

Melaloncha nigrita Borgmeier

(Fig. 8)

Melaloncha nigrita Borgmeier, 1959:185”186,

figs. 72-74, 76.

HOLOTYPE. ?, BRAZIL: Santa Catarina:
Nova Teutonia, 27.18°S, 52.38°W, v.1952, F.
Plaumann [LACM ENT 116531] (MZSP; exam-
ined).

RECOGNITION. Females of this species can
be recognized by the short processes of the trident
and the dark body color. They are most similar to
M. juxta species, from which they differ by the
characters in the key.

DESCRIPTION. Female. Body length 1.3-

I. 6 mm. Frons dark brown, lighter at dorsolateral
and anterior extremes, strongly reticulate. Mean
frontal width 0.45 head width. Flagellomere 1
yellow. Palpus yellow, palpal setae yellow, with 1
black, apical seta. Postocular setae black. Genal
setae black. Scutum dark brown. Pleuron dark
brown except anterior portion of anepisternum
light brown. Legs whitish-yellow. Venter of
foretibia lacking enlarged setae. Foretarsomeres
relatively elongate, narrow. Posteroventral mar-
gin of tarsomeres 1-2 without enlarged setae.
Large, lateral setae of foretarsomeres relatively
small, inconspicuous. Posterior (outer) tarsal claw
subequal to anterior claw. Mean costal length
0.44 wing length, range 0.39-0.45. Halter yellow.
Abdominal tergites black, with dull silver pollin-
osity. Ovipositor yellow basally, otherwise dark
brown, with short setae, slightly dorsally curved.
Lateral arms of trident short, relatively narrow.
Dorsomedial process of trident subequal in length
to lateral arms.

Male. Unknown.

HOST. Specimens were attracted to mixed
aggregations of bees in Argentina, where the most
likely host (based on size) was Plebeia sp.

GEOGRAPHICAL DISTRIBUTION. South-
eastern Brazil and northeastern Argentina.

OTHER MATERIAL EXAMINED. ARGEN-
TINA: Misiones: Reserva Vida Silvestre Urugua-i,
25.97°S, 54.irW, 6?, 10.xii.2003, 6$,

II. xii.2003, 5$, 12.xii.2003, B. Brown, G. Rung,
bee screen, honey spray, 3$, 17.xii.2003, 1$,
18.xii.2003, L. Gonzalez, G. Rung, honey spray,
400 m (LACM, MACN). BRAZIL: 3? paratypes,
same data as holotype, except vii.1946 (MCZC,
MZSP, USNM).

Melaloncha ungulata-suhgroup

DIAGNOSIS. Setae of at least foretarsomere 2
enlarged: inner (anterior) seta of elongate, outer
(posterior) thickened, curved (Fig. 51).

INCLUDED SPECIES. Melaloncha nigrifrons,
M. platypoda, plus the M. ungulata-series.

Brown and Rung: Revision of M. ungulata-^roup ■ 7

Melaloncha nigrifrons Borgmeier

(Fig. 9)

Melaloncha nigrifrons Borgmeier, 1971a:133-

134, figs. 173-174.

HOLOTYPE. ?, COSTA RICA: San Jose,
15.vii.[no year], H. Schmidt (MZSP; examined
but not bar-coded).

RECOGNITION. Both sexes can be recognized
by the yellowish body with the dark, shiny frons.
Additionally, females have the lateral arms of the
trident greatly elongate.

We collected males of this species for the first
time; they are recognizable by the shiny, dark-
colored frons with the yellowish-colored body,
and their co-occurrence with females. They key to
M. clavata Schmitz in the most recent key to
males (Borgmeier, 1971a), a species not consid-
ered part of the M. ungulata-group because
females lack a trident. The identification of the
male of M. clavata^ however, is questionable, as
the specimen was obtained from a single parasit-
ized bee, and no female Melaloncha specimens
were associated with it (Borgmeier, 1938). The
males of M. horgmeieri (below) also key to this
couplet but differ by their yellow frons and dark
brown foretarsomeres.

DESCRIPTION. Female. Body length 1.7-
1.9 mm. Frons varying from almost completely
dark brown or black, to having various amounts
of yellowish-brown at dorsolateral corners and on
setal bases of all frontal setae; shiny, but with fine,
reticulate sculpturing. Mean frontal width 0.43
head width. Flagellomere 1 white basally, yellow
apically. Palpus white, palpal setae black to
yellow. Postocular setae black dorsally, yellow
or black ventrally. Genal setae yellow to black.
Scutum anteriorly light brown, posteriorly dark
brown, with well-defined anteromedial projection
of dark color. Pleuron brownish-yellow. Legs
brownish-yellow. Venter of foretibia lacking
enlarged setae. Foretarsomeres relatively elongate,
narrow. Posteroventral margin of tarsomeres 1-2
without enlarged setae. Large, lateral setae of
foretarsomeres asymmetrical: posterior setae

shorter, thicker, curved; anterior setae longer,
thinner, straight. Posterior (outer) tarsal claw
slightly longer than anterior claw and directed
more outwards. Mean costal length 0.47 wing
length, range 0.44-0.52. Halter yellow. Abdom-
inal tergites anteriorly yellow, with black
posterior margins. Ovipositor brown basally,
darker brown apically, with short setae, slightly
dorsally curved. Lateral arms of trident elongate,
narrow, with narrower dorsoapical process.
Dorsomedial process of trident much shorter than
lateral arms.

Male. Frons black, with lighter color at dorsal
and ventral extremes and, in some specimens, on
setal bases. Flagellomere 1 yellow. Palpus white;
palpal setae black, well developed. Postocular and
genal setae black. Scutum on anterior two-thirds

8 ■ Contributions in Science, Number 507

light brown, on posterior two-thirds dark brown,
with or without a slight anteromedial projection
of dark color. Pleuron light brown. Legs yellow-
ish-brown, except apex of hind femur with small
posterodorsal margin of dark brown. Foretar-
somere 1 twice as long as broad; other foretar-
someres about as long as broad; pulvilli enlarged,
light brown. Abdominal tergite 1 black; tergites 2
and 6 mostly yellow with thin posterior black
band; tergites 3-5 black with central yellow spot
decreasing in size posteriorly. Venter of abdomen
yellow. Basal one-third of cercus brown, apical
two-thirds yellow. Surstylus with short apical
setae.

VARIATION. Besides the variation in color of
the frons and setae of the head noted above,
female (but not male) specimens from Finca
Montezuma have flagellomere 1 brown tipped.
Specimens from Bolivia are significantly larger
than the others, have the process of the lateral
arms of the trident smaller, and might represent
a separate species.

HOST. Flies were attracted to mixed aggrega-
tions of bees, but no attacks were seen.

GEOGRAPHICAL DISTRIBUTION. Costa
Rica to Argentina.

MATERIAL EXAMINED. ARGENTINA:
Misiones: Reserva Vida Silvestre Urugua-i,

25.97°S, 54.irW, 1?, ll.xii.2003, B. Brown,
G. Kung, bee screen, 400 m (MACN). BOLIVIA:
La Paz: 40 km N Caranavi, Cumbre Alto Beni,
3?, 13. iv. 2003, B. Brown, S. Marcotte, E.
Zumbado, honey spray (CBFC, LACM), 2?,
21.iv.2004, E. Zumbado (LACM). COSTA
RICA: Guanacaste: Rio Naranjo, Finca

Montezuma, 10.67°N, 85.06°W, 2c5, 4?,

26. ii. 2002, 1(5, 27.ii.2002, B. Brown, L. Gonza-
lez, K. Walker, E. Zumbado, honey-sprayed
undergrowth (INBC, LACM, MUCR); Puntare-
nas: Las Alturas, 8.95°N, 82.83°W, 1$,

vi.l992, P. Hanson, Malaise trap, 1500 m
(LACM); San Jose: San Antonio de Escazu,
9.90°N, 84.15°W, lc5, i-ii.l989, W. Eberhard,
Malaise trap, 1300 m (LACM), Zona Protectora
El Rodeo, 9.91°N, 84.28°W, 2$, l-5.viii.2001,
B. Brown, V. Berezovskiy, G. Kung, Malaise trap
(LACM).

Melaloncha platypoda n. sp.

(Figs. 10, 24)

RECOGNITION. This species can be distin-
guished from all other Melaloncha by the greatly
enlarged, flattened foretarsomeres. It is addition-
ally recognized within the M. ungulata-gcoxip by
the yellow coxal setae and the trident with a large
dorsomedial process and extremely short lateral
arms.

One male specimen was collected in copula
with a female. It is extraordinarily divergent from
the female, being almost entirely dark brown
(except the frons), with normal (for a male)

Brown and Kung: Revision of M. ungulata-gr oup

forelegs. In Borgmeier’s (1971a) key to males, it
runs to either M. genitalis or M. glabrifrons^
depending on whether one takes the first or
second lead in couplet 7, which specify a medium-
sized (2.5 mm) or small (1. 8-2.0 mm) specimen,
respectively. The terminalia of M. platypoda
differ from those of M. genitalis^ in which the
surstylar lobes are greatly elongate. The male of
M. glabrifrons is much lighter in color than that
of M. platypoda.

DESCRIPTION. Eemale. Body length 2.1-
2.5 mm. Frons yellow, except ocellar triangle black;
with fine, reticulate sculpturing. Mean frontal width
0.41 head width. Flagellomere 1 white basally,
yellow apically. Palpus yellow, palpal setae yellow.
Postocular setae black dorsally, yellow ventrally.
Genal setae yellow. Scutum anteriorly light brown,
posteriorly dark brown, with well-defined antero-
medial projection of dark color. Pleuron brownish-
yellow. Legs brownish-yellow. Coxal setae yellow.
Venter of foretibia lacking enlarged setae. Foretar-
someres extraordinarily modified: broad, flat, com-
bined length 1.4X length of foretibia. Posteroventral
margin of tarsomeres 1-2 without enlarged setae.
Large, lateral setae of foretarsomeres asymmetrical:
anterior seta longer, thicker. Posterior (outer) tarsal
claw subequal to anterior claw. Long setae of coxae
yellow (black in most other Melaloncha). Mean
costal length 0.50 wing length, range 0.48-0.53.
Halter yellow. Abdominal tergites anteriorly yellow,
with black posterior margins. Ovipositor yellow
basally, otherwise dark brown, with short setae,
slightly dorsally curved. Lateral arms of trident
extremely short, pointed. Dorsomedial process of
trident short, broad, apically truncate in lateral view.

Male. Body length 2.1 mm. Frons orange,
except setal bases dark brown and ocellar triangle
black. Flagellomere 1 orange. Palpus white;
palpal setae black, well developed. Postocular
and genal setae. Thorax and legs almost com-
pletely dark brown, except small spot of light
brown at posterior apex of postpronotal lobe;
foretibia and foretarsomeres 1-4 light brown.
Foretarsomere 1 about twice as long as broad;
other foretarsomeres about as long as broad;
pulvilli enlarged, yellow. Abdominal tergites
black, anteriorly with silvery pollinosity. Venter
of abdomen gray. Basal one-third of cercus
brown, apical two-thirds yellow. Surstylus with
short apical setae.

HOST. At the Wilson Botanical Garden in
Costa Rica, we observed this species attacking
Trigona nigerrima Cresson, T. corvina (Cocker-
ell), and T. fulviventris Guerin on flowers of the
exotic Brazilian palm Syagrus coronata. At other
sites they were attracted to mixed aggregations of
bees, but no attacks were observed.

GEOGRAPHICAL DISTRIBUTION. Costa
Rica to Bolivia.

DERIVATION OE SPECIEIC EPITHET.

Greek for “flat foot,” referring to the broad,
flattened tarsomeres of the female foreleg.

Contributions in Science, Number 507

HOLOTYPE. ?, COSTA RICA: San Jose:
Zurqui de Moravia, 10.05°N, 84.02°W, v.1992,
P. Hanson, Malaise trap, 1600 m [LACM ENT
100755] (LACM).

PARATYPES. BOLIVIA: La Paz: 40 km N
Caranavi, Cumbre Alto Beni, 15.83°S, 67.56°W,
19, 18.iv.2003, 4?, 19.iv.2003, B. Brown, S.
Marcotte, E. Zumbado, honey spray (CBFC,
LACM), 49, 16.iv.2004, 5 9, 17.iv.2004, 3 9,
21.iv.2004, B. Brown, E. Zumbado, honey spray
(LACM), Coroico, Hotel Don Quixote, 16.19°S,
67.72°W, 1 9 , 5.iv.2004, 4 9 , 6.iv.2004, B. Brown,
E. Zumbado, Phoenix palm flowers/honey spray
(LACM). COSTA RICA: Puntarenas: Las Alturas,
8.95°N, 82.83°W, 19, iii.1992, 19, vi.l992, P.
Hanson, Malaise trap, 1500 m (LACM), Wilson
Botanical Garden/Las Cruces, 8.79°N, 82.95°W,
29, 8.vi.l998, 29, 12.vi.l998, 49, 13.vi.l998,
79, 14.vi.l998, 49, 16.vi.l998, B. Brown, V.
Berezovskiy, flowers of Syagms coronata (INBC,
LACM, MCZC, MUCR, MZSP, NHRS, USNM),
1(5, 3 9 (including 1 in copula pair), 17.V.2000,
29, 18.V.2000, G. Rung, on honey baits (LACM);
San Jose, Zurqui de Moravia, 10.05°N, 84.02°W,
I9,ix-x.l990, I9,vii.l991, I9,iii.l992, 19,iv-
V.1993, P. Hanson, Malaise trap, 1600 m
(LACM). ECUADOR: Carchi: Chical, 0.93°N,
78.18°W, 1 9, 22.vii.1983, J. E. Rawlins (CMNH).

Melaloncha ungulata-inhagroup

DIAGNOSIS. Venter of tarsomeres 1-2 with
enlarged setae along posterior margin; enlarged
setae black, contrasting with other tarsal setae,
which are yellow.

The degree of specialization of these setae vary
among species: some have differentiated setae that
only differ slightly from other setae, whereas in
others there is a more marked separation (e.g..
Figs. 45-46 versus Figs. 58-59). The color
differentiation is easily seen in specimens.

INCLUDED SPECIES. Melaloncha diastata,
M. fuscipalpis, M. juxta, M. ustulata, and the
M. Caligula and M. ungulata-senes.

REMARKS. Excluding the species of the M.
ungulata-stries, this group includes some of the
smallest species of Melaloncha. These flies para-
sitize equally small bees, being parasitoids of
small Leurotrigona, Plebeia, and Trigonisca
species.

Melaloncha diastata n. sp.

(Figs. 11,25)

RECOGNITION. Identification of the 4 spe-
cies of small brown Melaloncha with a short
dorsomedial process of the trident is difficult and
is complicated by the scarcity of specimens. One
such species, M. fuscipalpis., is relatively easily
differentiated by the brown palpus. Two of the
remaining 3 species, M. diastata and M. ustulata,
are each represented by single specimens, whereas

Brown and Kung: Revision of M. ungulata-group ■ 9

there are 3 specimens of M. juxta. The single
specimen of M. ustulata has a much lighter brown
frons than the other 2 species, in which the frons
is nearly black, and the base of the forefemur in
M. ustulata is similar in color to the apical one-
half, not darkened like in the other 2 species. In
M. juxta the dorsomedial process is short and not
upturned. All these species are extremely similar,
and the main differences among them are docu-
mented in the key. More specimens are necessary to
determine if these differences are consistent, but for
now we treat the 3 as separate species.

DESCRIPTION. Female. Body length 1.6 mm.
Frons dark brown, lighter at dorsolateral and
anterior extremes, coarsely reticulate. Mean
frontal width 0.45 head width. Flagellomere 1
white basally, brown apically. Palpus white,
palpal setae black. Postocular setae black. Genal
setae black. Scutum dark brown. Pleuron dark
brown. Legs brownish-yellow, except bases of
femora and apical one-half of hind femur dark
brown. Venter of foretibia lacking enlarged setae.
Foretarsomere 1 elongate, twice as long as 2;
foretarsomeres 2-5 short, approximately as long
as broad. Posteroventral margin of tarsomeres 1-
2 with enlarged setae. Large, lateral setae of
foretarsomeres asymmetrical: anterior seta longer,
thicker. Posterior (outer) tarsal claw subequal to
anterior claw. Costal length 0.42 wing length.
Halter yellow. Abdominal tergites black, with
dull silver pollinosity. Ovipositor brown basally,
darker brown apically, setose, relatively straight.
Lateral arms of trident elongate, narrow, widely
spaced. Dorsomedial process of trident slightly
shorter than lateral arms; slightly curved dorsally
at apex.

Male. Unknown.

HOST. Unknown. The single specimen was
attracted to a mixed aggregation of bees attracted
to honey.

GEOGRAPHICAL DISTRIBUTION. Amazo-
nian Colombia.

DERIVATION OF SPECIFIC EPITHET. From
Greek diastasis for “separation,” referring to the
widely separated lateral arms of the trident.

HOLOTYPE. 9, COLOMBIA: Amazonas:
Amacayacu National Park, 3.82°S, 70.26°W,
12. hi. 2000, B. Brown, G. Kung, attracted to bees,
honey on cards [LACM ENT 127879] (UNCB).

Melaloncha fuscipalpis n. sp.

(Figs. 12, 26, 45-47)

RECOGNITION. This species can be recog-
nized by the minute size (relative to other
Melaloncha)., dark body color (including the
unusual brown color of the palpus), and the
shape of the trident.

DESCRIPTION. Female. Body length 1.4-
1.5 mm. Frons dark brown to black, coarsely
reticulate. Mean frontal width 0.48 head width.
Flagellomere 1 white basally, brown apically.

10 ■ Contributions in Science, Number 507

Palpus dark brown, palpal setae black. Postocular
setae black. Genal setae black. Scutum dark brown.
Pleuron dark brown. Legs mostly dark brown;
apical one-quarter of forefemur and apical one-half
of midfemur light brown, fore- and midtibia and all
tarsomeres light brown. Venter of foretibia lacking
enlarged setae. Foretarsomere 1 elongate, twice as
long as 2; foretarsomeres 2-5 short, approximately
as long as broad. Posteroventral margin of tar-
someres 1-2 with enlarged setae. Large, lateral
setae of foretarsomeres asymmetrical: anterior seta
longer, thicker. Posterior (outer) tarsal claw sub-
equal to anterior claw. Mean costal length 0.43
wing length, range 0.42-0.46. Halter yellow.
Abdominal tergites black, with dull silver pollinos-
ity. Ovipositor dark brown, setose, relatively
straight. Lateral arms of trident elongate, narrow.
Dorsomedial process of trident subequal in length
to lateral arms; slightly curved dorsally at apex.

Male. Unknown.

HOST. One fly was collected attacking a work-
er of Trigonisca (an undescribed species) while
still in the tube of the aspirator that collected both
specimens. It attacked with the ovipositor curved
beneath the body (as in Fig. 1).

GEOGRAPHICAL DISTRIBUTION. Southern
Peru.

DERIVATION OE SPECIFIC EPITHET. From
Latin fuscus for “dark,” referring to the color of
the palpus.

HOLOTYPE. ?, PERU: Madre de Dios:
Tambopata Research Center, 13.14°S, 69.61°W,
21.vii.2001, B. Brown, G. Kung, honey-sprayed
leaves [LACM ENT 075507] (MUSM).

PARATYPES. PERU: Madre de Dios: Tambo-
pata Research Center, 13.14°S, 69.6UW, 2$,
21.vii.2001, 2?, 22.vii.2001, 2$, 23.vii.2001,
2$, 24.vii.2001, B. Brown, G. Kung, honey-
sprayed leaves (LAGM, MUSM), 1?, 20-

23.vii.2001, B. Brown, G. Kung, Malaise trap
#5 (LACM).

Melaloncha juxta n. sp.

(Figs. 13, 27)

RECOGNITION. See Recognition for M.
diastata, above.

DESCRIPTION. Female. Body length 1.5-
1.6 mm. Frons dark brown to black, coarsely
reticulate. Mean frontal width 0.45 head width.
Flagellomere 1 white basally, yellow apically
(holotype with brown markings at ventral apex).
Palpus white, palpal setae black. Postocular setae
black. Genal setae black. Scutum dark brown.
Pleuron dark brown. Legs yellow, except coxae,
basal regions of femora, and apex of hind femur
dark brown. Venter of foretibia lacking enlarged
setae. Foretarsomere 1 elongate, twice as long as
2; foretarsomeres 2-5 short, approximately as
long as broad. Posteroventral margin of tarsome-
res 1-3 with group of short, thick setae. Large,
lateral setae of foretarsomeres asymmetrical:

Brown and Kung: Revision of M. ungulata-growp

anterior seta longer, thicker. Posterior (outer)
tarsal claw subequal to anterior claw. Mean
costal length 0.44 wing length, range 0.43-0.45.
Halter yellow. Abdominal tergites black, with
dull silver pollinosity. Ovipositor brown basally,
darker brown apically, setose, relatively straight.
Lateral arms of trident elongate, narrow closely
approximated. Dorsomedial process of trident
short, pointed.

Male. Unknown.

VARIATION. The paratype from Golombia
has thicker lateral arms of the trident. The
specimen from Costa Rica has the darkening of
the forefemur restricted to the extreme base of the
segment, unlike the Colombian specimens, in
which the basal one-half is darkened.

HOST. Unknown. In Colombia, the flies were
attracted to an aggregation dominated by Trigo-
nisca sp.

GEOGRAPHICAL DISTRIBUTION. Colom-
bia and Costa Rica.

DERIVATION OF SPECIFIC EPITHET.

Latin word for “near,” referring to the closely
approximated bases of the lateral arms of the
trident.

HOLOTYPE. $, COLOMBIA: Choco: PNN
Utria, Sendero Boroboro, 6.03°N, 77.32°W,
5.vii.2000, B. Brown, G. Kung, bee screen
[LAGM ENT 075842] (UNCB).

PARATYPES. GOLOMBIA: 1$, same data as
holotype (LAGM). COSTA RICA: Puntarenas:
5.5 km SW Rincon, Tropical Youth Center,
8.70°N, 83.51°W, 1$, ll.viii.2001, B. Brown,
V. Berezovskiy, E. Zumbado, honey-sprayed
leaves (LACM).

Melaloncha ustulata n. sp.

(Figs. 14, 28)

RECOGNITION. See discussion of M. dia-
stata. This species also differs from the other 4
small brown species by having a lighter brown
frons.

DESCRIPTION. Female. Body length 1.4 mm.
Frons brown, setal bases dark brown, coarsely
reticulate. Mean frontal width 0.45 head width.
Flagellomere 1 yellow. Palpus yellow, palpal setae
black. Postocular setae black. Genal setae black.
Scutum dark brown. Pleuron dark brown. Legs
light brown, except midcoxa, hind coxa, and apex
of hind femur dark brown. Venter of foretibia
with row of enlarged setae. Foretarsomere 1
elongate, twice as long as 2; foretarsomeres 2-5
short, approximately as long as broad. Poster-
oventral margin of tarsomeres 1-2 with group of
short, thick setae (that extends across venter of
tarsomere 1). Large, lateral setae of foretarsome-
res asymmetrical: posterior setae shorter, thicker,
curved; anterior setae longer, thinner, straight.
Posterior (outer) tarsal claw subequal to anterior
claw. Costa 0.46 wing length. Halter yellow.
Abdominal tergites anteriorly brown, with darker

Contributions in Science, Number 507

posterior margins and silver pollinosity. Ovipos-
itor yellow basally, otherwise dark brown, setose,
slightly ventrally curved. Lateral arms of trident
short, relatively narrow. Dorsomedial process of
trident subequal in length to lateral arms; slightly
curved dorsally at apex.

Male. Unknown.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Costa
Rica.

DERIVATION OE SPECIEIC EPITHET. Latin
for “darkened” or “browned,” referring to the
color of the frons.

HOLOTYPE. ?, COSTA RICA: Puntarenas:
3 km SW Rincon, 8.68°N, 83.48°W, xii.1989, P.
Hanson, Malaise trap, 10 m [LACM ENT
004669] (LACM).

Melaloncha caligula-stnts

DIAGNOSIS. Unusually small species of Mel-
aloncha (as are the 4 preceding species). Dor-
somedial process of trident slightly to greatly
enlarged, laterally flattened (Figs. 15-23).

INCLUDED SPECIES. Melaloncha claviapex,
M. debilis, M. setitibialis, and the M. caligula-
subseries.

Melaloncha claviapex n. sp.

(Fig. 15)

RECOGNITION. This species is easily recog-
nized by the elongate, straight dorsomedial pro-
cess of the trident, along with the clavate lateral
arm. Its trident is most similar to that of M.
setitibialis, but it lacks the elongate foretibial
setae of that species.

DESCRIPTION. Female. Body length 1.6 mm.
Frons dark brown, coarsely reticulate. Mean
frontal width 0.46 head width. Flagellomere 1
white basally, yellow apically. Palpus white,
palpal setae black. Postocular setae black. Genal
setae black. Scutum dark brown. Pleuron dark
brown. Legs brownish-yellow, except midcoxa,
hind coxa and apical one-quarter of hind femur
dark brown. Venter of foretibia lacking enlarged
setae. Foretarsomeres 2-5 short, approximately as
long as broad. Posteroventral margin of tarsome-
res 1-2 with group of short, thick setae. Large,
lateral setae of foretarsomeres asymmetrical:
anterior seta longer, thicker; setae of tarsomeres
2-3 unusually and distinctively enlarged, those of
tarsomere 4 relatively small. Posterior (outer)
tarsal claw slightly longer than anterior claw and
directed more outwards. Costa 0.46 wing length.
Halter yellow. Abdominal tergites black, with
dull silver pollinosity. Ovipositor brown basally,
darker brown apically, with short setae, slightly
dorsally curved. Lateral arms of trident elongate,
apically expanded and club-tipped. Dorsomedial
process of trident elongate, laterally flattened,
shorter than lateral arms.

Male. Unknown.

Brown and Kung: Revision of M. ungulata-^roup ■ 11

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Southern
Peru.

DERIVATION OF SPECIFIC EPITHET. From
Latin clava for “club” and apex for “tip,” refer-
ring to the club-tipped apices of the lateral arms
of the trident.

HOLOTYPE. ?, PERU: Madre de Dios:
Tambopata Research Center, 13.14°S, 69.6UW,
24.vii.2001, B. Brown, G. Kung, honey-sprayed
leaves [LACM ENT 074467] (MUSM).

Melaloncha debilis n. sp.

(Figs. 16, 48-51)

RECOGNITION. The species can be recog-
nized by the dorsomedial process of the trident,
which is small, knoblike, and supported by a short
stalk.

DESCRIPTION. Female. Body length 1.4-
1.7 mm. Frons dark brown, lighter at dorsolateral
and anterior extremes, coarsely reticulate. Mean
frontal width 0.47 head width. Flagellomere 1
yellow. Palpus yellow, palpal setae black. Posto-
cular setae black. Genal setae black. Scutum
anteriorly light brown, posteriorly dark brown.
Pleuron light brown, with darker markings. Legs
brownish-yellow. Foretibia with posteroventral
row of enlarged setae that are about equal in
length to depth of tibia. Foretarsomeres 2-5 short,
approximately as long as broad. Posteroventral
margin of tarsomeres 1-2 with group of short,
thick setae. Large, lateral setae of foretarsomeres
asymmetrical: posterior setae shorter, thicker,
curved; anterior setae longer, thinner, straight.
Posterior (outer) tarsal claw slightly longer than
anterior claw and directed more outwards. Mean
costal length 0.46 wing length, range 0.43-0.48.
Halter yellow. Abdominal tergites black, with
dull silver pollinosity. Ovipositor dark brown,
setose, slightly dorsally curved. Lateral arms of
trident broad. Dorsomedial process of trident
slender, longer than lateral arms, slightly expand-
ed apically.

Male. Unknown.

HOST. We observed this bee attacking Trigo-
nisca (an undescribed species) in Colombia;
otherwise, it has been attracted to mixed aggrega-
tions of bees.

GEOGRAPHICAL DISTRIBUTION. Colom-
bia and Costa Rica.

DERIVATION OF SPECIFIC EPITHET. Latin
for “weak,” referring to the relatively small,
poorly developed apex of the dorsomedial process
of the trident.

HOLOTYPE. ?, COSTA RICA: Heredia: La
Selva Biological Station, 10.43°N, 84.02°W,
16.ii-2.iii.l993, ALAS, Malaise trap M/07/022
[INBIOCRI001264310] (INBC).

PARATYPES. COLOMBIA: Choco: PNN

Utria, Sendero Boroboro, 6.03°N, 77.32°W, 2?,
5.vii.2000, B. Brown, G. Kung, bee screen

12 ■ Contributions in Science, Number 507

(LACM, UNCB). COSTA RICA: Heredia: La
Selva Biological Station, 10.43°N, 84.02°W, 6?,
3.iii.l993, ALAS, Malaise trap M/06/021, M/16/
031, 4?, 15.iii.l993, ALAS, Malaise trap M/06/
037, 2?, 2.iv.l993, ALAS, Malaise trap M/15/
062, 1?, 4.iv.l994, ALAS, Malaise trap M/12/
390, 1$, 14.ix.l995, ALAS, Malaise trap M/04/
450, 1.1V.1996, ALAS, Malaise trap M/07/608
(INBC, LACM, MUCR), 1?, 17.ii.2002, L.
Gonzalez, honey-sprayed undergrowth (LACM).

Melaloncha setitibialis n. sp.

(Fig. 17)

RECOGNITION. This species is most similar to
M. claviapex but differs by the shape of the trident
and by the long setae on the venter of the foretibia.

DESCRIPTION. Female. Body length 1.6-
1.7 mm. Frons dark brown, lighter at dorsolateral
and anterior extremes, coarsely reticulate. Mean
frontal width 0.4 head width. Flagellomere 1
yellow. Palpus white, palpal setae black. Posto-
cular setae black. Genal setae black. Scutum
dark brown. Pleuron dark brown except proepis-
ternum and anterior portion of anepisternum
light brown. Legs brownish-yellow except apex
of hind femur dark brown. Foretibia with poster-
oventral row of extremely enlarged setae that are
approximately twice width of tibia. Foretarsome-
res 2-5 short, approximately as long as broad.
Posteroventral margin of tarsomeres 1-2 with
group of short, thick setae. Large, lateral setae of
foretarsomeres asymmetrical: posterior setae

shorter, thicker, curved; anterior setae longer,
thinner, straight. Posterior (outer) tarsal claw
greatly enlarged, about twice size of anterior
claw. Mean costal length 0.47 wing length, range
0.45-0.49. Halter yellow. Abdominal tergites
black, with dull silver pollinosity. Ovipositor
dark brown, setose, slightly dorsally curved.
Lateral arms of trident elongate, apically expand-
ed and club-tipped. Dorsomedial process of
trident elongate, apically rounded and dorsoven-
trally flattened.

Male. Unknown.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Southern
Peru.

DERIVATION OF SPECIFIC EPITHET. Latin
seta for “bristle” and tibialis for “of the leg,”
referring to the long setae on the foretibia.

HOLOTYPE. $, PERU: Madre de Dios:
Tambopata Research Center, 13.14°S, 69.61°W,
22.vii.2001, B. Brown, G. Rung, honey-sprayed
leaves [LACM ENT 075454] (MUSM).

PARATYPE. 1 $ , same data as holotype
(LACM).

Melaloncha caligMla-suhsenes

DIAGNOSIS. Dorsomedial process with greatly
enlarged, flattened flange (Figs. 18-23).

Brown and Kung: Revision of M. ungulata-gr oup

INCLUDED SPECIES. Melaloncha caltgula, M.
cristula, M. culmena, M. spina, M. tambopaten-
sis, and M. tnparroensis.

Melaloncha caligula n. sp.

(Fig. 18)

RECOGNITION. This species is recognized by
the dorsomedial process of the trident, which is
greatly expanded but has only a minute notch
below.

DESCRIPTION. Female. Body length 1.6 mm.
Frons dark brown, lighter at dorsolateral and
anterior extremes, coarsely reticulate. Mean
frontal width 0.47 head width. Flagellomere 1
yellow. Palpus yellow, palpal setae yellow, with 1
black, apical seta. Postocular setae black. Genal
setae black. Scutum anteriorly light brown,
posteriorly dark brown, with well-defined ante-
romedial projection of dark color. Pleuron dark
brown, except proepisternum and anterior por-
tion of anepisternum light brown. Legs yellow,
apex of hind femur dark brown. Venter of
foretibia with row of enlarged setae. Foretar-
someres 2-5 short, approximately as long as
broad. Posteroventral margin of tarsomeres 1-2
with group of short, thick setae. Large, lateral
setae of foretarsomeres asymmetrical: anterior
seta longer, thicker. Posterior (outer) tarsal claw
slightly longer than anterior claw and directed
more outwards. Costa 0.43 wing length. Halter
yellow. Abdominal tergites black, with dull silver
pollinosity. Ovipositor brown basally, darker
brown apically, setose, slightly dorsally curved.
Lateral arms of trident broad, short. Dorsomedial
process of trident greatly enlarged, laterally
flattened, broadly attached, with only minute
notch below.

Male. Unknown.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Amazo-
nian Colombia.

DERIVATION OF SPECIFIC EPITHET. Latin
diminutive for “boot,” referring to the shape of
the trident in lateral view.

HOLOTYPE. $, COLOMBIA: Amazonas:
Amacayacu National Park, 3.82°S, 70.26°W,
12. hi. 2000, B. Brown, G. Kung, attracted to bees,
honey on cards [LACM ENT 127887] (UNCB).

Melaloncha cristula n. sp.

(Fig. 19)

RECOGNITION. The narrow dorsal crest of
the dorsomedial process of the trident is distinc-
tive.

DESCRIPTION. Eemale. Body length 1.5-
1.6 mm. Frons dark brown, lighter at dorsolateral
and anterior extremes, coarsely reticulate. Mean
frontal width 0.5 head width. Flagellomere 1
yellow. Palpus white, palpal setae black to brown,
with 1 black, apical seta. Postocular setae brown.

Contributions in Science, Number 507

Genal setae brown. Scutum dark brown. Pleuron
dark brown except anterior portion of anepister-
num light brown. Legs brownish-yellow except
apex of hind femur dark brown. Venter of
foretibia with row of enlarged setae. Foretar-
someres 2-5 short, approximately as long as
broad. Posteroventral margin of tarsomeres 1-2
with group of short, thick setae. Large, lateral
setae of foretarsomeres asymmetrical: posterior
setae shorter, thicker, curved; anterior setae
longer, thinner, straight. Posterior (outer) tarsal
claw greatly enlarged, about twice size of anterior
claw. Mean costal length 0.43 wing length, no
variation. Halter whitish-yellow. Abdominal ter-
gites black, with dull silver pollinosity. Ovipositor
brown basally, darker brown apically, setose,
slightly dorsally curved. Lateral arms of trident
elongate, apically expanded and club-tipped.
Dorsomedial process of trident greatly enlarged,
laterally flattened, with elongate stalk, tall dorsal
crest and pointed ventral apex.

Male. Unknown.

HOST. The holotype was attacking a worker of
Leurotrigona muelleri (Friese).

GEOGRAPHICAL DISTRIBUTION. Southern
Peru.

DERIVATION OF SPECIFIC EPITHET. From
Latin crista for “crest,” referring to the shape of
the dorsomedial process of the dorsomedial pro-
cess of the trident.

HOLOTYPE. ?, PERU: Madre de Dios,
Tambopata Research Center, 13.14°S, 69.61°W,
24.vii.2001, B. Brown, G. Kung, attacking bee
host [LACM ENT 075520] (MUSM).

PARATYPES. 1 $ , same data as holotype
except 22.vii.2001, honey-sprayed leaves
(LACM).

Melaloncha culmena n. sp.

(Fig. 20)

RECOGNITION. The dorsomedial process of
the trident is long, narrow, and dorsally truncated
in this species.

DESCRIPTION. Eemale. Body length 1.4 mm.
Frons brown, coarsely reticulate. Mean frontal
width 0.5 head width. Flagellomere 1 yellow.
Palpus yellow, palpal setae yellow. Postocular
setae brown. Genal setae brown. Scutum dark
brown. Pleuron dark brown. Legs brownish-
yellow except apex of hind femur with dark
brown spot. Venter of foretibia with row of
enlarged setae. Foretarsomeres 2-5 short, approx-
imately as long as broad. Posteroventral margin of
tarsomeres 1-2 with group of short, thick setae.
Large, lateral setae of foretarsomeres asymmetri-
cal: anterior seta longer, thicker. Posterior (outer)
tarsal claw slightly longer than anterior claw and
directed more outwards. Costa 0.44 wing length.
Halter whitish-yellow. Abdominal tergites black,
with dull silver pollinosity. Ovipositor brown
basally, darker brown apically, setose, slightly

Brown and Kung: Revision of M. ungulata-group ■ 13

dorsally curved. Lateral arms of trident broad.
Dorsomedial process of trident greatly enlarged,
laterally flattened, with elongate stalk; consists of
strongly sclerotized, dorsal support and small,
ventral, lighter brown flattened area; dorsal apex
slightly narrowed, rounded, knoblike; poster-
oventral portion of flattened area without con-
cavity; circular excision below stalk relatively
small.

Male. Unknown.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Amazo-
nian Brazil.

DERIVATION OE SPECIEIC EPITHET. From
Latin culmen for “top,” referring the dorsal
development of the dorsomedial process of the
trident.

HOLOTYPE. ?, BRAZIL: Amazonas: 60 km
N Manaus, Reserva Campina, 2.67°S, 60.02°W,
8-19.vi.l992, J. Vidal [LACM ENT 122639]
(INPA).

Melaloncha spina n. sp.

(Fig. 21)

RECOGNITION. This species can be recog-
nized by the shape of the dorsomedial process of
the trident, which has a small, posterodorsal
concavity.

DESCRIPTION. Eemale. Body length 1.4 mm.
Frons dark brown, lighter at dorsolateral and
anterior extremes, coarsely reticulate. Mean
frontal width 0.45 head width. Flagellomere 1
yellow. Palpus white, palpal setae black. Posto-
cular setae black. Genal setae black. Scutum dark
brown. Pleuron dark brown, except proepister-
num and anterior portion of anepisternum light
brown. Legs brownish-yellow, except apex of
hind femur dark brown. Venter of foretibia with
row of enlarged setae. Foretarsomeres 2-5 short,
approximately as long as broad. Posteroventral
margin of tarsomeres 1-2 with group of short,
thick setae. Large, lateral setae of foretarsomeres
asymmetrical: posterior setae shorter, thicker,
curved; anterior setae longer, thinner, straight.
Posterior (outer) tarsal claw slightly longer than
anterior claw and directed more outwards. Costa
0.43 wing length. Halter whitish-yellow. Abdom-
inal tergites black, with dull silver pollinosity.
Ovipositor brown basally, darker brown apically,
setose, slightly dorsally curved. Lateral arms of
trident broad. Dorsomedial process of trident
greatly enlarged, laterally flattened, with elongate
stalk; dorsal margin dark brown, ventrally with
lighter brown flange; posterodorsal apex with
small concavity; circular excision below stalk
relatively small.

Male. Unknown.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Southern
Peru.

14 ■ Contributions in Science, Number 507

DERIVATION OF SPECIFIC EPITHET. Latin
for “spine” or “backbone,” referring to the dark
ridge on the dorsal margin of the dorsomedial
process of the trident.

HOLOTYPE. ?, PERU: Madre de Dios:
Tambopata Research Center, 13.14°S, 69.61°W,
22.vii.2001, B. Brown, G. Kung, honey-sprayed
leaves [LACM ENT 075716] (MUSM).

Melaloncha tambopatensis n. sp.

(Fig. 22)

RECOGNITION. This species differs from
other M. culmena-suhstnts species by the dor-
somedial process of the trident, which is dorsoa-
pically rounded and has a small posteroventral
concavity.

DESCRIPTION. Female. Body length 1.3-
1.4 mm. Frons dark brown, lighter at anterior
extreme; coarsely reticulate. Mean frontal width
0.48 head width. Flagellomere 1 yellow. Palpus
white, palpal setae black. Postocular setae black.
Genal setae black. Scutum dark brown. Pleuron
dark brown with silvery pollinosity. Legs brown-
ish-yellow, except apex of hind femur dark
brown. Venter of foretibia with row of enlarged
setae. Foretarsomeres 2-5 short, approximately
as long as broad. Posteroventral margin of
tarsomeres 1-2 with group of short, thick setae.
Large, lateral setae of foretarsomeres asymmet-
rical: posterior setae shorter, thicker, curved;
anterior setae longer, thinner, straight. Posterior
(outer) tarsal claw slightly longer than anterior
claw and directed more outwards. Mean costal
length 0.44 wing length, range 0.41-0.46. Halter
yellow. Abdominal tergites black, with dull silver
pollinosity. Ovipositor brown basally, darker
brown apically, setose, slightly dorsally curved.
Lateral arms of trident broad. Dorsomedial
process of trident greatly enlarged, laterally
flattened, with elongate stalk; dorsal crest not
rising significantly above line of stalk, posteriorly
broadly rounded, truncate; posteroventral apex
with small concavity; circular excision relatively
large.

Male. Unknown.

HOST. One Peruvian specimen was collected
attacking a worker of Trigonisca scabiosa Albu-
querque.

GEOGRAPHICAL DISTRIBUTION. Amazo-
nian Brazil and Peru.

DERIVATION OF SPECIFIC EPITHET.

Named for the type locality.

HOLOTYPE. ?, PERU: Madre de Dios:
Tambopata Research Center, 13.14°S, 69.61°W,
24.vii.2001, B. Brown, G. Kung, honey-sprayed
leaves [LACM ENT 074437] (MUSM).

PARATYPES. BRAZIL: Amazonas: Manaus,
Reserva Ducke, 3.13°S, 60.02°W, 1$, 8-

15.iv.l992, J. Vidal, Arm. Cola 1-B-lOm (INPA).
PERU: 19, same data as holotype (LACM).

Brown and Kung: Revision of M. ungulata-group

Melaloncha tuparroensis n. sp.

(Fig. 23)

RECOGNITION. This species is characterized
by the dorsoapical process of the trident, which is
dorsally bluntly pointed, with an evenly convex
posterior margin.

DESCRIPTION. Female. Body length 1.3 mm.
Frons dark brown, lighter at dorsolateral and
anterior extremes, setal bases of ventral inter-
frontal setae also light brown; coarsely reticulate.
Mean frontal width 0.48 head width. Flagello-
mere 1 yellow. Palpus yellow, palpal setae brown.
Postocular setae brown. Genal setae brown,
largest setae darker. Scutum dark brown. Pleuron
light brown, with darker markings. Legs brown-
ish-yellow except apex of hind femur dark
brown. Venter of foretibia with row of enlarged
setae. Foretarsomeres 2-5 short, approximately
as long as broad. Posteroventral margin of tarso-
meres 1-2 with group of short, thick setae.
Large, lateral setae of foretarsomeres asymmetri-
cal: anterior seta longer, thicker. Posterior (outer)
tarsal claw slightly longer than anterior claw
and directed more outwards. Costa 0.44 wing
length. Halter yellow. Abdominal tergites black,
with dull silver pollinosity. Ovipositor brown
basally, darker brown apically, setose, slightly
dorsally curved. Lateral arms of trident
elongate, apically expanded and club-tipped,
truncate. Dorsomedial process of trident greatly
enlarged, laterally flattened, with elongate stalk,
tall dorsal crest and pointed ventral apex; dorsal
apex broadly pointed, posterior margin evenly
convex.

Male. Unknown.

HOST. The holotype was attracted to an
aggregation of Tetragona sp.

GEOGRAPHICAL DISTRIBUTION. Eastern
Colombia.

DERIVATION OF SPECIFIC EPITHET.

Named for the type locality.

HOLOTYPE. 9, COLOMBIA: Vichada: PNN
El Tuparro, 5.35°N, 67.86°W, 16.vi.2000, G.
Kung, bee screen [LACM ENT 055123] (UNCB).

Melaloncha (M.) ungulata-sents

DIAGNOSIS. Foretarsomere 1 with row of
long anteroventral setae. Posterior claw of foreleg
extremely elongate, 2.5 times length of anterior
claw. Ovipositor short, upturned, setose, relative-
ly uniform within group. Trident modified from
other M. ungulata~suhg\:o\xp species, with dor-
somedial process expanded into rounded bulb
(Figs. 32, 34), or dorsally flattened, shelflike
structure (Fig. 33); lateral arms extremely short
to medium length. The posterior portion of the
trident has posterior extension, formed by mem-
brane surrounding an elongate process that is
dorsally curved at apex (e.g.. Figs. 29-30).

PHYLOGENETIC RELATIONSHIPS. The re-
lationships among species within this group are

Contributions in Science, Number 507

mostly unknown. Most species are extremely
similar, except for small differences in the
structure of intersegment 7-8. A preliminary
molecular data set of mitochondrial 12S rRNA,
16S rRNA, NADH 1 and nuclear 28S rRNA
included 3 species of this subgroup and strongly
supported a closer grouping of M. castanea n. sp.
and M. strigosa n. sp. with each other than with
M. trita (B. Brown and P. Smith, in preparation).
Three other species, M. atrilingula, M. inicua, and
M. pegmata, are also hypothesized to belong to
a monophyletic group (below).

NOTES ON INCLUDED SPECIES. Two of the
male-only species described by Borgmeier could
potentially fall within the group treated herein:
M. genitalis and M. glabrifrons. Borgmeier’s
potentially spurious associations of these species
names with M. ungulata gives rise to considerable
confusion (discussed below).

Species that clearly belong to the M. ungulata-
series include M. borgmeieri and M. curtibrachia
and species of the M. ungulata-suhstnes.

Melaloncha genitalis Borgmeier

Melaloncha genitalis Borgmeier, 1934, p. 181,

plate 3, fig. 15.

HOLOTYPE. (5, BRAZIL: Espirito Santo:
Laranja da Terra, 21. iv. 1928, O. Conde, [LACM
ENT 100112] (MZSP).

RECOGNITION. This species is known from
a single male specimen from Espirito Santa,
Brazil. Borgmeier later (1971a) synonymized M.
ungulata^ a species known from both sexes (from
Nova Teutonia, Santa Catarina, Brazil), with M.
genitalis. This synonymy was based on the
supposed inadequacy of the differences between
males of the 2 species cited in his (1959) key with
this species. Our comparison of the terminalia of
M. genitalis with those of presumed males of M.
ungulata, however, shows little similarity: the
lower (surstylar) lobe of the left side of the
epandrium is much larger in M. genitalis. Based
on this observation, we remove M. ungulata from
synonymy with M. genitalis and reinstate it as
a valid species (below). Furthermore, there is no
support for the inclusion M. genitalis in the M.
ungulata-group.

Borgmeier (1971a) also noted further Brazilian
specimens that he attributed to M. genitalis;
whether these are M. genitalis or M. ungulata is
unknown.

GEOGRAPHICAL DISTRIBUTION. Brazil.

Melaloncha glabrifrons Borgmeier

Melaloncha glabrifrons Borgmeier, 1934, p. 178,

plate 3, fig. 11.

HOLOTYPE. $, COSTA RICA: Suiza de
Turrialba, 1921 [LACM ENT 122184] (Hungar-
ian Natural History Museum, Budapest, Hun-
gary). Note that Borgmeier’s original description

Brown and Kung: Revision of M. ungulata-grovrp ■ 15

gives the collection date as 4.vi.l921, but these
details are not on the specimen label.

RECOGNITION. This species is known from
a single male specimen from Suiza de Turrialba,
Costa Rica. Many years after description of the
holotype, Borgmeier (1959) also associated a se-
ries of male and female specimens from Nova
Teutonia, Brazil, with the name M. glabrifrons.
The males were considered to be the same species
based on coloration of the legs (a contrastingly
dark forefemur), a highly suspect character
especially when considering the numerous unde-
scribed species of Melaloncha. Comparison of the
female specimens from Brazil with those of the M.
ungulata-suhgroup from Costa Rica shows that
they are different and should be considered
separate species (see M. borgmeieri, below).
Furthermore, the male of M. borgmeieri has
now been collected (below) and shows that the
association of the M. glabrifrons males with
females from Brazil was entirely spurious, as the
newly collected males of M. borgmeieri have
a light-colored, not dark, forefemur. There is
therefore no credible evidence that M. glabrifrons
belongs in the M. ungulata-group.

GEOGRAPHICAL DISTRIBUTION. Costa
Rica.

Melaloncha borgmeieri n. sp.

(Figs. 29, 52-55)

Melaloncha glabrifrons: 1959, pp. 178-179 (spe-
cimens from Nova Teutonia, Brazil).

RECOGNITION. Females of this species are
distinguished from the only other species lacking
a bristlelike seta at the apex of foretarsomere 5,
M. curtibrachia, by the small, nonprotruding
knoblike dorsomedial process.

The male of this species was recently collected,
as we found 2 pairs in copula. This species cannot
be the same as the specimens of M. glabrifrons,
with which the females were formerly associated,
because they lack the strikingly darkened fore-
femur referred to by Borgmeier (1959:179). The
males of M. borgmeieri key to M. clavata Schmitz
in the most recent key to males (Borgmeier,
1971a), a species not considered part of the M.
ungulata-gmup because females lack a trident.
The identification of the male of M. clavata,
however, is questionable, as the specimen was
obtained from a single parasitized bee, and no
female Melaloncha specimens were associated
with it (Borgmeier, 1938). Males of M. nigrifrons
(above) also key to this couplet but are easily
distinguished by their dark brown frons and
yellow foretarsomeres.

DESCRIPTION. Female. Body length 1.5-
2.0 mm. Frons yellow, except anterior margin of
ocellar triangle black; with finely reticulate
sculpturing (Fig. 52). Mean frontal width 0.50
head width. Flagellomere 1 yellow. Palpus yellow.

16 ■ Contributions in Science, Number 507

with short yellow setae; apical seta brown. Dorsal
postocular setae brown; genal and other posto-
cular setae yellow. Scutum yellow with postero-
lateral black spot. Pleuron yellow. Legs yellow.
Foretibia without strongly differentiated rows of
dorsal setae; with generally bare area dorsally.
Foretarsomeres enlarged, flattened. Venter of
foretarsomere 1 with numerous enlarged setae
on anterior margin, although these setae not
strongly differentiated from other setae on venter
of tarsomere; posterior margin with row of larger
setae. Venter of foretarsomeres 2-3 with many
longer, spinelike setae on posterior margin;
apicoventral seta on anterior side greatly en-
larged. Venter of foretarsomere 4 without spine-
like setae, but with enlarged apical seta on
posterior side and enlarged seta at midlength
on anterior side. Dorsum of foretarsomere 5
without thick, apical seta. Posterior claw of
foreleg greatly enlarged, mean 2.33 times length
of anterior claw, range 2.25-2.50. Mean costal
length 0.47 wing length, range 0.45-0.50. Halter
yellow. Abdominal tergites yellow, with post-
erior black band. Venter of abdomen yellow.
Ovipositor basally yellow, apically black. Trident
of intersegment 7-8 with barely developed
dorsal bulge of dorsomedial process; lateral arms
short.

Male. Frons yellow, except anterior margin of
ocellar triangle black. Flagellomere 1 yellow.
Palpus white; palpal setae black, well developed.
Postocular and genal setae black. Scutum anteri-
orly light brown, posteriorly dark brown, with
well-defined anteromedial projection of dark
color. Pleuron light brown, with sinuous longitu-
dinal line of dark brown across entire length. Legs
yellowish-brown, except foretarsomeres and apex
of hind femur dark brown. Foretarsomere 1 twice
as long as broad; other foretarsomeres about as
long as broad; pulvilli enlarged, light brown.
Abdominal tergite 1 black; tergites 2 and 6 mostly
yellow with thin posterior black band; tergites 3-
5 black with central yellow spot decreasing in size
posteriorly. Venter of abdomen yellow. Basal one-
third of cercus brown, apical two-thirds yellow.
Surstylus with short apical setae.

HOST. At Urugua-i we observed these flies
attacking Plebeia sp. and Tetragonisca angustula
(Latreille). One fly attacked a worker of T.
angustula on Brown’s hand and was observed at
close range. The fly approached the bee directly in
front of the bee’s head. The fly was on foot, with
its ovipositor curved under the body and project-
ing anteriorly (as in Fig. 1). Suddenly, the fly
darted forward, apparently attacking the bee’s
head. The attack was extremely rapid and
knocked the bee over, somersaulting it backwards
onto its back. It remained in this position for
several seconds, weakly waving its legs and
appearing stunned.

GEOGRAPHICAL DISTRIBUTION. North-
ern Argentina, southeastern Brazil.

Brown and Kung: Revision of M. ungulata-%rowp

DERIVATION OF SPECIFIC EPITHET.

Named for the great phoridologist Thomas
Borgmeier.

HOLOTYPE. ?, ARGENTINA: Misiones:
Reserva Vida Silvestre Urugua-i, 25.97°S,
54.1 1°W, 28.xii.2003, L. Gonzalez, honey spray
[LACM ENT 188203] (MACN).

OTHER MATERIAL EXAMINED. ARGEN-
TINA: Misiones: Reserva Vida Silvestre Urugua-i,
25.97°S, 54.1 rW, 1$, 8.xii.2003, B. Brown, 1?,
10.xii.2003, G. Kung, bees on organic waste
(LACM), 3$, ll.xii.2003, 6$, 12.xii.2003, B.
Brown, G. Kung, bee screen (LACM), 43?,
17.xii.2003, 45?, 18.xii.2003, B. Brown, G.
Kung, L. Gonzalez, honey spray (EMUS, LACM,
MCZC, USNM), 1?, 21.xii.2003, 6?,

22.xii.2003, 2?, 26.xii.2003, ld',53? (including
1 in copula pair), 27.xii.2003, 12?, 28.xii.2003,
1(3, 22? (including 1 in copula pair),

29.xii.2003, L. Gonzalez, attracted to honey
spray, 400 m (LACM, MACN). BRAZIL: Santa
Catarina: Nova Teutonia, 27.18°N, 52.38°W,
3?, vii.I946, F. Plaumann (MZSP, USNM).

Melaloncha curtibrachia n. sp.

(Fig. 30)

RECOGNITION. Females of this species differ
from the only other species lacking a bristlelike
seta at the apex of foretarsomere 5, M. borg-
meieri, by the downturned apex of the dorsome-
dial process, and by the extremely short lateral
arms of the trident.

DESCRIPTION. Female. Body length 1.7-
1.8 mm. Frons yellow, except anterior margin of
ocellar triangle black; finely reticulate. Mean
frontal width 0.55 head width. Flagellomere 1
yellow. Palpus yellow, with short yellow setae;
apical seta black. Dorsal postocular setae black;
genal and other postocular setae yellow. Scutum
yellow with posterolateral black spot. Pleuron
yellow. Legs yellow. Foretibia with 2 rows of
slightly enlarged dorsal setae separated by dorsal
bare area. Foretarsomeres slightly enlarged, slight-
ly flattened. Venter of foretarsomere 1 with
numerous enlarged setae on anterior margin,
although these setae not strongly differentiated
from other setae on venter of tarsomere; posterior
margin with row of larger setae. Venter of
foretarsomeres 2-3 with many longer, spinelike
setae on posterior margin; apicoventral seta on
anterior side greatly enlarged. Venter of foretar-
somere 4 without spinelike setae, but with enlarged
apical seta on posterior side and enlarged seta at
midlength on anterior side. Dorsum of foretar-
somere 5 without thick, apical seta. Posterior claw
of foreleg greatly enlarged, 2.38 times length of
anterior claw (no variation). Mean costal length
0.51 wing length, range 0.48-0.54. Halter yellow.
Abdominal tergites yellow, with posterior black
band. Venter of abdomen yellow. Ovipositor
basally yellow, apically black. Trident of interseg-

Contributions in Science, Number 507

ment 7-8 with dorsomedial process not developed,
curved ventrally at apex; lateral arms extremely
short, difficult to see except in dorsal view.

Male. Unknown.

HOST. Unknown. The holotype was collected
at an aggregation of bees attracted to honey
spray.

GEOGRAPHICAL DISTRIBUTION. Costa
Rica.

DERIVATION OF SPECIFIC EPITHET. Based
on Latin words for “short arm,” referring to the
short lateral arms of the trident.

HOLOTYPE. ?, COSTA RICA: Puntarenas:
R. Privada Karen Mogensen, camino a sendero
El Viejo Nispero, 300-500 m, 8.vii.2003, Y.
Cardenas, red con aguamiel [INB0003738670]
(INBC).

PARATYPES. COSTA RICA: Guanacaste,
Nandayure, Bellavista, Cerro Azul, 27$, 28. ii-
5.iii.2004, W. Porras, honey spray, 1050 m,
#76449 (INBC, LACM), 3 km SE Rio Naranjo,
1$, 2-31.1.1992, F. D. Parker (LACM); San Jose:
trail to PN La Cangreja, 9.68°N, 84.37°W, 6$,
17.vii.2004, W. Porras, honey spray, 300 m,
#77716 (INBC, LACM).

Melaloncha ungulata-suhstvits

DIAGNOSIS. Dorsum of foretarsomere 5 with
enlarged seta at apex.

INCLUDED SPECIES. Melaloncha castanea,
M. laticlava, M. strigosa, M. trita, M. ungulata,
and the species of the M. atrilingula-mka.senes.

Melaloncha castanea n. sp.

(Fig. 31)

RECOGNITION. This species is most similar
to M. ungulata, from which it differs not only by
the darker color but also by the dorsomedial
process of the trident, which in dorsal view is
gradually broadened apically. In contrast, the
dorsomedial process of M. ungulata is narrower
at both midlength and apically.

Males were collected at the same bee nest
entrances as females and resemble them in general
appearance. Although no pairs were collected in
copula, we consider them to be conspecific. They
key to couplet 9 in Borgmeier’s latest (1971a) key
to males, which differentiates larger (2.5 mm)
from smaller (1. 8-2.0 mm) species. Because M.
castanea is intermediate, we tried both ways in the
key. Treating them as larger, they key to M.
genitalis, whose surstyli are massive and much
larger than those of M. castanea. Treating them as
small species leads to M. glabrifrons, from which
they differ in having the forefemur only partly
brown, but there are insufficient characters to
definitely exclude this possibility. The large geo-
graphic separation of the collecting localities of
the specimens, Costa Rica versus the mouth of the
Amazon River, however, suggests that they are
not conspecific.

Brown and Kung: Revision of M. ungulata-grow^ ■ 17

DESCRIPTION. Female. Body length 1.9-
2.3 mm. Frons yellow, except anterior margin
of ocellar triangle black; with fine, reticulate
sculpturing. Mean frontal width 0.42. Flagello-
mere 1 yellow, with dark brown encircling band.
Palpus white, with medium-sized, black setae.
Postocular and genal setae black. Scutum yellow-
ish-brown, with dark brown median band ex-
tending to anterior margin, shorter lateral dark
brown bands, and posteriorly dark brown.
Proepisternum and dorsal part of anepisternum
yellowish-brown; rest of pleuron dark brown.
Coxae, forefemur and apex of hind femur
dark brown; rest of legs yellowish-brown. Fore-
tibia with 2 irregular rows of enlarged dorsal
setae, separated by narrow bare area; bare
area less developed than in other species. Fore-
tarsomeres enlarged, flattened. Venter of foretar-
somere 1 with several rows of enlarged setae
on anterior margin; posterior margin with row of
longer setae. Venter of foretarsomeres 2-3 with
many small, spinelike setae on posterior margin;
apicoventral seta on anterior side elongate.
Venter of foretarsomere 4 without small, spine-
like setae, but with enlarged apical seta on
posterior side and enlarged seta at midlength
on anterior side. Dorsum of foretarsomere 5
with thick, apical seta. Posterior claw of foreleg
greatly enlarged, mean 2.70 times length of
anterior claw, range 2.5-2.75. Mean costal
length 0.51 wing length, range 0.49-0.54.
Halter yellow. Abdominal tergites mostly black,
with small anterior yellow markings. Venter of
abdomen gray. Trident of intersegment 7-8
slightly flattened anterodorsally; in dorsal view
gradually broadening apically to rounded, broad
apex; lateral arms medium-sized, rounded apical-

ly-

Male. Frons yellow, except ocellar triangle
black. Flagellomere 1 orange, with dark brown
band encircling basal one-third. Palpus white;
palpal setae black, well developed. Postocular and
genal setae black. Scutum dark brown, except
light brown at anterolateral corners. Pleuron dark
brown, except proepisternum and anterior and
dorsal portions of anepisternum light brown. Legs
light brown, except apex of hind femur dark
brown. Foretarsomere 1 twice as long as broad;
other foretarsomeres about as long as broad;
pulvilli enlarged, light brown. Abdominal tergites
black, anteriorly with silvery pollinosity. Venter
of abdomen gray. Basal one-third of cercus
brown, apical two-thirds yellow. Surstylus with
short apical setae.

HOST. All specimens were collected at en-
trances to the nests of Frieseomellita varia
Lepeletier and F. silvestri Friese. The bees were
nesting in the hollow metal tubing used to
construct a canopy tower, above the level of the
tallest trees.

GEOGRAPHICAL DISTRIBUTION. Eastern
Brazil.

18 ■ Contributions in Science, Number 507

DERIVATION OF SPECIFIC EPITHET. Latin
for “chestnut-colored,” referring to the brown
pleural color.

HOLOTYPE. $, BRAZIL: Para: Caxiuaha
Station, 1.73°S, 51.45°W, 5.X.2001, B. Brown,
A. Rodrigues, bee nests on canopy tower 60 m
aboveground ILACM ENT 075990] (MPEG).

PARATYPES. BRAZIL: Para: Caxiuaha Sta-
tion, 1.73°S, 51.45°W, 2(5, 3?, 5.X.2001, 5(5,
7$, 6.X.2001, 2(5, 3?, 7.X.2001, B. Brown, A.
Rodrigues, bee nests on canopy tower 60 m
aboveground (LACM, MPEG).

Melaloncha laticlava n. sp.

(Figs. 32, 37)

RECOGNITION. This species can be recog-
nized by the dorsomedial process of the trident,
which is an extremely broad, bulbous knob. It is
most similar to M. trita, whose knob is much
narrower in dorsal view, and in which the
posterior, elongate process is much shorter than
in M. latibrachia (compare Figs. 37 and 39).

DESCRIPTION. Female. Body length 2.2 mm.
Frons yellow, except anterior margin of ocellar
triangle black; with fine reticulate sculpturing.
Frons 0.45 head width. Flagellomere 1 yellow,
white at base. Palpus white, with short, yellow
setae. Dorsal postocular setae black; genal and
other postocular setae yellow. Scutum yellow,
with posterolateral dark spot. Pleuron yellow.
Legs brownish-yellow. Foretibia with 2 rows of
enlarged dorsal setae, separated by a large bare
area. Foretarsomeres enlarged, flattened. Venter
of foretarsomere 1 with several rows of en-
larged setae on anterior margin; posterior margin
with row of longer setae. Venter of foretar-
someres 2-3 with many small, spinelike setae on
posterior margin; apicoventral seta on anterior
side elongate. Venter of foretarsomere 4 without
small, spinelike setae, but with enlarged apical
seta on posterior side and enlarged seta at
midlength on anterior side. Dorsum of foretar-
somere 5 with thick, apical seta. Posterior claw of
foreleg greatly enlarged, 2.25 times length of
anterior claw. Costa 0.49 wing length. Halter
yellow. Abdominal tergites yellow, with post-
erior black band. Venter of abdomen yellow.
Ovipositor basally yellow, apically black. Trident
of intersegment 7-8 with broad, bulbous dor-
somedial process, lateral arms well developed,
broad.

Male. Unknown.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Amazo-
nian Ecuador.

DERIVATION OF SPECIFIC EPITHET. Latin
for “broad club,” referring to the swelling of the
dorsomedial process of intersegment 7-8.

HOLOTYPE. ?, ECUADOR: Sucumbios: Sa-
cha Lodge, 0.5°S, 76.5°W, 4-14.iii.1994, P.

Brown and Kung: Revision of M. ungulata-gr oup

Hibbs, Malaise trap, 270 m ILACM ENT
036619] (QCAZ).

Melaloncha strigosa n. sp.

(Figs. 33, 38)

RECOGNITION. The trident of this species is
distinctive in dorsal view, with its broadly pointed
dorsomedial process and long, curved lateral
arms.

DESCRIPTION. Female. Body length 2.4-
3.0 mm. Frons yellow, except anterior margin of
ocellar triangle black; with finely reticulate
sculpturing. Mean frontal width 0.40 head width.
Flagellomere 1 yellow, white at base. Palpus
white, with short, yellow setae. Dorsal postocular
setae black; genal and other postocular setae
yellow. Scutum yellow, posteriorly with dark
brown posterolateral spot. Pleuron yellow. Legs
brownish-yellow; apex of hind femur slightly
darker. Foretibia with 2 rows of enlarged dorsal
setae, separated by a large bare area. Foretar-
someres enlarged, flattened. Venter of foretar-
somere 1 with several rows of enlarged setae on
anterior margin; posterior margin with row of
longer setae. Venter of foretarsomeres 2-3 with
many small, spinelike setae on posterior margin;
apicoventral seta on anterior side elongate. Venter
of foretarsomere 4 without small, spinelike setae,
but with enlarged apical seta on posterior side and
enlarged seta at midlength on anterior side.
Dorsum of foretarsomere 5 with thick, apical
seta. Posterior claw of foreleg greatly enlarged,
mean 2.87 times length of anterior claw, range
2.63-3.11. Mean costal length 0.50 wing length,
range 0.49-0.51. Halter yellow. Abdominal ter-
gites yellow, with posterior black band. Venter of
abdomen yellow. Ovipositor basally yellow,
apically black. Trident of intersegment 7-8 with
dorsomedial process broad, apically pointed,
lateral arms well developed, elongate, strongly
divergent in dorsal view.

Male. Unknown.

HOST. The flies were attracted to an aggrega-
tion of bees, the commonest of which were
Partamona epiphytophila Pedro and Camargo
and Plebeia sp. No oviposition attempts were
seen.

GEOGRAPHICAL DISTRIBUTION. Bolivia.

DERIVATION OF SPECIFIC EPITHET. Latin
adjective for “thin,” referring to the shape of the
lateral arms of the trident.

HOLOTYPE. ?, BOLIVIA: La Paz: 40 km
N Caranavi, Cumbre Alto Beni, 15.83°S,
67.56°W, 14.iv.2001, B. Brown, G. Kung, hon-
ey-sprayed leaves, 1600 m [LACM ENT 128400]
(CBFC).

PARATYPE. 1 $ , same data as holotype
(LACM), 1?, 16.iv.2003, 1?, 18.iv.2003, B.
Brown, S. Marcotte, E. Zumbado (LACM), 7$,
14.iv.2004, 2?, 16.iv.2004, 9?, 17.iv.2004, 2?,

Contributions in Science, Number 507

19. iv. 2004, 1?, 21. iv. 2004, E. Zumbado, honey
spray (CBFC, LACM).

Melaloncha trita n. sp.

(Figs. 34, 39)

RECOGNITION. This is the commonest and
most widely distributed M. ungulata-stnts spe-
cies. It is distinctive for the bulbous, almost
spherical (except for the short apical process,
which forms a posterior point) dorsomedial pro-
cess of the trident, which is smaller than that of
M. laticlava.

DESCRIPTION. Female. Body length 1.7-
2.1 mm. Frons yellow, except anterior margin of
ocellar triangle black; with finely reticulate
sculpturing. Mean frontal width 0.43 head width.
Flagellomere 1 yellow, white at base. Palpus
white, with short, yellow setae. Dorsal postocular
setae black; genal and other postocular setae
yellow. Scutum yellow, posteriorly with dark
brown, ranging from posterolateral spot to almost
entire posterior one-third. Pleuron yellow. Fegs
brownish-yellow; apex of hind femur slightly
darker. Foretibia with 2 rows of enlarged dorsal
setae, separated by a large bare area. Foretar-
someres enlarged, flattened. Venter of foretar-
somere 1 with several rows of enlarged setae on
anterior margin; posterior margin with row of
longer setae. Venter of foretarsomeres 2-3 with
many small, spinelike setae on posterior margin;
apicoventral seta on anterior side elongate. Venter
of foretarsomere 4 without small, spinelike setae,
but with enlarged apical seta on posterior side and
enlarged seta at midlength on anterior side.
Dorsum of foretarsomere 5 with thick, apical
seta. Posterior claw of foreleg greatly enlarged,
mean 2.23 times length of anterior claw, range
1.88-2.50. Mean costal length 0.50 wing length,
range 0.47-0.53. Halter yellow. Abdominal ter-
gites yellow, with posterior black band. Venter of
abdomen yellow. Ovipositor basally yellow,
apically black. Trident of intersegment 7-8 with
rounded dorsomedial process, with small apical
point, lateral arms well developed.

Male. Unknown.

HOST. Females were observed attacking Ple-
beia jatiformis Cockerell (sensu Roubik) at Zona
Protectora El Rodeo. The flies attacked the bees
head-on, apparently ovipositing in the head,
similar to the behavior described for M. borg-
meieri.

GEOGRAPHICAL DISTRIBUTION. Wide-
spread, from Mexico to Bolivia.

DERIVATION OF SPECIFIC EPITHET. Latin
adjective tritus for “commonplace,” referring to
the widespread and common nature of this
species.

HOLOTYPE. ?, COSTA RICA: San Jose:
Zona Protectora El Rodeo, 9.91°N, 84.28°W,
7.viii.2001, B. Brown, G. Kung, honey-sprayed
leaves [LACM ENT 095408] (LACM).

Brown and Kung: Revision of M. ungulata-gr oup ■ 19

PARATYPES. BOLIVIA: La Paz: 40 km N
Caranavi, Cumbre Alto Beni, 15.83°S, 67.56°W,
5$, 14.iv.2004, 3?, 16.iv.2004, 1$, 17.iv.2004,
3$, 19. iv. 2004, 1?, 21. iv. 2004, E. Zumbado,
honey spray, 1600 m (CBFC, LACM); Coroico,
Hotel Don Quixote, 16.19°S, 67.72°W, 1?,
6.iv.2001, B. Brown, G. Kung, on Phoenix palm
flowers, 1750 m (LACM). BRAZIL: Amazonas:
Manaus, Reserva Ducke, 3.13°S, 60.02°W, 3&,
8-15.iv.l992, J. Vidal, 1-4-lOm, 1$, 6-

17.vii.l992, J. Vidal, Arm. Cola, l-B-20m (INPA,
LACM). COSTA RICA: Alajuela: 20 km S Upala,
1$, 10-21.V.1991, F. Parker (EMUS); Cartago:
Turrialba, 9.93°N, 83.67°W, 2?, 15-19.vii.1965,
P. Spangler (USNM). Guanacaste: Rio Naranjo,
Fmca Montezuma, 10.67°N, 85.06°W, 1$,

22.ii.2002, 1?, 23.11.2002, 1?, 26.ii.2002,

Brown, Walker, Gonzalez, Zumbado, honey-
sprayed undergrowth/bee screen (LACM,
MUCR); Heredia: La Selva Biological Station,
10.43°N, 84.02°W, 1?, 4.iv.l994, ALAS, Ma-
laise trap M/03/385, 1$, 24.X.1994, ALAS,
fogging FVK/30/16, 1?, 6.iii.2000, ALAS, Ma-
laise trap M/19/743 (INBC). San Jose: Zona
Protectora El Rodeo, 9.91°N, 84.28°W, 1?,
l.viii.2001, B. Brown, bee screen, 1 ? , 2.viii.2001,
1?, 5.viii.2001, 1?, 8.viii.2001, B. Brown, G.
Kung, honey-sprayed leaves, 1?, 1-5. viii. 2001,
1?, 3-8.viii.2001, B. Brown, V. Berezovskiy, G.
Kung, Malaise trap, 1000 m (LACM). ECUA-
DOR: Sucumbios: Sacha Lodge, 0.5°S, 76.5°W,
1?, 22.ii-4.iii.1994, P. Hibbs, Malaise trap,
270 m (LACM). MEXICO: Chiapas: Palenque,
3?, 10.ix.l974, G. Bohart, W. Hanson (EMUS,
LACM); Oaxaca: 35 mi W Tehuantepec, 1?,
16.ix.l974, G. Bohart, W. Hanson (EMUS).
PANAMA: Canal Zone: Barro Colorado

Island, 9.17°N, 79.83°W, 1?, 18-25.xi.l992,
J. Pickering, Malaise trap #727, 1?, 7-

14. iv. 1993, J. Pickering, Malaise trap #961,
2$, 11-18. V. 1994, J. Pickering, Malaise trap
#2424 (LACM, MIUP). PERU: Madre de
Dios, Tambopata Research Center, 13.14°S,
69.6HW, 1?, 20.vii.2001, 1?, 22.vii.2001, B.
Brown, G. Kung, honey-sprayed leaves (LACM,
MUSM); San Martin: San Antonio de Cumbaza,
6.40°S, 67.4HW, 1$, 5.vii.2004, B. Brown, E.
Corona, honey spray, 400 m (LACM). VENE-
ZUELA: Aragua: 2 km N Ocumare de la Costa,
1$, 21-22.vi.l976, A. S. Menke, D. Vincent
(USNM).

Melaloncha ungulata Borgmeier
(Figs. 35, 40)

Melaloncha ungulata Borgmeier, 1938, p. 45,

figs. 5-6, 12-13.

Melaloncha genitalis Borgmeier, 1934, p. 181,

plate 3 fig. 15; 1971, pp. 130-131 (in part).

LECTOTYPE. Because of past confusion of
this species with the name M. genitalis, we here

20 ■ Contributions in Science, Number 507

designate a lectotype to fix the concept of M.
ungulata: ?, BRAZIL: Santa Catarina: Nova
Teutonia, 27.18°N, 52.38°W, vii.1937, F. Plau-
mann [LACM ENT 122572] (MZSP).

RECOGNITION. Eemales are recognized by
the shape of the dorsomedial process of the
trident, which is squared in lateral view but
arrowhead-shaped in dorsal view.

Males were associated with the females by
Borgmeier, They are of similar size as the female
specimens, and some were collected at the same
time. We have not included them as paralecto-
types, as no definitive evidence {in copula pairs)
has yet been found.

DESCRIPTION. Female (note that colors in
this description are based a specimen collected
in 1967, not the lectotype; in the lectotype all
colors that are here described as yellow are
dark yellow to brown, apparently because of
their method of preservation). Body length 2.4-
2.6 mm. Frons yellow, except anterior margin
of ocellar triangle black; with finely reticulate
sculpturing. Mean frontal width 0.42 head
width. Flagellomere 1 yellow. Palpus white, with
short, yellow setae. Dorsal postocular setae
black; genal and other postocular setae yellow.
Scutum yellow, with small posterolateral dark
spot. Pleuron yellow. Legs brownish-yellow;
apex of hind femur not darkened. Foretibia
with 2 rows of enlarged dorsal setae, separated
by a large bare area; bare area in some speci-
mens with few scattered seta. Foretarsomeres
enlarged, flattened. Venter of foretarsomere 1
with several rows of enlarged setae on anterior
one-half; posterior margin with row of longer
setae. Venter of foretarsomeres 2-3 with many
small, spinelike setae on posterior margin; tar-
somere 2 with similar setae near apicoventral seta;
both tarsomeres with apicoventral seta on ante-
rior side elongate; tarsomere 3 with medium-
sized, medial seta near apex. Venter of foretar-
somere 4 without small, spinelike setae, but with
enlarged apical seta on posterior side and enlarged
seta at midlength on anterior side. Dorsum of
foretarsomere 5 with thick, apical seta. Posterior
claw of foreleg greatly enlarged, mean 2.43 times
length of anterior claw, range 2.36-2.50. Mean
costal length 0.48 wing length (all specimens).
Halter yellow. Abdominal tergites yellow, with
posterior black band. Venter of abdomen yellow.
Ovipositor basally yellow, apically black. Trident
of intersegment 7-8 with small, anteriorly flat-
tened dorsomedial process that is narrow in
dorsal view, lateral arms medium-sized, relatively
broad.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. South-
eastern Brazil.

OTHER MATERIAL EXAMINED. Paralecto-
types: BRAZIL: Santa Catarina: Nova Teutonia,
27.18°N, 52.38°W, vi.l937, 2$, vii.1937, 1?,
194.?, 1?, vi.l941, 1$, ii.l967 (LACM, MZSP).

Brown and Kung: Revision of M. ungulata-gr oup

Possible males: 2(5, vi.l937, 3(5, vii.1937
(LACM, MZSP).

Melaloncha atrilingula-inic2LScnts

DIAGNOSIS. Dorsomedial process of trident
flattened, with dark-colored lateral margins.

INCLUDED SPECIES. Melaloncha atrilingula,
M. inicua, and M. pegmata.

Melaloncha atrilingula n. sp.

(Fig. 41)

RECOGNITION. This species can be recog-
nized by the narrow, darkly sclerotized dorsome-
dial process, best seen in dorsal view.

DESCRIPTION. Female. Body length 2.3-
2.4 mm. Frons yellow, except anterior margin of
ocellar triangle black; with finely reticulate
sculpturing. Mean frontal width 0.43 head width
(both specimens). Flagellomere 1 yellow, white at
base. Palpus white, with short, yellow setae.
Dorsal postocular setae black; genal and other
postocular setae yellow. Scutum yellow, poster-
iorly with dark brown posterolateral spot.
Pleuron yellow. Legs brownish-yellow; apex of
hind femur slightly darker. Foretibia with 2 rows
of enlarged dorsal setae, separated by a large
bare area. Foretarsomeres enlarged, flattened.
Venter of foretarsomere 1 with several rows of
enlarged setae on anterior margin; posterior
margin with row of longer setae. Venter of
foretarsomeres 2-3 with many small, spinelike
setae on posterior margin; apicoventral seta on
anterior side elongate. Venter of foretarsomere 4
without small, spinelike setae, but with enlarged
apical seta on posterior side and enlarged seta at
midlength on anterior side. Dorsum of foretar-
somere 5 with thick, apical seta. Posterior claw
of foreleg greatly enlarged, mean 2.53 times
length of anterior claw, range 2.40-2.67. Mean
costal length 0.51 wing length, range 0.50-0.51.
Halter yellow. Abdominal tergites yellow, with
posterior black band. Venter of abdomen yellow.
Ovipositor basally yellow, apically black;
relatively straight in lateral view, not curved
dorsally as in other species. Trident of interseg-
ment 7-8 with narrow, black-margined dorsome-
dial process apically rounded in dorsal view;
lateral arms well developed, strongly divergent in
dorsal view.

Male. Unknown.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Amazo-
nian Brazil.

DERIVATION OF SPECIFIC EPITHET. Lat-
in, derived from words for “black” and “small
tongue,” referring to the dorsomedial process of
intersegment 7-8.

HOLOTYPE. ?, BRAZIL: Amazonas: Man-
aus, Reserva Ducke, 3.13°S, 60.02°W, 6-

17.vii.l992, J. Vidal, Arm. Cola, lB-20 [LACM
ENT 008393] (INPA).

Contributions in Science, Number 507

PARATYPE. 1$, same data as holotype
(LACM).

Melaloncha inicua n. sp.

(Fig. 42)

RECOGNITION. This species is similar to M.
pegmata but is smaller, the comb of setae on
tarsomere 4 is elongated, and the dorsomedial
process of the trident is slightly narrower,

DESCRIPTION. Eemale. Body length 1.8 mm.
Frons yellow, except anterior margin of ocellar
triangle black; with finely reticulate sculpturing.
Frons 0.45 head width. Flagellomere 1 yellow,
white at base. Palpus white, with short, yellow
setae. Dorsal postocular setae black; genal and
other postocular setae yellow. Scutum yellow,
posteriorly with dark brown posterolateral spot.
Pleuron yellow. Legs brownish-yellow; apex of
hind femur slightly darker. Foretibia with 2 rows
of enlarged dorsal setae, separated by a large bare
area. Foretarsomeres enlarged, flattened. Venter
of foretarsomere 1 with several rows of enlarged
setae on anterior margin; posterior margin with
row of longer setae. Venter of foretarsomeres 2-3
with many small, spinelike setae on posterior
margin; apicoventral seta on anterior side elon-
gate. Venter of foretarsomere 4 without small,
spinelike setae, but with enlarged apical seta on
posterior side and enlarged seta at midlength on
anterior side; setae of palisade on posterior
margin extremely long, about 4 times length of
those of tarsomere 3, and totaling about one-half
width of tarsomere. Dorsum of foretarsomere 5
with thick, apical seta. Posterior claw of foreleg
greatly enlarged, 3.25 times length of anterior
claw. Costa 0,52 wing length. Halter yellow.
Abdominal tergites yellow, with posterior black
band. Venter of abdomen yellow. Ovipositor
basally yellow, apically black; relatively straight
in lateral view, not curved dorsally as in other
species. Trident of intersegment 7-8 with slightly
narrowed, black-margined dorsomedial process
apically truncate in dorsal view; lateral arms well
developed, strongly divergent in dorsal view.

Male. Unknown.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Bolivia.
DERIVATION OF SPECIFIC EPITHET. The
name is based on the Rio Inicua near the type
locality and is treated as a noun in apposition.

HOLOTYPE. $, BOLIVIA: La Paz: Alto Rio
Beni, south of Rio Inicua, 1100m, 15-1 8. i. 1976,
L. Pena [LACM ENT 0956731 (AMNH).

Melaloncha pegmata n. sp.

(Fig. 43)

RECOGNITION. This species is similar to M.
inicua but differs by the extremely broad apex of
the dorsomedial process of the trident, the

Brown and Kung: Revision of M. ungulata-group ■ 21

unmodified palisade of setae on foretarsomere 4,
and the larger body size.

DESCRIPTION. Female. Body length 2.5 mm.
Frons yellow, except anterior margin of ocellar
triangle black; sculpturing finely reticulate with
few lateral punctures. Frons 0.38 head width.
Flagellomere 1 yellow. Palpus white, with short,
yellow setae. Dorsal postocular setae black; genal
and other postocular setae yellow. Scutum yellow
with small posterolateral dark brown macula.
Pleuron yellow. Legs brownish-yellow. Foretibia
with 2 rows of enlarged dorsal setae, separated
by a large bare area. Foretarsomeres enlarged,
flattened. Venter of foretarsomere 1 with several
rows of enlarged setae, plus enlarged apical seta
on anterior margin; posterior margin with row
of longer setae. Venter of foretarsomeres 2-3
with extremely dense areas of many spinelike
setae on posterior margin; apicoventral seta
on anterior side enlarged. Venter of foretarsomere
4 without small, spinelike setae, but with enlarged
apical seta on posterior side and enlarged seta
at midlength on anterior side; setae of palisade
on posterior margin only slightly longer than
those of tarsomere 3. Dorsum of foretarsomere 5
with thick, apical seta. Posterior claw of foreleg
greatly enlarged, 2.43 times length of anterior
claw. Costa 0.56 wing length. Halter yellow.
Abdominal tergites yellow, with posterior black
band. Venter of abdomen yellow. Ovipositor
basally yellow, apically black. Trident of in-
tersegment 7-8 with large, rounded, dorsally
flattened dorsomedial process, with broad, blunt
apex with thick, dark margin, lateral arms
elongate.

Male. Unknown.

HOST. Unknown.

GEOGRAPHICAL DISTRIBUTION. Costa
Rica.

DERIVATION OF SPECIFIC EPITHET.

Greek pegma for “thickened,” for the heavi-
ly sclerotized, shelflike process of intersegment
7-8.

HOLOTYPE. 9, COSTA RICA: Guanacaste:
3 km SE Rio Naranjo, 21-31. vii. 1992, F. D.
Parker [LACM ENT 1141311 (LACM).

KEY TO SPECIES (FEMALES ONLY)

1 Claws of foreleg extraordinarily asymmetrical,
with outer (posterior) claw about 2.25-3.25
times length of inner (anterior) claw; mostly
yellow species, but 1 brown species (M.
castanea) with well-developed dorsoapical seta

on foretarsomere 5 (Fig. 56)

M. ungulata-sents, (see key below)

- Claws of foreleg subequal or at most only
slightly asymmetrical, with outer claw less
than 1.5 times length of inner claw, but if
outer claw longer (2 times length of inner claw
in M. setitibialis and M. cristula) then brown

22 ■ Contributions in Science, Number 507

species lacking differentiated dorsoapical seta
on foretarsomere 5 2

2 Frons yellow to orange; central region of frons

in 1 species (M. adusta) sometimes brown, but
in this exceptional species, ventral margin and
lateral margins ventral to ventral fronto-
orbital setae orange and dorsum of ovipositor
bare 3

- Frons brown to black throughout, except

for some lighter color at ventral extremity
in some species and orange setal bases in
some specimens of M. nigrifrons-, lateral
margin of frons ventral to ventral fronto-
orbital setae dark colored; dorsum of ovipo-
sitor setose 6

3 Abdominal tergites completely yellow, with-
out substantial black markings

M. flava Borgmeier

- Abdominal tergites yellow with large black

markings to completely dark 4

4 Lateral arms of trident extremely short;

tarsomeres of foreleg extremely broad, flat-
tened; ovipositor dorsally with small setulae;
predominantly yellow-colored species; coxal
setae yellow M. platypoda n. sp.

- Lateral arms of trident as long as dorsomedial

process; tarsomeres of foreleg not strongly
flattened; ovipositor dorsally bare; predomi-
nantly yellow to mostly dark brown species;
coxal setae black 5

5 Abdominal tergites yellow and black; dor-

somedial process of trident as long as
lateral arms; lateral arms somewhat broad-
ened at apex; yellower species

M. acoma n. sp.

- Abdominal tergites almost completely black to

blackish-brown, with small areas of dark
orange in some specimens; dorsomedial pro-
cess of trident shorter than lateral arms; lateral
arms thin M. adusta n. sp.

6 Dorsomedial process elaborated at tip: ex-

panded, laterally flattened, deepened (Figs. 15-
23) 7

- Dorsomedial process simple, apically pointed

(Figs. 7-9, 11-14) 15

7 Elaboration of apex of dorsomedial process of

trident in the form of relatively small, rounded
apical lobe (Figs. 15-17) 8

- Elaboration of apex of dorsomedial process of

trident in the form of deep, flattened process
(Eigs. 18-23) 10

8 Row of posteroventral seta on foretibia

extremely long, about twice depth of ti-
bia M. setitibialis n. sp.

- Row of posteroventral setae on foretibia

relatively short, about equal to depth of
tibia 9

9 Lateral arm of trident elongate, narrow,
longer than dorsomedial process; dorsomedial

process straight (Eig. 15)

M. claviapex n. sp.

Brown and Kung: Revision of M. ungulata-^rowp

- Lateral arm of trident relatively short, thick; dor-
somedial process extended dorsally (Eig. 16) ...
M. debilis n. sp.

10 Dorsomedial process of trident with narrow

stalk proximal to flattened apex (Eig. 19-
23) 11

- Dorsomedial process of trident without nar-
row stalk (Eig. 18) M. caligula n. sp.

11 Dorsomedial process of trident relatively

narrow and strongly concave posteriorly
(Eig. 19) M. cristula n. sp.

- Dorsomedial process of trident broader and
less concave posteriorly (Eigs. 20-23) ... 12

12 Dorsomedial process of trident with narrow

dorsal point and small posterodorsal concavity
(Eig. 21); circular excision below stalk rela-
tively small M. spina n. sp.

- Dorsomedial process of trident more truncate

dorsally, or without posterior concavity; cir-
cular excision various 13

13 Dorsomedial process of trident dorsally form-

ing broad point, posteriorly evenly convex
(Eig. 23) M. tuparroensis n. sp.

- Dorsomedial process of trident dorsally form-

ing truncate or rounded apex; posterior mar-
gin not evenly curved (Eigs. 20, 22) 14

14 Dorsomedial process of trident relatively nar-
row; consists of strongly sclerotized, dorsal
support and small, ventral, lighter brown
flattened area; dorsal apex slightly narrowed,
knoblike; posteroventral portion of flattened
area without concavity; circular excision

below stalk relatively small (Eig. 20)

M. culmena n. sp.

- Dorsomedial process of trident relatively
broad; consists mostly of flattened, light
brown cuticle; dorsal apex flat, broader;
posteroventrally with small concavity; circular

excision larger (Eig. 22)

M. tambopatensis n. sp.

15 Pleuron partly to completely yellow or light

brown; coxae yellow to white; tarsomere 1 of
foreleg elongate 16

- Pleuron completely dark brown; coxae brown,

at least 1 coxa dark brown; tarsomere 1 of
foreleg short, broad 18

16 Abdominal tergites completely black; dor-

somedial process of trident as long as or
longer than lateral arms (Pigs. 7-8) 17

- Abdominal tergites black and yellow banded;

dorsomedial process of trident shorter than late-
ral arms (Pig. 9) M. nigrifrons Borgmeier

17 Dorsomedial process of trident longer than

lateral arms (Pig. 7) M. Candida n. sp.

- Dorsomedial process of trident equal in length

to lateral arms (Pig. 8)

M. nigrita Borgmeier

18 Palpus white to yellow 19

- Palpus dark brown . . . M. fuscipalpis n. sp.

19 Legs, including coxae, light brown to dark

brown 20

Contributions in Science, Number 507

- Legs, including coxae, whitish-yellow

M. nigrita Borgmeier

20 Basal one-eighth to one-half of forefemur dark
brown, apical one-half lighter in color; frons
dark brown, almost black; lateral arms of

trident longer than dorsomedial process

21

- Forefemur uniformly yellowish-brown; frons
medium brown in color; lateral arms of trident
subequal in length to dorsomedial process . . .
M. ustulata n. sp

21 Dorsomedial process slightly upturned at tip

(Fig. 11); apical one-half of forefemur brown-
ish-yellow; lateral arms of trident widely
separated below tip of dorsomedial process
(Fig. 25) M. diastata n. sp.

- Dorsomedial process relatively straight, not
upturned apically (Fig. 13); apical one-half of
forefemur more yellow in color; lateral arms
of trident close together (touching) below tip
of dorsomedial process (Fig. 27), although this

possibly is an artifact of drying

M. juxta n. sp.

KEY TO MELALONCHA UNGULATA-
SERIES EEMALES

1 Apex of foretarsomere 5 lacking a strongly

differentiated dorsal seta 2

- Apex of foretarsomere 5 with large, black,

strongly differentiated seta dorsally (Fig. 56) .
3

2 Apex of intersegment 7-8 not strongly down-
turned, with small dorsal knob (Fig. 29);
lateral arms larger . ... M, borgmeieri n. sp.

- Apex of intersegment 7-8 strongly down-
turned; lateral arms barely differentiated,

not noticeable (Fig. 30)

M. curtibrachia n. sp.

3 Pleural sclerites light brown to dark brown . .
M. castanea n. sp.

- Pleural sclerites yellow 4

4 Dorsomedial process in dorsal view with

darkened margins; dorsum of process flat . .
5

- Dorsomedial process rounded, bulging dorsal-
ly, without darkened margin 7

5 Dorsomedial process narrow, apically round-
ed in dorsal view (Fig. 41)

M. atrilingula n. sp.

” Dorsomedial process apically truncate,
broader (Figs. 42-43) 6

6 Small species, body length 1.8 mm; setae of

palisade on posterior margin of foretarsomere
4 greatly elongate, about 4 times length of
those on segment 3 M. inicua n. sp.

- Larger species, body length 2.5 mm; setae of

palisade on posterior margin of foretarsomere
4 only slightly longer than those of segment
3 M. pegmata n. sp.

7 Dorsomedial process extremely broad, apical-
ly pointed; lateral arms elongate, narrow.

Brown and Kung: Revision of M. ungulata-^roup ■ 23

apically divergent (Fig. 38)

M. strigosa n. sp.

- Dorsomedial process relatively narrow in
dorsal view; if broad, then not apically
pointed; lateral arms less elongate, thicker .
8

8 Dorsomedial process broad in dorsal view

(Fig. 37) M. laticlava n. sp.

- Dorsomedial process narrower in dorsal view

(Figs. 39-40) 9

9 Small species, body length 1. 7-2.1 mm; dor-

somedial process rounded, bulging dorsally
(Fig. 34) M. trita n. sp.

- Larger species, body length 2. 4-2. 6 mm;
dorsomedial process less bulbous, more flat-

topped (Fig. 35)

M. ungulata Borgmeier

ACKNOWLEDGMENTS

For technical assistance we thank V. Berezovskiy.
Illustrations were skillfully prepared by L. Mui. We
thank V. Berezovskiy, L. Gonzalez, S. Marcotte, K.
Walker, and E. Zumbado for field assistance. For
continuing help in conducting work in Costa Rica, we
thank P. Hanson and M. Zumbado. For help
obtaining permission to do research in Colombia,
we thank F. Fernandez and M. Sharkey; for help
obtaining permission in Bolivia, we thank J. Sar-
miento and S. Gaimari. Research in Peru was
facilitated by D. Feener and the friendly staff of the
Tambopata Research Center. Our work in Argentina
was aided by L. Elizalde, P. Folgarait, K. Schiaffino,
D. Rode, M. DiBitetti, and A. Bachmann. For
assistance in Brazil, we are grateful to R. Evans, M.
Martins, and A. Rodrigues. For identification of host
bees, we thank D. Roubik, C. Rasmussen, and J.
Camargo.

Research on Melaloncha was funded by National
Science Foundation grants DEB-0090031 to B. Brown
and DEB-0315271 to B. Brown and P. Smith as well as
a Research Experience for Undergraduates supplement
to support F. Gonzalez. Work in Colombia was
facilitated by NSF grants DEB-9972024 and 0205982
to M. Sharkey and B. Brown. NSF grants BSR-9025024,
DEB-9401069, DEB-9706976, and DEB-0072702
funded the AFAS project. J. Pickering’s work on Barro
Colorado Island was funded by the Smithsonian
Environmental Sciences Program. Work in Costa Rica
was supported in part by support to INBio by the
Biodiversity Resources Development Project (CR-GE-
39876), funded by Global Environmental Facilities
through its implementing agency the World Bank. NSF
grant DBI-02166 allowed purchase of a SEM at the
LACM.

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world. Flora and Fauna Handbook No. 11. Gai-
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Borgmeier, T. 1934. O genero Melaloncha Brues (Dipt.
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. 1938. Alguns Phorideos myrmecophilos de

Costa Rica e do Brasil e quatro especies novas de

24 ■ Contributions in Science, Number 507

Melaloncha Brues (Dipt. Phoridae). Revista de
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. 1959. Neue und wenig bekannte Phoriden aus

der neotropischen Region, nebst einigen Arten aus
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. 1968. A catalogue of the Phoridae of the world

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the Neotropical Region (Diptera, Phoridae). Studia
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. 1971b. Supplement to a catalogue of the

Phoridae of the world (Diptera, Phoridae). Studia
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Brown, B.V. 2001. Beginning to study the bee-killing
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. 2004a. Revision of the Melaloncha cingulata-

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Brown and Rung: Revision of M. ungulata-^roup

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. 2004b. Revision of the subgenus Udamochiras

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. 2005. Revision of the Melaloncha furcata-group

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Systematics and Evolution 36:241-258.

Gonzalez, L., and B.V. Brown. 2004. New species
and records of Melaloncha {Udamochiras) bee-
killing flies (Diptera: Phoridae). Zootaxa 730:
1-14.

McAlpine, J.F. 1981. Morphology and terminology —
adults. In Manual of Nearctic Diptera, vol. 1, J.F.
McAlpine, B.V. Peterson, G.E. Shewell, H.J.
Teskey, J.R. Vockeroth, and D.M. Wood, 9-63.
Ottawa: Agriculture Canada.

Received 1 January 2005; accepted 25 April 2005.

Contributions in Science, Number 507

Brown and Kung: Revision of M. ungulata-gr oup ■ 25

2

M. flava
M. acoma
M. adusta
M. Candida
M. nigrita
M. nigrifrons
M. platypoda
M. fuscipalpis
M. diastata
M. juxta
M. ustulata
M. Caligula - series
M. ungulata - series

M. nigrita
-subgroup

M. ungulata
-subgroup

Figure 2 Cladogram of hypothesized relationships of M. imgulata-group species. Numbers refer to characters
discussed in Phylogenetic Analysis

M. borgmeieri
M. curtibrachia
M. castanea
M. laticlava
M. strigosa
M. trita
M. ungulata

M. atrilingula
M. inicua
M. pegmata

Figure 3 Cladogram of hypothesized relationships of M. ungulata-series species. Numbers refer to characters
discussed in Phylogenetic Analysis

26 ■ Contributions in Science, Number 507

Brown and Kung: Revision of M, ungulata-^roxnp

11

M. platypoda

M. diastata

dorsomedial

process

lateral

M. flava

5 M. acoma

10

Figures 4-14 Ovipositors, left lateral. Note that Figure 5 is a fully extended ovipositor, whereas others are withdrawn

Contributions in Science, Number 507

Brown and Kung: Revision of M. ungulata-§r oup ■ 27

20 M. culmena

M. platypoda

M. diastata

Figures 15-25 Ovipositors: 15-23. left lateral; 24-25. dorsal

28 ■ Contributions in Science, Number 507

Brown and Rung: Revision of M. ungulata-^ronp

26

posterior

2.Q M. borgmeieri extension

30 M. curtibrachia

31 M. castanea

M. fuscipalpis

M. juxta

M. ustulata

32 laticlava

33 strigosa

34 M.trita

35 ungulata

36 M. atrilingula

M. laticlava M. strigosa

M. trita

M. ungulata M. atrilingula

M. inicua

M. pegmata

Figures 26-43 Female terminalia; 26-28. ovipositors, dorsal; 29-36. intersegment 7-8, left lateral; 37-43.
intersegment 7-8, dorsal

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Brown and Kung: Revision of M. ungulata-grouip ■ 29

Figures 44-49 Scanning electron micrographs: 44. M. acoma n. sp., foretarsomere 1, dorsal; 45-47. M. fiiscipalpis
n. sp. 45. foretarsomere 1, ventral, 46. foretarsomeres 2-4, ventral, 47. claw of foretarsomere 5; 48-49. M. debilis
n. sp., sculpturing of frons, 48. low magnification, 49. higher magnification

30 ■ Contributions in Science, Number 507

Brown and Kung: Revision of M. ungulata-gr oup

elongate anterior
seta N

enlarged

setae

thickened curved
posterior seta

S-3000 WD 5 . 3mm 10 . OkV x400 100m

13-Sep-04

Figures 50-55 Scanning electron micrographs: 50-51. M. debilis n. sp. 50. foretarsomeres 1-3, ventral, 51.
foretarsomeres 3-5, ventral; 52-55. M. borgmeieri n. sp. 52. frons, 53. foretarsomeres 1-3, ventral, 54. foretarsomeres
3-5, ventral, 55. foretarsomeres 4-5, ventral

Contributions in Science, Number 507

Brown and Kung: Revision of M. ungulata-^tou^ ■ 31

Figures 56-59 Scanning electron micrographs of M. strigosa n. sp.: 56. foretarsomere 5, dorsal; 57. foretarsomeres
4-5, ventral; 58. foretarsomere 1, ventral; 59. fortarsomeres 2-3, ventral

Natural

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